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VOL. 90 JANUARY 1988 ‘NO. i
95, 90674 (ISSN 0013-8797)
(> PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
ot WASHINGTON °
b 05 1988 PUBLISHED
apts “QUARTERLY
BURGER, J. F.—A new genus and two new species of Pangoniini (Diptera: Tabanidae) of
zoogeographic interest from Sabah, Malaysia ................00.0 000 cece eee eee ee 12
CLEMENT, S. L. and T. MIMMOCCHI—Occurrence of selected flower head insects of
CETIOUTEER SOISIIIAUS\AN Italy ANGUGLEECE | av be\s oh) lM b ahlasicters balers eizk, Al an lets eile wlele Kabres 47
DROOZ, A. T. and H. H. NEUNZIG—Notes on the biology of two Phycitines (Lepidoptera:
Pyralidae) associated with Toumeyella pini (Homoptera: Coccidae) on Pine ............ 44
GOEDEN, R. D.—Gall formation by the capitulum-infesting fruit fly, Tephritis stigmatica (Dip-
tera a Me DHnrTGde) Meee Sh Le ett ae eL Ee Era he RT PP ROME UN BERET MAD AS ML STS Ore i 57)
HALSTEAD, J. A.—Belaspidia longicauda, new species, the first Nearctic Belaspidia (Hyme-
MOPLETEAGhalGididae) cat Wl RU asd) Suna polis he FEAL OP REPEAL der eat fet be Pe eek 87
HANSSON, C.—A revision of the genus Mestocharis and a review of the genus Grahamia
(iymenopterd, BUlOophidazey sy Bitty tee Aaete e AWE eA RETRO URIBE NS eat erp: 28
HEYDON, S. L.—A review of the Nearctic species of Cryptoprymna Forster, with the description
of a new genus, Polstonia (Hymenoptera: Pteromalidae) .........................----- 1
MAcDONALD, J. F.—New synonyms pertaining to Chelifera and generic key for North Amer-
ican Llemerodromiinae: (Diptera: |Empididae) 2.00. Waki joke ia eee AAD st 98
MATHIS, W. N. and T. ZATWARNICKI—Studies on the systematics of the shore-fly tribe
DAPini (Diptera Ephydridae) Mansy Week Ue k M ME Lite LORIE EM) Bel Le Bea TPM Niel i 106
NEAL, J. W., Jk.—Unusual oviposition behavior on evergreen azalea by the Andromeda lace
bug Stephanitis takeyai (Drake and Maa) (Heteroptera: Tingidae) .................... 52
OSWALD, J. D.—A review of the South Pacific genus Austromegalomus Esben-Petersen (Neu-
roptera: Hemerobiidae) with a description of a new species from Rapa ................. 55
PETERSON, B. V., M. VARGAS V., and J. RAMIREZ-PEREZ—Simulium (Hemicnetha)
hieroglyphicum (Diptera: Simuliidae), a new black fly species from Costa Rica .......... 76
RAMIREZ-PEREZ, J., B. V. PETERSON, and M. VARGAS V.— Mayacnephia salasi (Diptera:
Simuliidae), a new black fly species from Costa Rica ...........0...0. 000020 e cece eee 66
STARK, B. P., S. W. SZCZYTKO, and B. C. KONDRATIEFF—The Cultus decisus complex
of Eastern North America (Plecoptera: Perlodidae) .................0...00..0 22200005.
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1988
F. EUGENE Woon, President WARREN STEINER, Program Chairman
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Publications Committee
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Title of Publication: Proceedings of the Entomological Society of Washington.
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PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 1-11
A REVIEW OF THE NEARCTIC SPECIES OF CR YPTOPRYMNA
FORSTER, WITH THE DESCRIPTION OF A NEW GENUS,
POLSTONIA (HYMENOPTERA: PTEROMALIDAE)
STEVEN L. HEYDON
Illinois Natural History Survey, Section of Faunistic Surveys and Insect Identification,
607 E. Peabody Drive, Champaign, Illinois 61820.
Abstract.—The genus Cryptoprymna Forster, herein reported from the Nearctic region
for the first time, is represented by two species, the Holarctic species C. atra (Walker)
and the Nearctic species C. dixiana n. sp. Cryptoprymna is redescribed. C. atra is reported
herein to be a parasitoid of the pupal stages of syrphids on conifers. A new genus, Polstonia,
is described with two included Nearctic species: P. quadriplana n. sp., the type species,
and P. pelagocorypha n. sp. A modification of Graham’s (1969) key to the genera of the
Sphegigasterini is presented to facilitate identification of the genus Po/stonia. Keys are
given to the Nearctic species of both genera.
This is the second, following Heydon and
LaBerge (in press), in a series of papers re-
vising the Nearctic miscogasterine Pter-
omalidae. The specimens upon which this
review 1s based were among the material
submitted to me by various collections for
that first paper, which was a revision of the
genus Sphegigaster Spinola. Terminology
and methodology follow those used by Hey-
don and LaBerge (in press) except that de-
scriptions of new species are based on the
type-specimens, the “‘sensillae’’ of the fu-
nicular and club segments are called mul-
tiporous plate (abbreviated MPP) sensillae,
and the specimens were examined under
fluorescent light so may appear more green
than described herein when viewed under
incandescent light.
This paper contains the first Nearctic rec-
ord for Cryptoprymna Forster. The Nearctic
fauna of Cryptoprymna contains one Hol-
arctic species, C. atra (Walker), and the new
Nearctic species C. dixiana. Because the
original description of the genus (Walker
1833) is now insufficient for distinguishing
this genus from several other similar genera
erected since 1833, I am redescribing the
genus based on my examination of three of
the four described species. The original de-
scription of C. atra is also very short so I
am presenting a detailed diagnosis for dis-
tinguishing this species from C. dixiana.
This diagnosis 1s based on the Nearctic
specimens of C. atra, which were compared
by me with specimens from the Palearctic
region from the British Museum of Natural
History.
The new genus Polstonia is described and
followed by descriptions of the two new
Nearctic species included in the genus, P.
quadriplana and P. pelagocorypha. The geo-
graphic range of this genus extends into the
Neotropical region since I have seen spec-
imens from South America that belong to
other species in this genus. None of these
Neotropical species will be described here
due to lack of sufficient material for a thor-
ough study of the fauna of this region.
Both these genera key to the Sphegigas-
terini in Graham (1969) and are similar in
2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
having the anterior margin of the clypeus
without any projecting denticles and retic-
ulate petioles which are distinctly longer than
wide. Relationships between these genera
and the other genera of the Miscogasterinae
will be discussed more fully in a later paper.
Cryptoprymna Forster
Prosodes Walker, 1833: 371, 374 (Preoc-
cupied by Eschscholtz 1829). Type-
species: Prosodes ater Walker 1833
(monotypy). Lectotype male in the
BMNH. Brulle, 1846: 582-583. Gahan
and Fagan, 1923: 121.
Cryptoprymna Forster, 1856: 52 (key), pp.
56, 59. Walker, 1872: 97, 98 (key, syn-
onymy). Ashmead, 1904: 330, 332, 372
(key). Nikol’skaya, 1952: 252 (key).
Schmiedeknecht, 1909: 375, 376, 380
(key, diagnosis). Gahan and Fagan, 1923:
41. Peck, Boucek, and Hoffer, 1964: 40
(key). Graham, 1969: 124, 140 (key, syn-
onymy). Dzhanokmen, 1978: 77, 80 (key).
Farooqiand Subba Rao, 1985: 260, 310G.
Farooqi and Subba Rao, 1986: 285.
Cryptoprymnus: Thomson, 1878: 17, 22.
Cresson, 1887: 75 (key). [Invalid emen-
dation]
Walker (1833) erected the genus Prosodes
in his Monographia Chalciditum. Forster
(1856) pointed out that the name Prosodes
had previously been proposed for a genus
of tenebrionid beetles (Eschscholtz 1829),
and he renamed the genus Crypfoprymna.
There are presently three described species:
C. atra (Walker 1833), a Holarctic species;
C. africanus Boucek (1976), from southern
Africa; and C. brama (Motschulsky 1863),
from southern Asia. I add a fourth species,
C. dixiana, from the southeastern United
States. Ashmead (1896) described Crypto-
prymna illinoensis from the Nearctic, but
this species was transferred to Callitula Spi-
nola (Delucchi 1955) and later synony-
mized with Callitula cyrnus Walker (Burks
1975).
Description.— Color: Head, mesosoma,
coxae, and petiole black, gaster dark brown.
Wing hyaline.
Female.— Head transversely oval in an-
terior view; 2x as wide as long; clypeus
subareolate, anterior margin truncate: gena
with broad concavity extending from mouth
margin to lower orbit; eye glabrous; occiput
concave or straight posteriorly, acarinate.
Antenna inserted below middle of face, just
above lower orbits; formula 1:1:2:6:3; scape
slender, length 8 x width, reaching or nearly
reaching median ocellus; club distinctly
wider than F6, sutures oblique, with a patch
of micropilosity on terminal or terminal two
segments, terminal spine or projection ab-
sent. Mandible 4-tooth, upper two smaller
and approximated. Mesosoma compact,
rounded dorsally in profile; pronotum with
neck short, collar with sharp transverse ca-
rina anteriorly and smooth except imme-
diately behind carina; mesoscutum with no-
tauli extending to its hind margin as
impressed lines; scutellum as long as wide,
frenal sulcus nearly obliterated; prepectus
acarinate; mesopleuron with upper epime-
ron smooth; propodeum as long as scutel-
lum, strongly arched, median carina and
plicae sharp, median panels alveolate with
some rugae, spiracles round, nucha unde-
veloped. Wing with basal cell and vein setu-
lose or bare; speculum present; relative
lengths of wing veins: marginal > post-
marginal > stigmal; stigma unenlarged, only
2-3 as wide as stigmal vein. Legs with
coxae relatively small; hind tibia with one
apical spur. Petiole much longer than wide,
cylindrical; areolate dorsally, strigose lat-
erally; with basal flange laterally and ven-
trally; lateral setal row sometimes present
as a few weak setae anteriorly. Gaster ovate;
T1 enlarged, nearly concealing succeeding
terga, hind margin convex; hypopygium
reaching to tip of gaster.
Male.—Similar to female, but antenna
with funiculus parallel-sided, all segments
elongate: club lacking area of micropilosity,
palps unmodified.
VOLUME 90, NUMBER |
@
Tat
9 0)
10)
Figs. 1-8.
fore wing showing arrangement of dorsal setae along basal vein and admarginal setae. 4, Female head (dorsal
view). C. dixiana n. sp. 5, Female propodeum and petiole. 6, Female head (anterior view). 7, Female head
(dorsal view). 8, Female fore wing showing bare basal vein and arrangement of ventral admarginal setae.
Diagnosis. — Cryptoprymna can be distin-
guished by the following unique combina-
tion of characters: edentate clypeus; large
area of micropilosity on club; carinate
pronotum; propodeum arched and as long
as scutellum, median carina and plicae de-
veloped; elongate sculptured petiole; en-
larged Tl; hypopygium extending to tip of
Cryptoprymna atra (Walker). 1, Female whole body. 2, Female propodeum and petiole. 3, Female
gaster, and loss of metallic coloration.
Though these character states are all apo-
morphic within the Miscogasterinae, none
of these characters is unique to this genus.
This combination of apomorphies 1s unique,
however.
Biology.—Little is known of the biology
of the species 1n this genus. Graham (1969)
4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mentions specimens of C. atra taken on
Abies sp. and Pinus sylvestrus L. Among the
Nearctic specimens of C. afra, is one from
Stockholm, Maine, mounted with a syrphid
pupa that has a lateral emergence hole and
a label reading “beaten from fir.” A speci-
men of C. dixiana n. sp. from Fort Pierce,
Florida, was reared from a similar pupa, but
there is no information on the plant source
of this pupa. It seems likely that species of
this genus are parasitoids of syrphids on co-
nifers.
Key To NEARCTIC SPECIES OF
CRYPTOPRYMNA FORSTER
1. Wing with basal cell and vein setate, ventrally
with a patch of setae behind the marginal vein
(Fig. 3). Occiput concave in dorsal view (Fig.
4). Petiole bare (Fig. 2) atra (Walker)
— Wing with basal cell and vein bare, ventrally
with only a single row of setae behind the mar-
ginal vein (Fig. 8). Occiput straight in dorsal
view (Fig. 7). Petiole with short lateral setal
row (Fig. 5) dixiana Nn. sp.
Cryptoprymna atra (Walker)
(Figs. 1-4)
Prosodes atra Walker, 1833: 375. Lectotype
male (designated by Graham 1969) in the
Westwood collection (BMNH) (not seen).
Westwood, 1840: 68-69. Haliday, 1842:
V, Plate C (figure). Walker, 1872: 94 (fig-
ure); 1873: 371 (figure). Gahan and Fa-
gan, 1923: 41, 121.
Cryptoprymnus cavigenaThomson, 1878:
22. Lectotype female (designated by Gra-
ham 1969) in the collection of Univer-
sitetets Zoologiska Institutionen, Lund
(not seen).
Cryptoprymna atra (Walker): Schmiede-
knecht, 1909: 380. Delucchi, 1955: 174
(synonymy). Boucek, 1961: 71 (distri-
bution). Graham, 1969: 140-141 (biolo-
gy, synonymy, distribution). Boucek,
1976: 14-15. Dzhanokmen, 1978: 80.
Diagnosis. —In addition to the characters
given in the key, female C. atra differ from
C. dixiana in the following ways: the trun-
cate portion of the clypeus of C. atra has a
concave anterior margin and the anterior
lateral corners are sharp while the anterior
margin is straight and the corners rounded
in C. dixiana; the antennal flagellum of C.
atra is longer, 0.89 + (S.E.) 0.014 (n = 4)
times as long as the head width compared
with 0.81 times in C. dixiana; the antennal
club of C. atra is more slender, 2.1 + 0.21
times as long as wide compared to 1.6 times
in C. dixiana; and the wings of C. atra are
longer, 2.2 + 0.13 times the mesosomal
length versus 1.7 times in C. dixiana.
Biology.—The specimen from Stock-
holm, Maine, was reared from the pupa of
a syrphid which was “beaten from fir.”
Nearctic material examined (CNC, INHS,
USNM): Canada. BRITISH COLUMBIA:
Terrace, 8-VIII-1960, 1 ¢. NEW BRUNS-
WICK: Acadia Experiment Station (Fred-
ricton) 1-17-VH-1970, 1 4, 13-VIII-1970,
1 6. QUEBEC: Messines, 10-VII-1947, 1 4;
Parke Reserve (near St. Eleuthere), 1 3-VIII-
1957, 1 6. United States. MAINE: Stock-
holm, 6-VI-1955, 1 2°. MICHIGAN: Isle
Royale, 3-7-VHI-1936, 1 °. OREGON:
Saddleback Mt. (near Rose Lodge), 1 1-VIII-
LOGIS le
The records of C. atra from Greenland
cited by Bakkendorf (1955) are in error
(Boucek 1961).
Cryptoprymna dixiana,
NEw SPECIES
(Figs. 5-8)
Description. — Holotype female: Color.
Body black with mesosoma tinged blue,
scutellum and propodeum tinged gold. An-
tenna with scape brownish yellow; pedicel
and flagellum brown. Legs reddish brown,
femora and mid tibia with dark bands; tarsi
light brown, pretarsus dark brown. Head
and mesosoma with scattered, short (one
half ocellar diameter) white setae.
Sculpture. —Clypeus, median area of face
subareolate; face laterally and dorsally, frons,
vertex finely alveolate; gena coriaceous.
Mesosoma with pronotal collar with trans-
verse row of punctures posterior to anterior
VOLUME 90, NUMBER 1
transverse Carina; mesoscutum alveolate,
side lobes more finely so than median lobe;
scutellum areolate. Gaster T1 polished; T7
coriaceous.
Structure.— Mesosomal length 0.85 mm.
Relative lengths of head, mesosoma, gaster;
16:42.5:33. Head broadly oval in anterior
view (Fig. 6), width 1.2 height (33:27),
2.1 length (33:16); clypeus with anterior
margin nearly straight mesally, anterior cor-
ners rounded; eye height 1.4 length (16:
11), 2.1 x malar length (16:7.5); POL 1.3 x
OOL (8:6), 1.5 x LOL (8:4); occiput straight
in dorsal view (Fig. 7). Antenna inserted a
quarter of the way up eye; scape length 0.88 x
eye height (14:16), reaching to median ocel-
lus, slightly recurved; length of pedicel plus
flagellum 0.82 head width (27:33); rela-
tive lengths of segments (annelli omitted,
club taken as a unit) scape = 14:3:2.5:3:3:
2.5:2.5:2:8; widths of F1, F6, club as 2:4:5;
F1-2 elongate, F3-—4 quadrate, F5—6 trans-
verse; MPP sensillae two thirds length of
segment, arranged in single row; club length
1.6 x width (8:5), asymmetrically curved to
outside, sutures oblique, area of micropi-
losity extending to midway down C2, C3
pointed apically. Mesosoma with meso-
scutal length 0.48 x width (12:25); scutellar
length 0.93 width (13:14); propodeum
with costula rugiform, nucha coriaceous,
supracoxal flange drawn out over base of
hind coxa. Wing length 2.4 x width (71:30);
basal cell and basal vein bare (Fig. 8); costal
cell with single row of setae; relative lengths
of submarginal, marginal, postmarginal, and
stigmal veins = 30:14:10:7. Petiole 0.96 x
as long as propodeum (13:13.5) (Fig. 5);
length 2.4 maximum width (13:5.5); with
a central and diverging lateral carinae on
basal one fourth; lateral setal rows present
as a patch of a few setae anteriorly. Gaster
1.6 as long as wide (35:22); T5-6 with
distal fringe of setae; hypopygium with short
erect white setae.
Allotype male: Color. Similar to female
but funiculus, club dark brown; femora dark
brown, lighter distally. Structure. Antenna
with flagellum parallel-sided, length of ped-
icel plus flagellum 1.3 x head width (39:31);
scape length 0.80 x eye height (12:15). Pet-
iole longer (petiole 1.2 propodeal length
[14:12]), more slender (petiolar length 2.8 x
width [14:5]). Gaster with terminal seg-
ments glabrous.
Diagnosis.— Characters for separating C.
dixiana from C. atra are given in the key
to Nearctic species and in the discussion
section for C. atra. C. dixiana can be dis-
tinguished from C. africana by the same
characters given in the key for distinguish-
ing C. dixiana from C. atra, except that C.
dixiana and C. africana both lack setae on
the basal cell and basal vein. The straight
occiput and lack of setae on the basal cell
of C. dixiana distinguishes that species from
C. brama.
Biology.—The allotype male from Ft.
Pierce, Florida, is mounted with a syrphid
pupa.
Etymology.—The name is a latinization
of Dixie, referring to the southeastern United
States distribution of this species.
Type material.— Holotype female is from
Andrews, South Carolina, and was collected
8 May 1963 by R. D. Eikenbary (USNM).
The allotype male is from Ft. Pierce, Flor-
ida, and was collected 26 April 1955 by
Holtzburg (USNM).
Polstonia, New GENUS
Type-species.—Polstonia quadriplana
Heydon. The gender is feminine. It is my
pleasure to name this genus in honor of Jane
Polston with whom I have spent many hours
collecting.
Description.—Color: Head, mesosoma,
and coxae dark blue to green; metasoma
dark reddish brown to black. Wing hyaline.
Female: Face slightly bulging viewed in
profile; clypeus subareolate, anterior margin
straight or slightly produced: genal concav-
ity short, reaching only one fourth the dis-
tance to lower orbit; eye bare, bulging in
anterior view; ocellar triangle width 1.5 x
length; occiput acarinate, moderately con-
cave. Antenna inserted below middle of face,
6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
just above a line between lower orbits; for-
mula 1:1:2:6:3; scape slender (length 7 x
width), extending to mid ocellus or higher;
funicular segments with MPP sensillae in a
single row; club with ventral patch of mi-
cropilosity and terminal spinelike protu-
berance on C3. Mandible 4-toothed, upper
two smaller and approximated or equally
spaced. Mesosoma arched dorsally; prono-
tum with collar lacking anterior transverse
carina, smooth strip along hind margin oc-
cupying a third to a half median length;
mesoscutum with notauli present as shallow
furrows, traceable to hind margin as strip
of distinct texture; scutellum as long as wide,
with 4-6 pairs of lateral setae, frenal sulcus
obscure or absent; prepectus acarinate; me-
sopleuron with upper epimeron smooth;
propodeum with plicae and median carina
complete and distinct, median panels al-
veolate-rugose, nucha obscurely sculptured
crescent. Wing with basal vein setate; specu-
lum present; relative lengths of wing veins:
marginal > postmarginal > stigmal; stigma
small, width only 2-3 width of stigmal
vein. Hind tibia with one apical spur. Pet-
iole sculptured dorsally, length 2-3 x width;
lateral setal row extending nearly entire
length of petiole, setae projecting perpen-
dicularly. Metasoma ovate, plicate ventral-
ly near insertion of petiole; Tl and T2 sub-
equal in length, distinctly longer than the
succeeding terga, Tl with hind margin
straight or sinuate.
Male: Similar to female but club lacking
area of micropilosity, palpi unmodified.
Diagnosis.—The possession by Polstonia
of an edentate clypeus, genal concavities, a
thirteen-segmented antenna, the female an-
tennal club with terminal spine and ventral
patch of micropilosity, propodeum with
distinct median carina and plicae, and an
elongate and reticulate petiole makes this
genus phenetically similar to Toxeuma
Walker. It differs from Toxeuma by having
an acarinate pronotal collar, notauli obscure
posteriorly, frenal suture obscured, and
lengths of Tl and T2 subequal. In these
characters, Polstonia resembles Sphegigas-
ter. However, Sphegigaster species have a
bidentate clypeus, never have the female
antennal club with a terminal spine and only
rarely with a ventral patch of micropilosity,
their propodeum lacks the median carina
and plicae, and the hind margin of gastral
T1 is broadly concave. In contrast, Polston-
ia species have an edentate clypeus, the an-
tennal club in the female with a terminal
spine and ventral patch of micropilosity, a
distinct median carina and plicae on the
propodeum, and gastral Tl has a nearly
straight hind margin. The elongate petiole
with complete lateral rows of setae that stick
out perpendicularly is the one unique apo-
morphic character defining this genus.
Polstonia can be included in Graham’s
(1969) key to the Sphegigasterini by mod-
ifying the first half of couplet one so it goes
to couplet la instead of 2, and then inserting
the following couplet after the first:
la. Clypeus simple (Fig. 9). Petiole with lateral
row of setae extending at least half its length
(Figs, WiGand )2)) cs. 1s. . Polstonia Heydon
— Clypeus bidentate or tridentate. Petiole with
a short row of setae extending less than half
its length
th
Key TO SPECIES OF
POLSTONIA HEYDON
1. Petiole less than 2.3 x as long as wide, rounded
dorsally, and areolate with reticulations only
2 as long as wide. Propodeum with area be-
tween the basal foveae relatively smooth and
divided into four equal sized regions (Fig. 11).
Usually only the hind femur with basal dark
tened dorsally, and strigulate dorsally with re-
ticulations three or more times as long as wide.
Propodeum areolate between basal foveae,
sublateral carinae weak or absent (Fig. 12). All
femora with dark bands basally ............
Sees ceseee . pelagocorypha n. sp.
Polstonia quadriplana, New SPECIES
(Figs. 9-11)
Description. — Holotype female: Color.
Head, mesosoma, coxae dark green with
coppery reflections; occiput, neck, pleural
VOLUME 90, NUMBER 1
Figs. 9-10. Polstonia quadriplana n. sp. 9, Female head (anterior view). 10, Female whole body.
regions, petiole darker; propodeum paler.
Gaster black with greenish reflections. An-
tenna with scape brownish yellow; pedicel
brown; flagellum black. Mandible brownish
yellow; teeth reddish brown. Legs brownish
yellow; strong dark bands on mid tibia and
hind femur with greenish reflections, fore
femur with weak broad dark band: pretarsus
darker. Wing veins pale brown. Head, dor-
sum of thorax with distinct brown setae.
Sculpture.—Clypeus subareolate; face,
frons, vertex, occiput alveolate; neck alveo-
late; mesoscutum, scutellum, axilla coarsely
alveolate; propodeum with median panels
areolate-rugose; petiole finely areolate, cells
2x as long as wide; gastral terga 4-7 sub-
imbricate.
Structure.— Mesosomal length 0.96 mm.
Relative lengths of head, mesosoma, me-
tasoma = 15:48:42. Head width 1.2 x height
Female propodeum and petiole.
8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(37:30) (Fig. 9), 2.5 x length (37:15); clypeus
with anterior margin slightly produced; eye
height 1.3 width (18:13.5), 2.0x malar
length (18:9); POL 1.4x OOL (7:5), 2.3 x
LOL (7:3). Antenna with length of pedicel
plus flagellum 0.97 head width (36:37);
relative lengths of segments (annelli omit-
ted, club counted as a unit) scape = 17:6:5:
4:3.5:2.5:3:3:7.5; widths of Fl, F6, club as
3:4.5:5: funicular segments narrowed at
bases; Fl-2 elongate, F3 quadrate, F4-6
transverse; MPP sensillae prominent; club
length 1.5 < width (7.5:5), sutures oblique.
Mesosomal length 1.7 x width (48:29); scu-
tellar length 0.97 x width (15.5:16); dorsel-
lum tilted nearly perpendicularly with re-
spect to lateral metanota (Fig. 10), anterior
edge regularly rounded, surface with little
sculpture; propodeum with spiracles round,
groove between basal foveae smooth and
divided by the median and two short sub-
median carinae into four subequal areas (Fig.
11). Wing length 2.2 x width (85:38); rela-
tive lengths of submarginal, marginal, post-
marginal, and stigmal veins = 35:18:16:11;
basal vein marked by row of eight setae;
basal cell with one seta distally on left wing.
Petiole length 1.9 width (13:7); 1.1 x as
long as propodeum (13:12); rounded dor-
sally, median carina complete and sharp;
length of setae in lateral setal rows a half to
three fourths width of petiole. Metasoma
broadly oval; length 1.4 x width (42.5:32.5);
basal region with scattered setae laterally;
T4-7 with submarginal row of setae.
Allotype male: Color similar to female ex-
cept head and dorsum of mesosoma lacking
coppery reflections; antennal flagellum
brown. Sculpture similar to holotype. Me-
sosomal length 0.98 mm. Antenna with
pedicel plus flagellum 1.2 x head width (43:
35); relative lengths of segments 15:5:5:4.5:
4.5:4.5:4:4:9; width of Fl, F6, club as 3:3.5:
3.5. Petiole length 2.0 x width (14:7). Wings
with basal vein having five setae on right
wing and four on left.
Variation.—Female mesosomal length
varies between 0.76 and 0.96 mm. Body
color varies from bluish black, to greenish
black, to dark green with coppery reflections
like the holotype. A female from Ohio has
all the femora with dark bands basally. The
groove between the basal foveae is some-
times weakly sculptured posteriorly, but
there is always a smooth strip of at least one
spiracular outside diameter along anterior
margin of propodeum. There are occasional
specimens with two strong sublateral cari-
nae on one side or the other. Male meso-
somal length varies between 0.90 and 0.98
mm. The body color variation is similar to
that of females.
Diagnosis. —In addition to the characters
given in the key, the vertex of female P.
quadriplana is usually nearly concolorous
with the dorsum of the mesosoma; in P.
pelagocorypha, the vertex is distinctly paler.
The antenna of female P. quadriplana has
each funicular segment narrowed basally,
the prominent MPP sensillae give the seg-
ments a coarse texture, and the club varies
between 1.4 and 2.0 times as long as wide.
The antenna of female P. pelagocorypha has
cylindrical funicular segments with a rather
smooth texture, and the club varies between
2.0 and 2.6 times as long as wide. In male
P. quadriplana, the combined length of the
pedicel and flagellum is between 4.1 and 4.8
times as long as the club length; it is between
3.6 and 3.9 times as long as the club in P.
pelagocorypha. The dorsellum of P. quad-
riplana 1s usually smooth and tilted nearly
perpendicularly with respect to the metano-
ta. In P. pelagocorypha, the dorsellum is
usually alveolate and in nearly the same
plane as the metanota. The lateral setae on
the petiole are shorter in P. quadriplana,
only a half to three fourths the width of the
petiole (Fig. 11); they are nearly as long as
the petiole width in P. pelagocorypha (Fig.
12). The petiole always has a distinct and
complete median carina in P. guadriplana
(Fig. 11); in P. pelagocorypha, the median
carina may be lacking or incomplete, or there
may be multiple fine longitudinal rugae (Fig.
12).
VOLUME 90, NUMBER 1
Biology.—The host(s) is unknown; how-
ever, the numerous specimens from Nova
Scotia were collected during a study in which
apple trees were fumigated and the arthro-
pods on them were collected on sheets be-
neath the trees (W. R. M. Mason, pers.
comm.).
Etymology.—The name comes from the
Latin words quadrus, meaning fourfold, and
planus, meaning flat or level, and refers to
the four smoothish areas along the anterior
margin of the propodeum, which are diag-
nostic of this species.
Type material.— Holotype female is from
Mt. Ste. Marie Low, Quebec, and was col-
lected 20 September 1965 by J. R. Vock-
eroth (CNC). The allotype male is from
Cooper’s Rock State Forest (near Morgan-
town), West Virginia, and was collected 22
June 1964 by O. Peck (CNC). Fifty-three
paratypes are as follows (CNC, INHS,
USNM): Canada. BRITISH COLUMBIA:
Cultus Lake, 14-VII-1948, 1 @. NEW
BRUNSWICK: Kouchibouguac National
Park, 12-IX-1977, 1 2. NOVA SCOTIA:
Aldershot, 4-VII-1952, 2 8, 1 9, 15-VH-
1952, 1 6, 7 9, 28-VII-1952, 11 9, 8-VII-
1952, 4 2, 18-VII-1950, 4 2, 9-IX-1950, 1
2. ONTARIO: Innisville, 18-VIII-1963, 1
2; Simcoe, 19-VI-1939, 1 92. QUEBEC: Lac
Brulle, 15-VII-1946, 1 3, Mt. Ste. Marie
Low, 20-IX-1965, 3 2; Old Chelsea, 3-VII-
1969, 1 4, S-VIII-1969, 9 2, United States.
NEW YORK: Lake Placid, 15-VIII-1896,
1 2; Otter Lake (near Meridian), 25-VII-
1946, | ¢. OHIO: Barberton, 30-VI-1936,
1 9. VIRGINIA: Monterey, 22-VI-1964, | 2.
Polstonia pelagocorypha,
New SPECIES
(Fig. 12)
Description. — Holotype female: Color.
Head with face, frons dark green; vertex
blue-green; occiput greenish black. Antenna
with scape brownish yellow; pedicel, flagel-
lum dark reddish brown. Mandible yellow-
ish brown; teeth reddish. Mesosoma with
dorsum dark green; pleural area, coxae, pro-
podeum, petiole, gaster bluish black. Legs
brownish yellow except trochanters, basal
two thirds of fore and mid femora, hind
femur brown (hind femur with traces of me-
tallic coloring); pretarsus black. Wing with
veins pale brown. Head and mesosoma with
pale brown setae.
Sculpture.—Pattern similar to P. guad-
riplana except texture delicate, particularly
on head, and petiole strigulate dorsally, cells
three or more times as long as wide.
Structure.—Mesosomal length 0.92 mm.
Head width 1.2 x height (34:27), 2.3 x length
(34:14.5); clypeus with anterior margin
slightly produced and reflexed: eye height
1.2 length (15:12), 1.7 x malar length (15:
APOE <x OOL.(7.5:5), 2.1 x OOL GS:
3.5). Antenna with length of pedicel plus
flagellum 1.0 x head width (34.5:34); rela-
tive lengths of antennal segments (annelli
omitted, club counted as a unit) scape = 15:
5:3.512.5:3:2.523:2.5:10:5; widths of Fl, F6,
club as 3:3.5:4; funicular segments cylin-
drical; MPP sensillae fine, club length 2.6 x
width (10.5:4), sutures only slightly oblique.
Mesosoma 1.7 as long as wide (46:26.5);
scutellar length 0.86 width (12:14); dor-
sellum in same plane as metanota, anterior
margin scalloped, finely alveolate: propo-
deum with groove between basal foveae
subareolate, with short weak sublateral ca-
rinae (Fig. 12). Wing length 2.2 x width (89:
41); relative lengths of submarginal, mar-
ginal, postmarginal, stigmal veins as 32:18.5:
17:10; basal vein marked by row of nine
setae on left wing; one seta in basal cell of
left wing. Petiole length 3.2 x width (16:5),
1.3 as long as propodeum (16:12); flat-
tened dorsally; median carina visible only
in posterior third: length of lateral setae
nearly equal to width of petiole (Fig. 12).
Gaster fusiform, length 1.5 x width (35:23);
succeeding terga withdrawn beneath T2
(specimens air-dried).
Allotype male: Color. Similar to holotype
but paler, dorsum of mesosoma green with
faint yellowish reflections, antennal pedicel
10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and flagellum brown, bands on femora dark
but extending only one third length of mid
and three fourths length of hind femora.
Mesosomal length 0.84 mm. Antenna with
pedicel plus flagellum 0.98 x as long as head
width (32.5:33); relative lengths of segments
as 13:4.5:4:3.5:3.5:3.5:3.5:3.5:9; parallel-
sided, widths of Fl, F6, club as 3:3:3. Pet-
iole length 2.6 width (16.6). Wing with
basal cell bare except for a couple of setae
adjacent to setal row on basal vein on right
wing.
Variation. — The female mesosomal length
varies between 0.77 and 0.96 mm. Body
color varies from bluish black to dark green.
The female from North Carolina has the
vertex green; in the other females it is blue-
green like the holotype. The groove between
the basal foveae of the propodeum is some-
times crossed by one or more weak carinae,
but these are less than one spiracular outside
diameter in length. The petiole varies be-
tween 2.6 and 3.0 times as long as wide, and
its dorsal surface is either acarinate, with
weak or incomplete median carina, or with
several long longitudinal rugae. The male
mesosomal length varies between 0.67 and
0.94 mm. The color of the males varies from
green with coppery reflections to bluish
black. The male from Illinois has a brown-
ish yellow flagellum and very weak dark
bands on the femora. The petiole varies be-
tween 2.7 and 3.0 times as long as wide.
Diagnosis.—For a detailed diagnosis see
that of P. quadriplana above.
Biology.—The host(s) of this species is
unknown.
Etymology.—The name is from the Greek
words pe/agos, meaning sea, and koryphe,
meaning top of the head, and refers to the
sea-green vertex of the female.
Type material.—Holotype female
(AMNH) is from 1.5 miles SW of Lolo Hot
Springs, Montana, and was collected 22 July
1978 by N. L. Herman. Allotype male (CNC)
is from Whiteface Mountain, New York,
and was collected 19 July 1962, by J. C.
Chillcott. Thirteen additional paratypes are
as follows (CMNH, CNC, FSCA, INHS,
USNM): Canada. ALBERTA: Edmonton,
20-VI-1937, 1 6. NEW BRUNSWICK:
Fundy National Park, 10-VII-1970, 1 9°;
Kouchibouguac National Park, 10-XI-1977,
1 2. QUEBEC: Duchensay, 5-VII-1953, 1 @.
SASKATCHEWAN: White Fox, 18-VII-
1944, 1 2. United States. ALASKA: Mata-
nuska, 6-X-1945, | 4; Palmer, 1-VIII-1948,
1 6. ILLINOIS: MacLean Co., 30-V-1883,
1 2; Urbana, 10-VI-1928, 1 6. MICHIGAN:
Manistee Co., 5-VII-1957, 1 6. NORTH
CAROLINA: Lake Junaluska, 27-V-1954,
1 ¢. WEST VIRGINIA: Spruce Knob,
5-VIII-1960, 1 4, Weston, 13-18-IX-1938,
Id:
ACKNOWLEDGMENTS
I thank the following persons for the loan
of material: M. Favreau, American Mu-
seum of Natural History (AMNH), New
York, NY; Dr. G. Wallace, Carnegie Mu-
seum of Natural History (CMNH), Pitts-
burgh, PA; Dr. C. M. Yoshimoto, Canadian
National Collection (CNC), Ottawa, ON; Dr.
H. Weems, Florida State Collection of Ar-
thropods (FSCA), Gainesville, FL; Dr. G.
W. Byers, Snow Entomological Collection
(SEC), Lawrence, KS; and Dr. E. E. Grissell,
United States National Museum (USNM),
Washington, D.C. I would like to express
my appreciation to J. K. Bousman and Dr.
G. L. Godfrey of the Illinois Natural His-
tory Survey (INHS), Champaign, IL, and
two anonymous readers for reviewing this
paper. I would like to thank my advisor, Dr.
W. E. LaBerge (INHS) for help and en-
couragement, J. Sherrod for assistance with
the illustrations, and the staff at the Center
of Electron Microscopy at the University of
Illinois at Urbana-Champaign.
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Peck, O., Z. Boucek, and A. Hoffer. 1964. Keys to
the Chalcidoidea of Czechoslovakia. Mem. Ento-
mol. Soc. Can. 34: 1-120.
Schmiedeknecht, O. 1909. Hymenoptera family
Chalcididae. Jn P. Wytsman, Genera Insectorum
97: 1-550.
Thomson, C. G. 1878. Hymenoptera Scandinaviae.
5. Pteromalus (Svederus) continuatio. Lund.
Walker, F. 1833. Monographia Chalcidum. Art.
XLIII. Entomol. Mag. 1: 367-384.
1872. Notes on Chalcididae. Part VI. Hor-
moceridae, Sphegigasteridae, Pteromalidae, Elas-
midae, Eulophidae, Entendonidae, Tetrastichidae,
Trichogrammidae, pp. 89-105. London.
. 1873. Economy of Chalcididae. Entomolo-
gist 6: 322-324.
Westwood, J. O. 1840. Synopsis of the genera of
British insects, pp. 1-158. [Bound with: J. O.
Westwood, An introduction to the modern clas-
sification of insects, Vol. 2.] London.
1842. [No Title] Entomologist 1: v—
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 12-19
A NEW GENUS AND TWO NEW SPECIES OF PANGONIINI
(DIPTERA: TABANIDAE) OF ZOOGEOGRAPHIC
INTEREST FROM SABAH, MALAYSIA
JOHN F. BURGER
Department of Entomology, Nesmith Hall, University of New Hampshire, Durham,
New Hampshire 03824.
Abstract. —A new genus of Pangoniini, Mesopangonius Burger, 1s described from Mount
Kinabalu, Sabah, Malaysia. This is the first representative of this tribe known to occur
in the Oriental Region. Although possibly derived from unspecialized Laurasian Pango-
niini, its long, slender proboscis is characteristic of specialized genera of the tribe. Two
new species, philipi and brackleyae are described in the genus, and a key is provided.
A recent small collection of Tabanidae
from Mount Kinabalu, Sabah, Malaysia, by
entomologists from the Smithsonian Insti-
tution, Washington, D.C., yielded two re-
markable new species in an undescribed ge-
nus of the tribe Pangoniini. These are the
first representatives of this tribe known from
the Oriental Region, and are particularly in-
teresting because they combine anatomical
features of both generalized and specialized
genera of Pangonuini.
Mesopangonius Burger, New GENUS
Type species.—Mesopangonius philipi
Burger, Sabah, MALAYSIA, by original
designation.
Medium-sized (12-16 mm long) rather
slender to moderately stout-bodied Esen-
beckia-like species with well-developed
ocelli; eyes bare with no color pattern; fron-
tal index 2.5-3.0. Antennal bases closely ap-
proximated; flagellum subulate, bearing 8
annuli; basal annulation enlarged (Figs. 2C,
4C), about twice as long as high; apical an-
nulation greatly elongated, about one-half
Scientific Contribution Number 1485 from the New
Hampshire Agricultural Experiment Station.
length of remaining annulations combined.
Proboscis slender, length 1.5-1.9 times
height of head; labella long, slender and
sclerotized (Figs. 2A, 4A); 2nd maxillary
palpomere short and subcylindrical or
somewhat flattened on outer surface, and
bearing a shallow concavity. Legs long and
slender, hind tibial spurs well-developed.
Base of vein R, with a long spur, basal sec-
tion of Cu bare or with 2-3 scattered setulae,
Sc bare dorsally and ventrally. Female gen-
italia (Fig. 2E-G): ninth tergite entire, rel-
atively broad and heavily sclerotized lat-
erally, narrowed and weakly sclerotized
medianly; tenth tergite divided medially;
cerci rounded apically, length and width
subequal; eighth sternite shield-shaped, very
weakly sclerotized; apical lobes of the an-
terior gonapophyses deeply divided medi-
ally, about as long as wide, distance between
lobes about two-thirds width of individual
lobes; arms of the genital fork with wing-
like expansions apically; distal ends of sper-
mathecal ducts membranous, unexpanded.
Mesopangonius philipi Burger,
NEw SPECIES
Female (Fig. 1).—Length: body 12-14
mm; wing 13-15 mm. Front (Fig. 2B) yel-
VOLUME 90, NUMBER 1
Fig. 1.
Mesopangonius philipi sp. n., female. (8 x)
lowish brown laterally, slightly diverging
below, with broad dark brown pollinose
ridge in middle extending from subcallus to
vertex, bordered by an irregular row of semi-
erect black setulae; frontal index 2.8-3.2,
divergence index 1.2—1.3. Ocelli large and
prominent, borne on a conspicuous tuber-
cle; vertex depressed below upper margin
of head. Subcallus, gena and face brown pol-
linose, gena sparsely clothed with brown
hairs; face moderately produced, upper lat-
eral surface with patch of brown hairs; beard
rather sparse, with brown hairs anteriorly,
pale yellowish ones posteriorly. Antenna
(Fig. 2C) yellow brown, slightly darker api-
cally; bases closely approximated, distance
between them distinctly less than width of
scape; scape and pedicel yellowish brown
pollinose, black setose; flagellum subulate,
with 8 annuli; basal annulation enlarged,
about one-third broader than succeeding
annulation and bearing a dense tuft of black
setulae at apex of upper margin, apical an-
nulation three times length of penultimate
annulation. Proboscis slender (Fig. 2A),
length 1.7-1.9 times head height; labella long
and slender, sclerotized. Maxillary palpus
(Fig. 2D) with apical palpomere short and
slender, subcylindrical, length less than one-
fifth that of proboscis, bearing long, black
semi-erect setae on outer surface, basal pal-
pomere slightly broader than apical seg-
ment, bearing long, black semi-erect setae.
Eye bare, unpatterned (relaxed), rather
coarsely faceted.
Mesonotum light brown, bearing semi-
erect brown hairs, except pale yellowish
white ones anteriorly near the head; noto-
pleural lobe concolorous with mesonotum;
scutellum slightly paler; pleuron paler yel-
lowish brown, bearing pale yellow hairs, ex-
cept dark brown ones posteriorly on mesan-
14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2.
Antenna. (32 x) D, Maxillary palpus. (32 =
and anterior gonapophyses. (60 x) G, Genital fork and caudal ends of spermathecal ducts. (90 x)
episternum. Legs slender, elongate,
unicolorous pale yellowish brown, bearing
mixed pale yellowish and dark brown hairs;
apical spurs on hind tibia nearly as long as
those on mid-tibia. Wing lightly brown tint-
ed throughout; R, with long spur; cells r;
and m, open to wing margin. Halter light
brown.
Mesopangontius philipi sp. n., female. A, Profile of head. (16) B, Frontal view of head. (16x) C,
) E, Eighth, 9th tergites, cerci, dorsal view. (60 =) F, Eighth sternum
Abdomen pale greenish brown, with some
yellowish tones intermixed but without def-
inite pattern; tergite | slightly paler; tergites
5-7 slightly darker; all tergites bearing pre-
dominantly dark hairs, with some pale yel-
lowish ones intermixed anteriorly and lat-
erally; ventral surface concolorous.
Holotype 2, MALAYSIA: Sabah; Kina-
VOLUME 90, NUMBER 1
Fig. 3.
balu National Park, Headquarters area, el.
1560 m, 9 Sept. 1983, G. F. Hevel & W. E.
Steiner (National Museum of Natural His-
tory, Washington, D.C. (NMNH)).
Paratypes, MALAYSIA: 2 8, 8, 13 Sept.
1983. Same data as holotype (NMNH; J. F.
Burger Collection).
I take great pleasure in naming this species
for the late Cornelius Becker Philip, inde-
fatigable student of Tabanidae, who con-
tributed much to our knowledge of Oriental
Tabanidae.
Mesopangonius brackleyae sp. n., female. (8 x)
Mesopangonius brackleyae Burger,
New SPECIES
Holotype female.—Length: body 15.6
mm; wing 16 mm (Fig. 3). Front (Fig. 4B)
dark brown pollinose, slightly diverging be-
low, middle with a poorly-defined raised
ridge from subcallus to vertex, bearing an
irregular median subshining black area and
bordered by an irregular row of black set-
ulae; frontal index 2.5, index of divergence
1.2. Ocelli large and prominent, borne on a
16
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4.
conspicuous tubercle at vertex. Subcallus
concolorous with front. Gena and face gray
pollinose, except face shining dark brown
along lower margin; face moderately pro-
duced, dorsolateral surfaces with patch of
dark brown hairs; beard mostly pale yellow-
ish, except dark brown hairs anteriorly.
Scape and pedicel of antenna yellowish gray
pollinose, antennal bases closely approxi-
mated, distance between them distinctly less
than width of scape; flagellum subulate (Fig.
4C), with 8 annuli, yellowish brown, apical
annulation dusky brown; basal annulation
conspicuously enlarged, twice as broad as
second annulation, bearing 1-2 setulae on
upper surface; apical annulation 2.5 times
Mesopangonius brackleyae sp. n., female. A,
C, Antenna. (32x) D, Maxillary palpus. (32 =)
Profile of head. (16) B, Frontal view of head. (16x)
length of penultimate annulation. Proboscis
slender (Fig. 4A), length 1.5 times head
height, labella long and slender, sclerotized.
Maxillary palpus (Fig. 4D) short, apical pal-
pomere brown, distinctly flattened, length
one-fourth that of proboscis, bearing long
black setae on outer surface and narrow bare
median concavity. Eye bare, unpatterned
(relaxed), relatively coarsely faceted.
Mesonotum and scutellum subshining
dark brown, densely clothed with semi-erect
yellowish hairs; postpronotal lobe reddish
gray pollinose; notopleural lobe reddish;
pleuron grayish pollinose, except mesanepi-
sternum and anterior half of katepisternum
with blackish tones, densely yellow pilose.
VOLUME 90, NUMBER |
Legs slender, elongate; coxa and femur dark
brown, black pilose, except apex of femur
paler; tibia pale brown, bearing yellow hairs;
hind tibial spurs well-developed, subequal
to mid-tibial spurs; tarsus basally concol-
orous with tibiae, darker brown apically.
Wing light brownish tinted on anterior half,
subhyaline posteriorly; R, with long spur;
cells r; and m, open to wing margin. Halter
pale brown.
Tergite 1 of abdomen entirely pale yel-
lowish brown, pale yellow pilose: tergites 2—
4 dark brown on basal three-fourths, con-
trastingly paler brown on apical fourth, the
dark and light areas bearing black and pale
yellow hairs respectively; tergites 5-7 dark
brown, black-haired, with conspicuous yel-
low-haired posterior margins (Fig. 3); ven-
tral surface of abdomen with similar pattern
as dorsum, except sternite 2 predominantly
light brown, and only basal halves of ster-
nites 3-4 dark brown.
Holotype °, MALAYSIA: Sabah; Kina-
balu National Park, Headquarters area, el.
1560 m, 9 Sept. 1983, G. F. Hevel & W. E.
Steiner (National Museum of Natural His-
tory, Washington, D.C.).
I take pleasure in naming this striking
species for my good friend and colleague,
Frances Brackley, a specialist of the Orchi-
daceae, who first sparked my interest in dip-
terous pollinators of alpine plants.
The following key will separate the species
of Mesopangonius described above:
1. Abdominal tergites 2-5 dark brown, with paler
brown apices and complete yellow-haired in-
cisures; all femora dark brown, contrasting with
light brown tibiae; apical palpomere of max-
illary palpus somewhat flattened, outer surface
with a bare, shallow concavity brackleyae, n. sp.
— Abdominal tergites light brown throughout,
without strongly contrasting markings or hairs;
all femora and tibiae concolorous light brown;
apical palpomere of maxillary palpus narrow
and subcylindrical, lacking a bare concavity on
the outer surface .. philipi, n. sp.
DISCUSSION
Mesopangonius resembles Esenbeckia
Rondani, a predominantly Neotropical ge-
nus, but differs in having the basal annu-
17
lations of the flagellum not forming a par-
tially-fused, enlarged plate, a more slender
proboscis with very narrow, elongate label-
la, and with cell r; of the wing widely open
to the wing margin. The female terminalia
also are similar to Esenbeckia, differing pri-
marily in the broader ninth tergite (Fig. 2E),
the more rounded cercus, and the more
widely separated apical lobes of the anterior
gonapophyses (Fig. 2F).
Mackerras (1955) subdivided the genera
of Pangoniini into generalized (Group |) and
specialized (Group 2) moieties. Those gen-
era considered to be more generalized have
the r; cell of the wing open, proboscis stout
and subequal to head height, the labella dis-
tinctly enlarged and unsclerotized, and the
body usually slender or with the abdomen
parallel-sided. Most of the genera included
have a south or north temperate relict dis-
tribution in montane or desert environ-
ments. Fourteen of 18 genera in Mackerras’
Group | occur in coastal or desert North
America (5), the mountains of Chile and
Argentina (5), and in Australia (4). Two gen-
era occur in Japan, and one each in Brazil
and southern Africa.
Genera of Pangoniini considered to be
specialized have cell r; closed, or strongly
narrowed apically, proboscis slender, as long
as to much longer than head height, labella
narrow, sclerotized, and sometimes very
long, and the body usually stoutly-built.
These genera have a predominantly south-
ern Palearctic, amphi-Mediterranean (Pan-
gonius), or a new world tropical and sub-
tropical (Esenbeckia) distribution. Mack-
erras considered Austroplex Mackerras, from
Australia, to be a link between the two
groups because of its basally expanded an-
tennal flagellum, but most of its features can
be considered generalized.
Mesopangonius also has a preponderance
of primitive features, eyes bare and coarsely
faceted, cell r; of the wing open, relatively
slender body (somewhat stouter in brack-
leyae), basal flagellar annulations not con-
solidated into an enlarged plate, relatively
narrow, unspecialized maxillary palpus, the
18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
long, slender legs, and the broad and un-
divided ninth tergite. The principal spe-
cialized features are the long, slender pro-
boscis, the long, narrow, sclerotized labella,
and the deeply divided and relatively widely-
separated apical lobes of the anterior gon-
apophyses. The preponderance of primitive
features suggests that Mesopangonius 1s
closer to the generalized group of genera,
and that the elongate, narrow proboscis and
labella may be an adaptation to a particular
trophic niche. Mesopangonius also occurs
in a montane “temperate” area on Mount
Kinabalu at the northern end of the Crocker
Range in Sabah, an environment similar to
that where some genera of Pangoniini in
Mackerras’ Group | occur.
Since Borneo is a continental island as-
sociated with the Laurasian plate, Meso-
pangonius may be derived from a general-
ized Laurasian pangoniine stock. Genera of
Pangoniini presently known from Eurasia,
other than Mesopangonius, are Stonemyia
Brennan (Japan, Southwestern Asia [Cau-
casus], and possibly China), Nagatomyia
Murdoch & Takahasi (Japan), and Pan-
gonius Latreille (amphi-Mediterranean).
Mesopangonius differs most conspicuously
from Sfonemyia in having a more slender
body, a longer, more slender antennal fla-
gellum, with only the basal annulation en-
larged and a much longer apical annulation,
a longer, more slender proboscis with a long,
narrow, sclerotized labella, shorter palpus,
legs longer and more slender, ventral surface
of scutellum without bristles, R, of the wing
with a long spur, the larger, deeply-divided
lobes of the anterior gonapophyses, and the
caudal ends of the spermathecal ducts mem-
branous and delicate (Fig. 2G). It shares few
features with Nagatomyia other than the
presence of ocelli, the slender body and the
open cell r;. Mesopangonius differs from
Pangonius in having a more slender body,
ocelli larger and more prominent, basal an-
nulation of flagellum broader, apical pal-
pomere shorter relative to proboscis length,
and with the outer concavity, when present,
very shallow, the legs more slender and
elongate, cell r; of the wing open, and the
lobes of the anterior gonapophyses larger
and more deeply-divided medianly.
Mesopangonius bears little resemblance
to generalized Pangoniini associated with
the Australian plate, as one might expect
from tectonic evidence. Ectenopsis Mac-
quart has coarse eye facets and a narrow
cylindrical apical palpomere, but otherwise
has little in common with Mesopangonius.
Some species of Fidena Walker, a genus with
specialized features in the Scionini that has
radiated extensively in the Neotropical Re-
gion, have a proboscis configuration like that
of Mesopangonius, but otherwise are quite
distinct.
The only other representatives of the Pan-
goniinae known from the Oriental Region
are in the Philolichini (Philoliche Wiede-
mann). Of these, only species in the sub-
genus Buplex Austen of Philoliche share even
a superficial resemblance to Mesopango-
nius, and then only because they have ocell,
and some species have a narrow, elongate
proboscis. However, they are presently re-
stricted to southern Africa. The Oriental
species of Philoliche lack ocelli, have strong-
ly produced faces, closed wing cells, lobes
of the anterior gonapophyses more widely
separated, and other features that clearly ex-
clude them from close relationship with
Mesopangonius.
It is remarkable that Mesopangonius re-
mained uncollected and unknown for so
long. Its chance discovery suggests that oth-
er representatives of the Pangoniini possibly
may be found in remaining “temperate is-
lands” within tropical Asia, given sufficient
patience and collecting, before such refuges
disappear.
ACKNOWLEDGMENTS
I thank G. B. Fairchild, Bureau of Ento-
mology, Florida Department of Agriculture,
L. L. Pechuman, Cornell University, and
H. J. Teskey, Biosystematics Research
Centre, Ottawa, for their helpful comments
VOLUME 90, NUMBER | is)
about characters of the Pangoniini. I also LITERATURE CITED
thank Dz S. Chandler and Scott Sherman, Mackerras, I. M. 1955. The classification and distri-
University of New Hampshire, for review- bution of Tabanidae (Diptera) II. History: Mor-
ing the manuscript, and Tess Feltes, Ports- phology: Classification: subfamily Pangoniinae.
mouth, New Hampshire, for preparing the Australian Journal of Zoology 3: 439-511.
illustrations.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 20-27
APTERONA HELIX (LEPIDOPTERA: PSYCHIDAE), A PALEARCTIC
BAGWORM MOTH IN NORTH AMERICA: NEW DISTRIBUTION
RECORDS, SEASONAL HISTORY, AND HOST PLANTS
A. G. WHEELER, JR. AND E. RICHARD HOEBEKE
(AGW) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg,
Pennsylvania 17110; (ERH) Department of Entomology, Cornell University, Ithaca, New
York 14853.
Abstract.—New distribution records for Apterona helix (Siebold), an Old World bag-
worm moth detected in California in 1940 and in New York in 1962, are given for Michigan
and Pennsylvania, and additional New York records are provided; populations in western
and eastern North America are thought to represent separate introductions from Europe.
Seasonal history, habits, and host plants of this parthenogenetic, polyphagous species are
reported for populations observed in New York and Pennsylvania, and North American
distribution and economic importance in Europe and North America are reviewed. Char-
acters facilitating recognition of this psychid are provided, and the unusual helicoid or
snail-like larval case is illustrated.
Members of the psychid genus Apterona
Milliére are endemic to the Palearctic re-
gion, with 7 species occurring in the Med-
iterranean area, west and central Europe,
Asia Minor, Crimea, Caucasus, Iranian Pla-
teau, and southcentral Asia (Kozhanchikov
1956). Apterona helix (Siebold), a parthe-
nogenetic species, is widely distributed in
central and southern Europe, ranging east
to European USSR and Kirgiz SSR in cen-
tral Asia and south to Iran (Strand 1912,
Davis 1964).
The first confirmed North American re-
cord of 4. helix was based on an infestation
discovered at a private residence in Nevada
City, California, in June 1940 (Kiefer 1940,
Robinson 1953). In the western states, A.
helix now occurs in the northern half of
California (Eichlin 1985) and in portions of
Idaho, Nevada, Utah (Davis 1964), Oregon
(Every 1970), and Washington (Suomi
1986).
Apterona helix is also known in eastern
North America, having been detected near
Albany, New York (Loudonville), in June
1962 (Davis 1964); a second infestation was
soon discovered in Albany. Eastern popu-
lations are believed to be the result of an
independent European introduction rather
than to have originated from the established
western U.S. populations (Davis 1964).
Lenox, Massachusetts (Adamski 1984), is
the only other eastern record.
The name 4. crenulella (Bruand) has been
used in early (and some current) literature
on this adventive species in North America.
Some European workers considered helix to
be a parthenogenetic form of crenulella,
whereas others argued that they are distinct
species. In revising the Western Hemi-
sphere Psychidae, Davis (1964) retained he-
lix asa “‘facultative, parthenogenetic form,”
noting interfertility between crenulella and
helix might be expected but that “many
VOLUME 90, NUMBER 1
questions remain unanswered.” Apterona
helix is now accorded specific rank (Davis
1983, 1987).
Here we review its status as an econom-
ically important species, give additional lo-
calities for 4. helix in New York, and report
Michigan and Pennsylvania as new state
records. We summarize our observations on
seasonal history, habits, and host plants in
the East and give morphological characters
allowing this immigrant species to be rec-
ognized in the North American fauna.
ECONOMIC IMPORTANCE
The European literature indicates that 4.
helix occasionally is injurious. It has been
implicated in causing damage to apple, hor-
ticultural crops, and olive (Rev. Appl. Ento-
mol. (A) 3: 393, 1915; 4: 210, 1916; 56:
553, 1968).
In Utah, 4. helix has been observed skele-
tonizing leaves of apple trees and causing
extensive damage to many range plants; it
sometimes injures various cultivated plants
and becomes a nuisance when it congregates
on the walls and windows of homes (Tib-
betts and Knowlton 1952, Knowlton and
Roberts 1968). There also are records of
severe damage to cherry foliage in an or-
chard (Knowlton 1961), leafmining injury
to corn (Knowlton and Parrish 1965), and
destruction of green color in three acres of
barley and four acres of alfalfa (Knowlton
1966). In Idaho, 4. helix was extremely
abundant in alfalfa and sweetclover, causing
considerable skeletonizing of the foliage
(Gittins 1958). Marshall (1970) recorded
heavy damage to strawberry plants in Ne-
vada. In California, where this insect has
been called the garden bagworm, consid-
erable damage to several commercial crops
such as apple, cruciferous vegetables, and
chrysanthemums and other plants grown for
cut flowers has occurred when populations
are high (Keifer 1947, Robinson 1953).
Suomi (1986) reported that baby’s breath
(Gypsophila, Caryophyllaceae) used in dried
flower arrangements was so heavily infested
21
in one Washington county that plant ma-
terial could not be shipped out of state.
There are no reports of damage by 4. helix
to cultivated plants in eastern United States,
although Adamski (1984) stated that cases
were found attached to planted flowers and
vegetables, ornamentals, and shade trees. In
New York, the cases have attracted notice
when they attach to houses (Davis 1964).
Each year homeowners submit larval cases
to Cornell University’s Insect and Plant
Disease Diagnostic Laboratory; their con-
cern is with large number of cases that ac-
cumulate on houses and the paint that is
sometimes removed when cases are pulled
off (Klass 1983).
DISTRIBUTION IN EASTERN
NorTH AMERICA
In addition to published records from the
Albany, New York, area and Lenox, Mas-
sachusetts, the following new records are
available. Michigan, Pennsylvania, and
some of the New York records are based on
our collecting; other New York records
(those without collector names and mostly
lacking exact localities) were obtained from
the Insect and Plant Disease Diagnostic
Laboratory, Cornell University. Voucher
specimens have been deposited in the insect
collections of Cornell University and the
Pennsylvania Department of Agriculture.
MICHIGAN: Kent Co., Wyoming, 5 May
1986, E. R. Hoebeke; Grand Rapids, 6 May
1986, ERH.
NEW YORK: Albany Co., nr. Colonie,
Pine Bush, 30 June 1984, ERH and A. G.
Wheeler, Jr. Broome Co., Binghamton, 23
July 1983, ERH and AGW. Chemung Co.,
Rt. 17 N. of Wellsburg, 27 June and | Au-
gust 1982, AGW. Clinton Co., August 1983.
Columbia Co., June 1985. Dutchess Co.,
Stanfordville, October 1984. Erie Co., Ton-
awanda, 12 June 1983 and 22 June 1985,
ERH. Essex Co., May, November 1977.
Livingston Co., Dansville, June 1980.
Greene Co., East Windham, June 1984.
Monroe Co., Greece, 31 July 1982, ERH
22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and AGW. Onondaga Co., Solvay, 26 June
and 4 September 1982, 14 May 1983, ERH
and AGW. Ontario Co., Canandaigua, April
1979. Rensselaer Co., Troy, June 1973.
Schenectady Co., August 1973. Tompkins
Co., Ithaca, 20 April 1987, ERH. Ulster Co.,
May and September 1982. Wayne Co.,
Clyde, 25 June 1983, ERH and AGW.
PENNSYLVANIA: Erie Co., Erie, 11 July
1985, AGW. Lackawanna Co., Carbondale,
28 June 1985, AGW. Lebanon Co., I-81 at
junc. I-78 NW of Jonestown, 20 August 1982
and March—August 1983, AGW. Mercer Co.,
Sharon, 29 July 1987, AGW. Susquehanna
Co., Thompson, 28 June 1985, AGW.
Our collections of A. helix in Broome,
Chemung, Erie, Monroe, Onondaga, Tomp-
kins, and Wayne counties in New York; Erie,
Lackawanna, Mercer, and Susquehanna
counties in Pennsylvania; and Kent County
in Michigan were made along or near rail-
road right-of-ways. In fact, populations were
nearly always found only in a small area
adjacent to active or abandoned trunk lines.
In Lebanon Co., Pennsylvania, A. helix ap-
parently is restricted to less than a mile of
road near the junction of interstate high-
ways 78 and 81.
It seems reasonable to assume that spread
of this flightless, parthenogenetic moth in
North America is largely dependent on
commerce, especially rail traffic. Mature
larvae are known to leave low-growing hosts
and to attach to a suitable, usually higher,
substrate for pupation (Robinson 1953). A
larva could easily attach its case to a rail car
in storage and the pupa, female, eggs, or
overwintering larvae could be transported
many miles to initiate a new colony. In one
California orchard a new infestation of J.
helix was attributed to its introduction as
cases attached to a “‘small private spray rig”
that had been used in the infested area (Ar-
mitage 1953).
SEASONAL HISTORY AND HABITS
To obtain information on phenology of
this psychid in eastern North America, col-
lections were made in 1983 from a roadside
planting and from guardrails at the junction
of interstate highways 78 and 81 in Lebanon
Co., Pa. Observations were made and cases
collected on 30 March; 1, 13, 22, and 27
April; 5, 11, and 18 May; 2 and 21 June;
15 July; and 3 August. Populations at the
site were not as large as the hundreds of old
cases adhering to guardrails might suggest,
and on several sample dates the bagworms
were not easily found on vegetation; cases,
which “resist weathering to an amazing de-
gree” (Robinson 1953), probably persist for
years in protected places.
Life stages present in the population were
approximated by dissecting small numbers
of cases (usually only 5-10) on each sample
date and, for larvae, measuring widths of
head capsules. Such measurements suggest-
ed four larval instars: I, 0.28—0.30 mm wide
(n = 40); II, 0.36-0.40 mm (n = 16); III,
0.48-0.54 mm (n = 10); and IV, 0.64—0.80
mm (n = 9).
As Robinson (1953) reported for A. helix
in California, young larvae overwinter with-
in the female’s pupal skin. Larvae appar-
ently construct septa within this empty shell
so that each is enclosed in its own cell (Davis
1964). In Pennsylvania during late March—
early April, 14 occupied cases that were col-
lected on guardrails and examined in the
laboratory contained first instars. As many
as 42 and 49 larvae emerged from a single
case. Several cases were coated with clover
mites, Bryobia praetiosa Koch, and their egg
shells. When cases were brought into the
laboratory, larvae soon emerged, began to
construct their own cases from sand and soil
of old cases, and to feed on foliage, the dam-
age appearing as tiny, circular transparent
areas (Fig. 1).
On 13 April, first instars, about | mm
long and without cases, were active on
guardrails, with a few larvae in small cases
observed on nearby spotted knapweed,
Centaurea maculosa Lam. (Asteraceae).
These cases consisted merely of a few sand
grains on the body. Some knapweed leaves
VOLUME 90, NUMBER |
Figs. 1-3.
cases) feeding on black medic, Medicago lupulina, causing tiny, circular transparent areas. 2, Feeding injury by
mature larvae to leaves of common mullein, Verbascum thapsus. 3, Feeding injury by mature larvae to black
medic.
showed slight feeding symptoms similar to
those observed in the laboratory. First in-
stars were the only stage found in cases tak-
en on 22 and 27 April (n = 7, 10) (Fig. 6).
Of 5 larvae collected on 5 May, 3 were first
and 2 were second instars; by 11 May only
one first instar was present in a collection
that contained 8 second instars; and only
second instars were present on 18 May (n =
5). Cases observed during May were larger,
consisting of a white silken sac impreg-
nated with grains of sand and soil (Fig. 4a).
Robinson (1953) described these cases as
having the form of an inverted J or U. In
May damage on knapweed foliage became
more obvious.
Feeding damage to foliage by larval stages of Apterona helix. 1, Early-stage larvae (with small
Larvae feed mainly at night (Davis 1964).
A larva feeds by using silk to fasten its case
to a leaf and emerging through an opening
near the bottom of the case (see Fig. 4b). It
chews a hole in the adaxial or abaxial sur-
face, inserts its head in the opening, and
scrapes out tissue between the leaf surfaces,
making a nearly oval mine. Davis (1964)
noted that this injury closely resembles that
made by lepidopteran larvae of the genus
Coleophora (Coleophoridae). Two, three, or
more windowlike areas were observed on
some small leaves (Figs. 2, 3). According to
Robinson (1953), fecal material is expelled
through a lateral aperture in the upper or
smallest whorl of the case.
24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
-)
Tas
a
Figs. 4-5. Larval cases of Apterona helix. 4a, Case of early-stage larva, consisting of small silken case
impregnated with grains of sand and soil. 4b, Mature larval case, usually of 2! to 32 whorls; arrow indicates
large basal opening through which the larva emerges to feed; scale line = 5 mm. 5, Aggregation of larval cases
on trunk of tree sapling.
Ten cases collected on 2 June contained
third instars; all nine taken on 21 June yield-
ed fourth instars (Fig. 6). Mature larvae oc-
cupy the lower whorl (Robinson 1953, Da-
vis 1964) of the helicoid or snail-like cases
(Fig. 4b). On 15 July few active larvae were
observed on knapweed, but closed cases were
apparent and clumped on small trees (Fig.
5); several cases opened in the laboratory
were found to contain pupae. Mature larvae
also ascended guardrails for pupation, where
current-season cases could not be distin-
guished in the field from those of previous
seasons. At other localities utility poles were
used as pupation sites, with large numbers
of old cases occurring in cracks and crevices.
VOLUME 90, NUMBER |
[Zan [Fee [WAR[ APR MAY | JUN | JUL | AUG] SEP | ocr | NOV] DEC]
GUEHWItTERING 1ST ST INSTARS NN J 1ST INSTARS
SSS ~
.
2ND INSTARS
3RD INSTARS bie
4TH INSTARS
Fig. 6.
Ist instars are overwintering inside the female case.)
Females were found in several cases col-
lected on 3 August. Robinson (1953) re-
ported that yellowish-white eggs (about 25/
case) were laid in the pupal skin that re-
mains in the lower whorl and hatch in about
3 weeks; the diapausing larvae remain in-
side the case through the winter.
Seasonal history of the Pennsylvania pop-
ulation studied was nearly the same as that
reported for 4. Ae/ix in northern California
(Robinson 1953). Overwintered first-instar
larvae became active in mid-April and fed
until early to mid-July, with females ap-
pearing to oviposit in August.
Host PLANTS
Spotted knapweed, the most frequently
infested host at the study site in Lebanon
Co., Pennsylvania, was the most common
host plant observed along railroad lines in
New York and Pennsylvania. White and
yellow sweetclover, Melilotus alba Medik.
and M. officinalis Lam. (Fabaceae), also were
infested. The large population of A. helix at
Solvay, New York, severely damaged these
1ST INSTARS
No" OVERWINTERING 1ST INSTARS NS
Inferred seasonal history of Apterona helix in Pennsylvania. (Note: stippling indicates period when
plants and several unidentified hosts. Species
that showed occasional injury or moderate
to heavy damage at only one or a few sites
included green foxtail, Setaria viridis (L.)
Beauv. (Poaceae); mugwort, Artemisia vul-
garis L. (Asteraceae); black mustard, Bras-
sica nigra (L.) Koch, and peppergrass,
Lepidium virginicum L. (Brassicaceae);
blueweed, Echium vulgare L. (Boraginace-
ae); common mullein, Verbascum thapsus
L., and yellow toadflax, Linaria vulgaris
Mill. (Scrophulariaceae); common evening-
primrose, Oenothera biennis L. (Onagra-
ceae); cinquefoil, Potentilla sp. (Rosaceae);
buckhorn plantain, Plantago lanceolata L.,
and broadleaf plantain, P. major L. (Plan-
taginaceae); knotweed, Po/ygonum sp. (Po-
lygonaceae); and crownvetch, Coronilla
varia L., black medic, Medicago lupulina L.,
and alsike clover, Trifolium hybridum L.
(Fabaceae).
Crownvetch was the dominant plant
species at the Pennsylvania study site, but
only slight feeding on leaflets of a few plants
was observed. Along the railroad in New
26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
York common plants that generally were
avoided were hedge bindweed, Calystegia
sepium (L.) R. Br. (Convolvulaceae); lambs-
quarters, Chenopodium album L. (Cheno-
podiaceae); chicory, Cichorium intybus L.,
and horseweed, Conyza canadensis (L.)
Cronq. (Asteraceae); and Queen Anne’s-lace,
Daucus carota L. subsp. carota (Apiaceae).
Apterona helix, although a polyphagous
insect known from various wild and culti-
vated plants (Robinson 1953, Davis 1964),
fed mainly on low-growing herbs in New
York and Pennsylvania. In California, how-
ever, Robinson (1953) noted that “‘foliage
of shrubs and trees were freely attacked later
in the season.”’ We also observed some feed-
ing on shrubs and trees at the study site and
along railroad tracks, but these plants served
primarily as sites for pupation (Fig. 5).
RECOGNITION FEATURES
The most characteristic morphological
attribute of this small psychid moth is the
larval case (Figs. 4a, 4b); it alone will enable
recognition of this introduced bagworm
moth in North America. The small spiraled
case measures approximately 3—5 mm in
diameter and 4-5 mm in depth. The mature
larval case typically has 2’ to 3% whorls
and is constructed of silk overlain with mi-
nute earthen particles. There are three open-
ings present in the case: a small apical open-
ing, a large basal opening through which the
larva is able to crawl and feed, and a large
slitlike opening in the uppermost whorl
through which the shrivelled female re-
portedly emerges after oviposition (Davis
1964).
Males of A. helix are not known. The par-
thenogenetic females are highly specialized,
larviform, wingless or with reduced wings,
with a small head with or without small
antennal rudiments, with small pigment
spots instead of eyes, and with leg rudiments
without claws (Kozhanchikov 1956). The
females never leave the spiral case.
ACKNOWLEDGMENTS
We thank Carolyn Klass (Cornell Uni-
versity) for allowing us to use records of A.
helix submitted to the Insect and Plant Dis-
ease Diagnostic Laboratory, Gary Miller
(Auburn University) for illustrating the lar-
val cases in Figs. 4a and 4b, Alan Snook
(Pennsylvania Dept. of Agriculture) for dis-
secting cases and measuring head capsule
widths of larvae, Maureen E. Carter (CU)
for assistance in the field, James Stimmel
(PDA) for helping make field collections and
for reproducing Figs. 1-3 and 5, and Karl
Valley (PDA) for reviewing an early draft
of the manuscript.
LITERATURE CITED
Adamski, D. 1984. An unusual oviposition site for
Amphipyra tragopogonis (L.) (Noctuidae). J. Lep-
id. Soc. 38: 317-318.
Armitage, H. M. 1953. Current insect notes. Calif.
Dep. Agric. Bull. 42: 39-41.
Davis, D. R. 1964. Bagworm moths of the Western
Hemisphere (Lepidoptera: Psychidae). U.S. Natl.
Mus. Bull. 244. 233 pp.
1983. Psychidae (Tineoidea), pp. 7-8. Jn R.
W. Hodges et al., eds., Check List of the Lepi-
doptera of America North of Mexico. E. W. Clas-
sey Ltd. & Wedge Entomol. Res. Found.
1987. Psychidae (Tineoidea), pp. 366-369.
In F. W. Stehr, ed., Immature Insects. Kendall/
Hunt Publ. Co., Dubuque, Iowa.
Eichlin, T. 1985. Imported bagworm, Apterona cren-
ulella. Calif. Plant Pest Dis. Rep. 4(4): 115.
Every, R. W. 1970. A psychid moth (4pterona cren-
ulella). U.S. Dep. Agric. Coop. Econ. Insect Rep.
20(40): 700.
Gittins, A. R. 1958. A psychid moth (Apterona cren-
ulella). U.S. Dep. Agric. Coop. Econ. Insect Rep.
8(28): 607.
Keifer, H. H. 1940. Systematic entomology, pp. 241-
245. In D. B. Mackie, Chief. Division of Plant
Industry, Bureau of Entomology and Plant Quar-
antine. Calif. Dep. Agric. Bull. 29.
1947. Systematic entomology, pp. 168-173.
In H. M. Armitage, Chief. Division of Plant In-
dustry, Bureau of Entomology and Plant Quar-
antine. 28th Ann. Rep. Per. Ending Dec. 31, 1947.
Calif. Dep. Agric. Bull. 36.
Klass, C. 1983. Comments from diagnosticians.
Entomology, pp. 3-4. /n Insect and Plant Disease
VOLUME 90, NUMBER |
Diagnostic Laboratory Annual Report, 1983. Coop.
Ext. Cornell Univ., Ithaca, N.Y.
Knowlton, G. F. 1961. A psychid (Apterona crenu-
lella). U.S. Dep. Agric. Coop. Econ. Insect Rep.
11(27): 598.
1966. A psychid (Apterona crenulella). U.S.
Dep. Agric. Coop. Econ. Insect Rep. 16(22): 479,
482.
Knowlton, G. F. and D. S. Parrish. 1965. A psychid
(Apterona crenulella). U.S. Dep. Agric. Coop. Econ.
Insect Rep. 15(28): 744.
Knowlton, G. F. and R. S. Roberts. 1968. Range
entomology, a subject of increasing importance.
Utah State Univ. Entomol. Mimeo Ser. 133. 7 pp.
Kozhanchikov. 1956. Fauna of the U.S.S.R., Lepi-
doptera, Vol. III, No. 2 (Psychidae). Zool. Inst.
Akad. Nauk SSSR (n.s.), No. 62 (Translated from
27
Russian, 1969, Israel Program for Scientific Trans-
lations, Jerusalem. 525 pp.).
Marshall. 1970. A psychid moth (Aplerona crenulel-
la). U.S. Dep. Agric. Coop. Econ. Insect Rep.
20(25): 407.
Robinson, D. 1953. Garden bagworm, Apferona
crenulella (-helix) in Nevada and Placer counties,
California. Calif. Dep. Agric. Bull. 42: 25-33.
Strand, E. 1912. Psychidae, pp. 353-370. Jn A. Seitz,
Gross-Schmetterlinge der Erde, Fauna palaearct.
Bd. 2. A. Kernen, Stuttgart.
Suomi, D. 1986. Have you seen this bug?, pp. 4-5.
In Co. Agent Entomol. Newsl. No. 2. Wash. State
Univ., Pullman.
Tibbetts, T. and G. F. Knowlton. 1952. Apterona
crenulella in Utah (Lep.: Psychidae). Entomol.
News 63: 211.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 28-36
A REVISION OF THE GENUS MESTOCHARIS AND A REVIEW
OF THE GENUS GRAHAMIA (HYMENOPTERA, EULOPHIDAE)
CHRISTER HANSSON
Department of Systematic Zoology, Lund University, Helgonavagen 3, S-223 62 Lund,
Sweden.
Abstract.—The Nearctic and Palearctic species of the genus Mestocharis Forster are
revised. Three species are recognized, one exclusively Nearctic (M. tropicalis), one exclu-
sively Palearctic (M. maculata) and one found in both regions (M/Z. bimacularis). Mesto-
charis nearctica Yoshimoto is a new junior synonym of M. bimacularis. Hosts are known
only for MM. bimacularis, a gregarious endoparasitoid in eggs of larger Dytiscidae (Co-
leoptera). The two known species of Grahamia are reviewed and characters are presented
to facilitate their separation. Both species are recorded as new to the Nearctic Region.
The biology is known only for G. clinius, a parthenogenetic endoparasite of Haplodiplosis
equestris (Diptera, Cecidomyiidae) in Europe.
Mestocharis has been treated in two rath-
er recent papers, one dealing with the Eu-
ropean (Boucek et al. 1963) and the other
with the North American species (Yoshi-
moto 1976). Boucek et al. recognized two
species (M. bimacularis (Dalman) and ™.
maculata (Forster)) and established two new
combinations and three new synonyms.
They dealt also with the intraspecific vari-
ation of taxonomically important charac-
ters and presented a key. Yoshimoto de-
scribed two new species (V/. nearctica and
M. tropicalis), compared them to the Eu-
ropean species, and presented a key to the
Nearctic species. The material I have at my
disposal indicates that the intraspecific vari-
ation described in those papers was under-
estimated. The consequences of this under-
estimation are that some characters used for
separating the species are unsafe and that
one of the species is invalid. The biology
is known only for M. bimacularis (Jackson
1958, 1960, 1964), an endoparasite in eggs
of larger Dytiscidae (Coleoptera). Two other
Nearctic species were described as Mesto-
charis (M. wilderi Howard, M. williamsoni
Girault), but both have been transferred to
Pediobius Walker (Burks 1958: 68-69).
The genus Grahamia was erected to in-
clude Entedon clinius Walker and Graham-
ia tatrica Erdés (Erdés 1966). Later, Boucek
and Askew (1968) catalogued the Palearctic
Eulophidae and added new distribution rec-
ords and corrected some previous host rec-
ords for the genus. Hansson (1985) specu-
lated that the two species might be the same
because of high intraspecific variation in the
length of flagellar segments, the only char-
acter separating the two species, but made
no definite decision regarding the validity
of the two species. Because I have had access
to a fairly large sample of these species from
Europe and from North America, I con-
clude that the two species are valid. A key
is presented to facilitate their identification.
The host is known only for G. clinius, a
parthenogenetic endoparasite of Haplodi-
plosis equestris (Diptera, Cecidomyiidae) 1n
Europe.
Morphological terms used are explained
VOLUME 90, NUMBER 1
in Hansson (1985), the exception being
POO, the distance between posterior edge
of hind ocelli and occipital margin. Abbre-
viations of museums and private collections
used in the text were as follows: BMNH =
British Museum (Natural History), London,
England; CH = collection of the author;
CNC = Canadian National Collections,
Ottawa, Canada; DAFZ = Department
of Agriculture and Forest Zoology, Hel-
sinki, Finland; INHS = Illinois Natural His-
tory Survey, Champaign, Illinois, USA;
HNHM = Hungarian Natural History Mu-
seum, Budapest, Hungary; LUZM = Lund
University Zoological Museum, Lund, Swe-
den; SMNH = Swedish Museum of Natural
History, Stockholm, Sweden; USNM = Na-
tional Museum of Natural History, Wash-
ington, D.C., USA.
Genus Mestocharis Forster
Mestocharis Forster, 1878: 50. Type-species:
Entedon bimacularis Dalman, 1820: 181
(= Mestocharis cyclospila Forster, 1878:
50), by original designation.
Diagnosis.—Species of Mestocharis are
distinguished from other genera of Ente-
dontinae by the following combination of
characters: both sexes with two small and
discoid anelli; antennal scrobes adjoining
on horizontal line of frontal fork; mandibles
tridentate; transverse pronotal carina ab-
sent; postmarginal vein about as long as
stigmal vein; anterior part of propodeum
with two conspicuous indentations lateral-
ly; anteromedian part of propodeum with a
large and more or less triangular projection.
Remarks.—The monophyly of Mesto-
charis is shown through the following syn-
apomorphies: 1) anterior part of propo-
deum with two conspicuous indentations
laterally; 2) anteromedian part of propo-
deum with a large triangular projection.
KEY TO THE HOLARCTIC SPECIES OF
MESTOCHARIS
bho
2. Costal cell with a complete row of setae on
underside; 2nd tergite smooth and shiny
ne M. maculata (Forster)
— Costal cell bare; 2nd tergite microreticulate (Fig.
1) hs ; 3
3. Seventh tergite 1.2-2.0= as long as width of
base of same tergite (Fig. 8); scutellum smooth
and shiny medially along its entire length
M. tropicalis Yoshimoto
— Seventh tergite 0.4-1.0= as long as width of
base; scutellum with weaker reticulation me-
dially, but not smooth (Figs. 1, 6)
M. bimacularis (Dalman)
4. Costal cell with a row of setae on underside,
antenna mainly testaceous =
M. maculata (Forster)
— Costal cell bare, apical part of scape and the
antenna beyond usually infuscate
M. bimacularis (Dalman)
Mestocharis bimacularis (Dalman)
Figs. 1-7
Entedon bimacularis Dalman, 1820: 181.
Entedon arisha Walker, 1839: 121-122. Syn.
Boucek et al., 1963: 5.
Mestocharis cyclospila Forster, 1878: 50.
Syn. Bouéek et al., 1963: 5.
Mestocharis militaris Rimsky-Korsakov,
1933: 232, 244-245. Syn. Bouéek et al.,
1963: 5.
Mestocharis nearctica Yoshimoto,
756-757. New synonymy.
Mestocharis bimacularis (Dalman), Bouéek
et al., 1963: 5.
Diagnosis.—Costal cell bare; second ter-
gite microreticulate; seventh tergite 0.4 to
1.0 as long as wide at base; scutellum re-
ticulate all over; femora usually more or less
infuscate.
Description.— Female: Entire antenna
dark, except scape more or less pale at base.
Face and clypeus golden-green, frons com-
pletely golden-purple or golden-green below
frontal fork. Vertex, mesoscutum and scu-
tellum golden, coppery, golden-green or
golden-red. Coxae dark and metallic, major
part of femora dark and more or less me-
tallic, tibiae varying from brownish to yel-
low, base of tibiae occasionally dark, fore-
tarsi dark, mid- and hindtarsi with basal
1976:
Figs. 1-8. Mestocharis spp. 1-7, M. bimacularis. 1, Habitus, female. 2, Front view of head, female. 3, Same,
male. 4, Female antenna. 5, Male antenna. 6, Female gaster. 7, Male petiolus and gaster. 8, M. tropicalis, female
gaster.
30
VOLUME 90, NUMBER 1
three segments pale and with 4th segment
dark. Forewing with a large fuscous spot just
below stigmal vein, usually also with two
much fainter spots below that spot (see re-
marks for comments on these spots). Pro-
podeum and gaster golden-green. Length of
body: 1.6-2.6 mm. Ratios height of eye/
malar space/width of mouth opening = 3.1/
1.0/2.0. Malar space 2.5-3.0x as wide as
width of scape. Frons below fork with very
weak reticulation and almost smooth, part
between toruli and antennal scrobes raised
In a conspicuous, blunt elevation. Frons
above fork smooth and shiny in lower part,
in upper part reticulate with low and narrow
septa, and with small meshes. Horizontal
line of frontal fork shaped like a V. Inner
orbit of compound eye with | to 2 rows of
setae. Vertex reticulate with very low and
very narrow septa inside ocellar triangle,
outside smooth and shiny. Ratios POL/
OOL/POO = 6.0/3.1/1.0. Occipital margin
with a +strong carina behind ocellar tri-
angle. Ratio width of head/width of thorax
across shoulders = 1.1. Mesoscutum and
scutellum reticulate with high and wide sep-
ta, median part of scutellum with lower sep-
ta. Forewing with speculum closed, base of
submarginal vein with a short row of setae
on underside. Propodeum with a strong and
complete median carina, plicae complete or
missing in anterior part of propodeum, pro-
podeal surface rather strongly reticulate, also
with irregular carinae. Propodeal callus with
6-9 and lateral part of propodeum inside
spiracular sulcus with 3-10 setae. Petiolar
foramen rounded. Petiolus transverse, me-
dian part with considerably raised carinae,
lateral corners protruding and sharp. Shape
of gaster varying from ovate (Fig. 6) to lan-
ceolate (Fig. 1). First tergite smooth and
shiny, tergites 2-5 microreticulate in ante-
rior 3/4, tergites 6-7 with stronger reticu-
lation. Mean ratio length of thorax + pro-
podeum/length of gaster = 0.82 + 0.104,
n= 10.
Male: Scape pale with apical part dark,
pedicel and two first flagellar segments oc-
31
casionally paler than remaining flagellum,
which is dark. Face and frons golden-green
or -blue, vertex golden-red. Thorax golden-
green, -blue or -red. Coxae dark, remainder
of legs yellowish, hindfemur usually dark at
base. Forewing immaculate. Metallic col-
oration much brighter than in female. Length
of body: 1.3-1.8 mm. Frons above fork and
vertex smooth and shiny. Ratios height of
eye/malar space/width of mouth opening =
4.6/1.0/2.6. Malar space as wide as width
of scape. Ratio width of head/width of tho-
rax across shoulders = 1.3. Propodeal callus
with 4-7 and lateral part of propodeum in-
side spiracular sulcus with 2-7 setae. Petio-
lus like in female, but varying in shape from
transverse to as long as wide. Mean ratio
length of thorax + propodeum/length of
gaster = 1.06 + 0.087 n = 10.
Remarks.— The dark spots on the female
forewings are apparently characters that de-
velop with age. Newly emerged females have
immaculate forewings, while the same fe-
males 8-13 days later have clearly visible
spots. In females that have lived 9-10
months the spots are exceptionally dark
(Jackson 1964).
The shape of the female gaster is variable
in this species, varying from ovate (Fig. 6)
to lanceolate (Fig. 1). This character (ac-
tually, the shape of last tergite) was used in
Bouéek et al. (1963) to separate bimacularis
from maculata: the last tergite was about
halfas long as its basal width in bimacularis
and as long or longer as its basal width in
maculata. In Swedish specimens of bimacu-
/aris last tergite varies from slightly less than
half as long to as long as its basal width.
Yoshimoto (1976) separated nearctica
from bimacularis through several characters
presented in a table. However, these char-
acters are either so variable intraspecifically
that they have no taxonomic value or are
misinterpreted. The propodeal carinae are
variable characters. The median carina is
usually strong, wide and complete, but in a
few specimens it is narrower. The diagonal
carina extending from the triangular pro-
32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
jection to the sides of the petiolar foramen
varies from strong and complete to com-
pletely missing. The hind margin of the first
tergite is usually sinuate while the hind mar-
gin of the second tergite varies from almost
straight to sinuate. The characters of the
male antenna of nearctica must have been
misinterpreted by Yoshimoto. A male para-
type of nearctica (Can., Ont., Ottawa
19.vu1.1939, O. Peck) that I saw showed the
following characters: the scape is yellow with
apical fourth infuscate and the first flagellar
segment is about 1.8 x as long as wide. The
measurements of the fifth flagellar segment
are correct (1.7 x as long as wide), but this
is also about the same size usually encoun-
tered in bimacularis. Consequently I regard
nearctica as a synonym of bimacularis.
Material examined.—Two paratypes of
M. nearctica (1 2 and 1| 4, in CNC); Canada:
British Columbia 2 2 (CNC, LUZM): New-
foundland | ¢ (BMNH): Nova Scotia | 2
(CH); Ontario 1 2 (LUZM). Finland: 3 ¢
(DAFZ). Sweden: 40 ¢ 24 6 (CH, LUZM).
USA: Michigan | ¢é (USNM). Lectotype 2
E. bimacularis (not seen) in SMNH.
Hosts.—This species is a solitary or gre-
garious endoparasitoid in eggs of Dytiscidae
(Coleoptera). The size of the egg restricts
the number of wasps that can develop suc-
cessfully, in larger eggs, e.g. those of Dytis-
cus marginalis, up to 12 wasps have been
reared, while in smaller eggs, e.g. those of
Ilybius ater, only one wasp developed (Jack-
son 1964). Imagines of /. bimacularis are
most frequently encountered in pond- or
marsh vegetation.
Distribution.— Widespread in Europe
(Boucek and Askew 1968), in the Nearctic
Region this species is recorded from both
Canada (Alberta, Manitoba, Ontario and
Quebec (Yoshimoto 1976); British Colum-
bia, Newfoundland and Nova Scotia) and
the United States (Michigan).
Mestocharis maculata (Forster)
Eulophus maculatus Forster, 1841: 41-42.
Pleurotropis maculata (Forster), Erdos,
1956: 38-39.
Mestocharis maculata (Forster), Bouéek et
al., 1963: 9.
Diagnosis.—Costal cell with a complete
row of setae on underside: second tergite
smooth and shiny; femora more or less pale
testaceous. In other characters M. maculata
is very similar to M. bimacularis, and the
description of bimacularis is otherwise ap-
plicable to maculata.
Material examined.—USSR: Moldavian
SSR 1 4; Yugoslavia: Beograd | 2. Both
specimens are from the Bouéek collection.
Lectotype 2 E. maculatus (not seen) in the
Forster collection in Vienna.
Distribution.—Europe: Czechoslovakia,
Germany, Hungary and USSR (Bouéek and
Askew 1968).
Mestocharis tropicalis Yoshimoto
Fig. 8
Mestocharis tropicalis Yoshimoto,
Ti
Diagnosis.—Seventh tergite 1.2 to 2.0x
as long as width of base of same tergite;
scutellum smooth and shiny medially along
its entire length; costal cell bare; femora pale.
Males are unknown for this species.
Remarks.—This species is similar to its
two congeners but can be separated from
them using the characters given in the key.
The 7th tergite shows a great deal of intra-
specific variation. I have seen only two spec-
imens, both females, one from Florida
(paratype) and one from Illinois. The spec-
imen from Florida had ratio length/basal
width of 7th tergite = 1.2, and the specimen
from Illinois = 2.0. There is a great gap
between these two measurements, but when
more material turns up this gap may be filled.
The shape of the gaster 1s more lanceolate
than in M. bimacularis, which leaves no
doubt that tropicalis is a valid species. Like
M. bimacularis, tropicalis also has a fuscous
spot just below the stigmal vein, but not the
two fainter spots present in bimacularis.
Material examined.— Paratype 2 (CNC);
1 2 from USA, Illinois, Champaign Co.
(INHS). Holotype 2 (not seen) in CNC.
1976:
VOLUME 90, NUMBER 1
Distribution.—The United States (Flori-
da, Illinois).
Genus Grahamia Erdos
Grahamia Erdés, 1966: 406. Type-species:
Entedon clinius Walker, 1839: 90, by
original designation.
Diagnosis. —Species of Grahamia are dis-
tinguished from other genera of Entedon-
tinae by the following combination of char-
acters: all flagellar segments free; antenna
with only one small and discoid anellus;
mandibles tridentate; pronotal collar with-
out transverse carina; costal cell narrow;
postmarginal vein about 2 x as long as stig-
mal vein.
Remarks.—The monophyly of Grahamia
is shown through the single discoid anellus,
a synapomorphy for the genus. Two things
might argue against the value of this char-
acter. First, it is clearly a reduction (the ple-
siomorphic character state is three anelll,
present in the closely related genus Chry-
socharis Forster). Reductions are some-
times regarded as poor apomorphic char-
acter states. Secondly, this character state
(one discoid anellus) occurs in other closely
related genera (some Chrysonotomyia Ash-
mead and Closterocerus Westwood). Never-
theless, I prefer to keep Grahamiaa separate
genus from Chrysocharis, the genus to which
Grahamia shows closest morphological af-
finity. Among the species-groups of Chry-
socharis (sensu Hansson 1985), Grahamia
comes closest to the mediana-group. Gra-
hamia has, however, some characters that
disagree with this assessment: its single dis-
coid anellus of the antenna, all flagellar seg-
ments free, and gallmidges as hosts. Chry-
socharis has three anelli, two apical flagellar
segments fused in species of the mediana-
group, and leafminers as hosts.
KEY TO THE SPECIES OF
GRAHAMIA (FEMALES)
1. First segment of flagellum about 2* and 4th
segment about 1.5 = as long as wide (Fig. 12);
malar space about 1.5= as wide as width of
scape; metallic coloration of body compara-
tively dull .. ..G. clinius (Walker)
— First segment of flagellum about 1.5 * and 4th
segment 1 x as long as wide (Fig. 10); malar
space narrower (as wide as width of scape);
metallic coloration of body brighter
G. tatrica Erdés
Grahamia clinius (Walker)
Figs. 11-12
Entedon clinius Walker, 1839: 90.
Chrysocharis clinius (Walker), Graham,
1959: 196.
“Chrysocharis”’ clinius (Walker), Graham,
1963: 204.
Grahamia clinius (Walker), Erdés, 1966:
407.
Diagnosis. — Flagellar segment | about 2 x
and 4 about 1.5= as long as wide; malar
space 1.5 as wide as width of scape; me-
tallic coloration of body dull; larger species
(1.6-2.1 mm).
Description.— Female: Entire antenna
dark, scape occasionally paler in basal half.
Face and clypeus golden-green. Frons and
vertex metallic purple, below fork some-
times golden-purple. Mesoscutum golden-
green and scutellum metallic purple. Coxae
dark brown and weakly metallic. Femora
predominantly dark, fore- and midtibiae
varying from pale to infuscate, hindtibia
predominantly pale, tarsi + infuscate, 4th
segment dark. Wings hyaline or weakly in-
fuscate. Propodeum and gaster golden-green.
Length of body: 1.6-2.1 mm. Ratios height
of eye/malar space/width of mouth open-
ing = 3.9/1.0/2.8. Malar space 1.5 x as wide
as width of scape. Frons below fork reticulate
with low and rather narrow septa, meshes
small, above fork shiny and almost smooth.
Horizontal line of frontal fork almost
straight. Inner orbit of compound eye with
one row of setae. Vertex reticulate with very
low and very narrow septa. Ratios POL/
OOL/POO = 3.3/2.2/1.0. Occipital margin
with a carina behind ocellar triangle. Ratio
width of head/width of thorax across shoul-
ders = 1.3. Mesoscutum reticulate with low
and rather narrow septa, meshes small. Scu-
tellum reticulate with low to very low and
narrow to very narrow septa, 1.e. with finer
34
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
| 10
Nes)
Ne:
ii
Sy
Figs. 9-12. 9-10, Grahamia tatrica, female. 9. Habitus. 10, Antenna. 11-12, G. clinius, female. 11, Front
view of head. 12, Antenna.
VOLUME 90, NUMBER 1
reticulation than mesoscutum. Anterome-
dian part of propodeum with a weak tri-
angular fovea, propodeal surface +reticu-
late, with or without a weak median carina.
Propodeal callus with two setae. Petiolar fo-
ramen rounded. Petiolus small and trans-
verse. Gaster elongate, ratio length of tho-
rax + propodeum/length of gaster = 0.67-
0.87.
Remarks.—The species Tetrastichus
idothea Walker, 1844: 409, was regarded as
a possible synonym of G. clinius by Graham
(1961: 62).
Material examined.—Lectotype EF. cli-
nius 2 (BMNH Type No. 5.2025); BRD: 1
2 ex Hapl. equestris (USNM). Canada: Brit-
ish Columbia 3 2? (CNC, LUZM). Sweden:
1 2 (CH).
Host.—Grahamia clinius is known as an
endoparasitoid in larvae of Haplodiplosis
equestris (Diptera, Cecidomyiidae) (Baier,
1963/64, as Chrysocharis seiuncta). The sex
ratio of reared G. clinius (2:6 = 100:1) (Baier
1963/64) suggests that the species propa-
gates parthenogenetically.
Distribution. — Grahamia clinius is wide-
spread in Europe (Boucek and Askew 1968),
and now for the first time recorded from the
Nearctic Region (Canada, British Colum-
bia).
Grahamia tatrica Erdos
Figs. 9-10
Grahamia tatrica Erdés, 1966: 407.
Chrysocharis atripes Szelenyi, 1979: 178.
Syn. Hansson, 1985: 97.
Diagnosis.—Flagellar segment | about
1.5 and 4 about | x as long as wide; malar
space as wide as width of scape; metallic
coloration of body brighter; smaller species
(1.3-1.5 mm).
Remarks.—Apart from the distinguish-
ing characters G. tatrica is very similar to
G. clinius and the description of c/inius is
applicable to tatrica.
Material examined. —Paratypes 2 2 G. ta-
trica (HNHM Nos. 6061 & 6062); Canada:
Nova Scotia | ? (LUZM),; Finland: 10 2 (CH,
35
DAFZ); Sweden: 2 2 (CH); USA: Michigan
1 @ (USNM), West Virginia | @ (LUZM).
Holotype 2 (not seen) in HNHM.
Distribution.—Grahamia tatrica 1s re-
corded from Europe (Finland, Hungary (Er-
dds 1966), Sweden, Switzerland (Erdés
1966)), Canada (Nova Scotia) and the
United States (Michigan, West Virginia).
This species was previously not recorded
from the Nearctic.
ACKNOWLEDGMENTS
I am grateful to Z. Boucek (BMNH),
G.A.P. Gibson (CNC), S. L. Heydon (INHS),
L. Huggert (same address as author), M. Ko-
ponen (DAFZ), J.S. Noyes (BMNH), J. Papp
(HNHM), M. E. Schauff (USNM), and C.
M. Yoshimoto (CNC) for loan of type- and
additional material. My work with this pa-
per was partly financed through a grant from
the O. Larsén Foundation.
LITERATURE CITED
Baier, M. 1963/64. Zur Biologie und Gradologie der
Sattelmiicke Haplodiplosis equestris Wagner (Dip-
tera, Cecidomyiidae). Z. Angew. Entomol. 53: 216-
273:
Bouéek, Z. and R. R. Askew. 1968. Palearctic Eu-
lophidae (excl. Tetrastichinae). Index of Ento-
mophagous Insects. Le Francois, Paris. 254 pp.
Boucéek, Z., M. W. R. de V. Graham, and G. J. Kerrich.
1963. A revision of the European species of the
genus Mestocharis Forster (Hym., Chalcidoidea,
Eulophidae). Entomologist 96: 4-9.
Burks, B. D. 1958. Chalcidoidea, pp. 62-84. Jn
Krombein et al., eds., Hymenoptera of America
North of Mexico: Synoptic Catalog Agric. Mono-
gr. 2. Suppl. 1.
Dalman, J. W. 1820. Foérs6k till uppstallning af in-
sectenfamiljen Pteromalini, i synnerhed med af-
seende pa de i Swerige funne arter. Tab. 2. K.
Svenska VetenskAkad. Handl. 41: 123-174, 340-
385.
Erdos, J. 1956. Additamente ad cognitionem faunae
Chalcidoidarum in Hungaria et regionibus finiti-
mis. VI. 19. Eulophidae. Fol. Entomol. Hung. (s.n.)
9: 1-64.
1966. Nonnulae Eulophidae novae Hunga-
ricae (Hymenoptera, Chalcidoidea). Ann. Hist. Nat.
Mus. Natl. Hung. Zool. 58: 395-420.
Forster, A. 1841. Beitrage zur Monographie der
Pteromalinen Nees. Aachen. 46 pp.
1878. Kleine Monographien parasitischer
36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Hymenopteren. Verh. Naturh. Ver. Preuss. Rhein.
35: 42-82.
Graham, M. W. R. de V. 1959. Keys to the British
genera and species of Elachertinae, Eulophinae,
Entedontinae and Euderinae (Hym., Chalcidoi-
dea). Trans. Soc. Br. Entomol. 13: 169-204.
1961. The genus Aprostocetus Westwood,
sensu lato (Hym., Eulophidae); notes on the syn-
onymy of European species. Entomol. Mon. Mag.
97: 34-64.
1963. Additions and corrections to the Brit-
ish list of Eulophidae (Hym., Chalcidoidea), with
descriptions of some new species. Trans. Soc. Br.
Entomol. 15: 167-275.
Hansson, C. 1985. Taxonomy and biology of the
Palearctic species of Chrysocharis Forster, 1856
(Hymenoptera: Eulophidae). Entomol. Scand.
Suppl. 26: 1-130.
Jackson, D. J. 1958. A further note on a Chrysocharis
(Hym. Eulophidae) parasitizing the eggs of Dytis-
cus marginalis L. and a comparison of its larva
with that of Caraphractus cinctus Walker (Hym.
Mymaridae). J. Soc. Br. Entomol. 6: 15-22.
1960. Revised determination of a Eulophid
(Hym., Chalcidoidea) bred from eggs of Dytiscus
marginalis L. Entomologist 93: 181.
1964. Observations on the life-history of
Mestocharis bimacularis (Dalman) (Hym. Euloph-
idae), a parasitoid of the eggs of Dytiscidae. Opusc.
Entomol. 29: 81-97.
Rimsky-Korsakov, M. N. 1933. Methoden zur Un-
tersucherung von Wasserhymenopteren. Handb.
Biol. ArbMeth. 9: 227-258.
Szelenyi, G. 1979. Four new species of Chrysocharis
Forster, 1856 (Hymenoptera: Chalcidoidea: Eu-
lophidae). Acta Zool. Acad. Sci. Hung. 25(1-2):
177-181.
Walker, F. 1839. Monographia Chalciditum. I. Lon-
don. 333 pp.
1844. On the species of Chalcidites inhab-
iting the Arctic Region. Ann. Mag. Nat. Hist. 14:
331-342, 407-410.
Yoshimoto, C. M. 1976. Synopsis of the genus Mes-
tocharis Forster in America north of Mexico
(Chalcidoidea: Eulophidae). Can. Entomol. 108:
755-758.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 37-43
GALL FORMATION BY THE CAPITULUM-INFESTING FRUIT FLY,
TEPHRITIS STIGMATICA (DIPTERA: TEPHRITIDAE)
RICHARD D. GOEDEN
Department of Entomology, University of California, Riverside, California 92521.
Abstract.— Known heretofore only as a capitulum-infesting species, Tephritis stigmatica
(Coquillett) also is reported here to develop in galls formed on branches and stems of
Senecio douglasii (deCandolle) (Asteraceae) in southern California. Two generations of
galls occur each year on S. douglasii, with immature larvae in juvenile, F, galls constituting
at least part of the overwintering population. One or more larvae feed gregariously in
interconnected, central longitudinal, short-branched feeding tunnels in the expanded pith.
As many as six individuals, including five puparia, were found in one F, gall. Florets
(including ovaries) were fed upon in ungalled capitula and F, and F, galls were formed
on branches and stems of S. douglasii at some locations, but gall-formation is the much
less common and less widespread mode of development. Galls are described and pictured.
Tephritis stigmatica (Coquillett) is the only
tephritid among the 17 species in this genus
currently described from North America
north of Mexico for which the life history
and behavior have been reported in any de-
tail (Foote 1960a, Tauber and Toschi 1965,
Foote and Blanc 1979). Tauber and Toschi
(1965) studied this species reared from ca-
pitula of Senecio integerrimus Nuttall (As-
teraceae), acommon perennial shrub in cen-
tral- and northeastern-montane California
(Munz and Keck 1959, Munz 1968). In
1981, I first reared what appeared to be 7.
stigmatica or an undescribed congener from
galls on branches and stems of Senecio
douglasii (deCandolle), another common
perennial shrub in the same genus, but
widely distributed below ca. 1500 m in Cal-
ifornia (Munz and Keck 1959, Munz 1968).
The capitula of S. douglasii also are widely
infested by 7. stigmatica (Foote and Blanc
1963, Tauber and Toschi 1965, Wasbauer
1972).
Tephritis dilacerata Loew is fairly well
known from studies by Berube (1978) and
Shorthouse (1980), but this European species
is an obligate gall former in capitula and
stems of its host, unlike the unique host-
plant relationship reported herein. Several
other species of Tephritis are known to form
capitulum and stem galls; others to infest
capitula without forming galls, but none to
do both (Freidberg 1984). The purpose of
this report is to describe gall formation by
T. stigmatica as a unique alternative mode
of development for this tephritid heretofore
known only from capitula.
MATERIALS AND METHODS
Galls on branches and stems of S. doug-
lasii from which adults recently had emerged
and galls containing larvae or puparia were
sampled at two locations in southern Cali-
fornia south of Lamont Peak at Spanish
Needle Creek, Sequoia Nat. Forest, Kern
Co., on 24 VII 1984, 7 VIII 1984, and 3 III
1987; and 2 km south of Pearblossom, San
Bernardino Co., on 22 IV 1985 and 12 I
1987.
Fully-grown larvae and puparia extracted
38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
from some of these galls were placed indi-
vidually in glass shell vials with perforated
plastic caps and held for adult emergence at
room temperature in a covered bell jar at
near-saturation R.H. The external mor-
phology of adults reared from these and sev-
eral other collections of galls were compared
at magnifications of up to 50 to 7. stig-
matica adults reared from capitula of S.
douglasii and other Asteraceae sampled
throughout southern and central California
during 1980-86.
All host-plant identifications were made
by or confirmed by A. C. Saunders, Curator
of the Herbarium of the University of Cal-
ifornia, Riverside. The plant nomenclature
used is that of Munz and Keck (1959) and
Munz (1968, 1974); the insect nomencla-
ture, that of Foote (1959, 1960a).
RESULTS
Taxonomy.—The 26 4 and 32 2 reared
from puparia dissected from branch and
stem galls (18 4 and 29 2 were reared from
the first Pearblossom sample) showed no
consistent differences compared to adults of
T. stigmatica reared from capitula. The lar-
vae feed on the florets (including ovaries)
but do not gall the capitula of S. douglasii
(personal observation). Other than body size,
body parts examined, measurements taken,
and ratios calculated in this study (Table 1)
were the same as those used by Quisenberry
(1951) of Nearctic Tephritis, including T.
stigmatica, and by Foote (1959) in his di-
agnosis of the most recently described
species of Tephritis.
Foote and Blanc (1963, p. 72) termed 7.
stigmatica “the largest of the California Te-
phritis species.” T. stigmatica adults reared
from branch and stem galls on S. douglasii
were larger on the average than adults reared
from capitula of both S. douglasii and S.
integerrimus (Table 1). The size of adults
reared from capitula of these and other As-
teraceae also varied considerably (Table 1).
This probably reflected the different nutri-
tional value of the capitula of varying size
and maturity (S. N. Thompson, in /itt. 1987),
as well as the degree of larval development
when the heads were sampled. A tiny male
reared from a capitulum of Haplopappus
venetus (Humboldt) Blake ssp. vernonioides
(Nuttall) Hall, an uncommon and appar-
ently nutritionally ill-suited host (as well as
a new host genus and species record, Was-
bauer 1972) at Cardiff-by-the-Sea, San Di-
ego Co., on 15 x 1980, illustrates the ex-
treme effect of host-plant unsuitability on
adult size in 7. stigmatica (Table 1). Dif-
ferences in mean head-part measurements
as indices of body size differences (Foote
1960a) between males and females and
among flies reared from galls versus capitula
of the same or different host plants alone
were insufficient to warrant description of
the gall-forming flies as a separate species.
F. L. Blanc, California Department of Food
and Agriculture (Retired), Sacramento, con-
firmed that specimens in Table | were 7.
stigmatica (in litt. 1987).
Galls.— Tephritis stigmatica overwinters
in southern California at least partly as im-
mature larvae in F, galls formed mostly on
low axillary branches on S. douglasii (Fig.
la). Adults that emerge from heads in late
summer and fall also overwinter. Young F,
galls examined in February (mid-winter) 1n
1987 at Pearblossom and early March 1987
at Spanish Needle Creek were a third to half
of full size. Most F, galls (Fig. 1b), like un-
galled, elongating current season branches,
showed darker (purple) coloration than galls
and branch growth formed later in the year.
The pigmentation could favor solar energy
adsorption and facilitate larval and pupal
development as well as adult emergence
during cooler months of the year. Galls and
ungalled branches that develop during the
hot summer are mostly light green (Fig. 1a,
1d). As many as five axillary branches bore
F, galls along one 10-cm basal section of
stem (Fig. la). In contrast, F, galls usually
were isolated on upper parts of aerial stems
and usually contained only one or two te-
phritids (Fig. 1d). Dissection of 72 juvenile
VOLUME 90, NUMBER |
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40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
overwintering F, galls yielded mostly 2nd
instars and small to half-grown 3rd instars.
A few fully-grown larvae and puparia also
were found in F, galls collected in early
March at Spanish Needle Creek. From one
to five individuals in the same or different
stages of growth, e.g. 2nd instars and pu-
paria, were found together in overwintering
F, galls within interconnected, longitudinal,
short-branched feeding tunnels.
Fifty overwintered, fully-formed F, galls
collected at Pearblossom in April, 1985,
contained an average of 1.5 (range, 1-6) ful-
ly-grown larvae and intact or empty puparia
(Fig. 1c). All but one of the 50 galls were
spindle-shaped (Fig. 1b, 1d). These galls
either were sessile or born on pedicels of
various lengths, up to 5 mm long. These F,
galls tapered apically as a result of continued
apical meristem growth and branch elon-
gation distal to the gall; whereas, death of
the apical meristem from larval feeding re-
sulted in a club-shaped gall, or curved gall
as a lateral branch assumed apical domi-
nance (Goeden and Ricker 1981, Goeden
1987). Dissections suggested that multiple
Ovipositions by one or more F, females in
an axillary bud initiated the formation of a
F, gall during the previous fall; whereas, a
female emerging from these galls the fol-
lowing spring probably oviposited in the
apical meristem of an upright aerial stem to
initiate F, galls. Capitula production was
observed to be delayed on F, gall-bearing
plants sampled at Spanish Needle Creek in
early August, 1984.
Branch and stem galls are formed from
expanded pith tissue, the branches and stems
swelling to about three times normal di-
ameter (Fig. 1b, 1d). Vascular strands were
deflected outward and followed the gall con-
tour. Irregular masses of hypertrophied
parenchyma cells lined the walls of the feed-
ing chamber(s), resembling round, shiny
globules which the larvae scored with
mouthhooks while feeding. Frass lined the
feeding tunnels of the late-stage 3rd instars
(Fig. lc, 1g). Only part of the gall mass was
consumed by a single larva (Fig. le, 1g);
although, the interiors of smaller galls, es-
pecially those containing two or more lar-
vae, were largely consumed (Fig. 1b). Forty-
two, spindle-shaped, overwintered, fully
formed F, galls collected at Pearblossom in
1985 averaged 2.4 + 0.1 (range, 1.4—-3.2)
cm in length and 6.5 + 0.2 (range, 4-10)
mm in greatest diameter. These galls incor-
porated as many as six nodes as indicated
by the number of lateral branches arising
thereon. The fully-grown larva extends its
feeding tunnel distally and outward to, but
not through, the epidermis leaving a round,
thin, 2-mm dia., translucent window to the
outside through which the emerging adult
eventually exits (Fig. 1f, 1h, 11). Once the
window is formed, the larva returns to its
feeding tunnel and pupariates, often with
the anterior part of the puparium projecting
into the exit tunnel, and usually facing the
window (Fig. 1g). The larval predecessor of
one of 75 (1.3%) puparia examined in over-
wintered Pearblossom galls tunneled basal-
ly in constructing its exit tunnel. A maxi-
mum of three windows was formed in one
gall from Pearblossom that contained five
puparia and one larva; otherwise, most galls
contained a single window through which
as many as three flies emerged. Exit holes
Fig. 1.
—
Galls of Tephritis stigmatica on Senecio douglasii: a, overwintering, juvenile, axillary-branch, F, galls
(0.8 x); b, fully-formed, dark-pigmented, spindle-shaped, F, gall with lateral window (2.1 =); c, same gall as in
b opened to expose four puparia within (3.3); d, fully-formed, compound, F, gall showing lateral exit hole
(0.8 x); e, same gall as in d opened to expose two empty puparia in interconnected, central-longitudinal feeding
tunnel and two lateral exit tunnels (1.3 x); f, fully-grown larva just having completed an exit tunnel ending in
circular window (4.5 «); g, puparium at juncture of short feeding tunnel and lateral exit tunnel (4.6 x ); h, closeup
view of broken, epidermal window on gall surface (3.6); i, newly-emerged female reared from gall (7.6).
VOLUME 90, NUMBER 1 4]
42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
usually were constructed in the distal third
or half of the overwintered Pearblossom galls
at, below, or between the nodes (Fig. 1b, 1d,
le, 1f, 1g, lh). Either one or both sexes of
flies (Fig. 11) emerged from individual F,
galls.
Among 39 F, galls collected at Spanish
Needle Creek, only one (2.8%) was club-
shaped, and the remainder, spindle-shaped.
The 32 spindle-shaped F, galls from which
flies had emerged in the field in the fall were
4.7 + 0.3 cm (range, 2.0-7.5) long and 8 +
| (range, 5-10) mm wide at their widest.
These spindle-shaped galls incorporated an
average of 4 (range, 1-8) nodes. The 39 F,
galls contained one or two, empty, or intact
and parasitized, puparia in interconnected
central-longitudinal feeding tunnels (Fig. le).
Thirty-three of these galls had contained a
single tephritid and six had contained two
tephritids (Fig. le), which as noted above
was considerably less than the number of
individuals in F, galls from Pearblossom
(Fig. lc). The feeding tunnels in these F,
galls averaged 1.3 + 0.2 (range, 0.5-3.5) cm
in length and were 2 mm wide. The exit
tunnels averaged 3 (range, 2-5) mm in length
and also were circular and 2 mm in cross-
sectional width. Ten intact and mostly par-
asitized empty puparia in these galls aver-
aged 4.2 + 0.1 (range, 3.6—4.6) mm in length
and 1.9 + 0.0 (range, 1.7—2.0) mm in width.
An unidentified species of Eurytoma (Hy-
menoptera: Eurytomidae) was reared from
these puparia.
Galls were collected from S. douglasii at
the following locations in addition to those
noted above: 2 4 reared from galls collected
south of Hesperia at Mojave River Forks,
San Bernardino Nat. Forest, San Bernar-
dino Co., on 21 IV 1981: 8 4 reared from
galls collected at Orcutt, Santa Barbara Co.,
on 23 VI 1981; 1 dand | 2 reared from galls
collected at Cajon Junction, San Bernardino
Co., on 8 VII 1981. This represents only a
small fraction of uncounted demes of S.
douglasii examined, but found to lack galls
during my many wide-ranging field trips
throughout southern California during
1981-1986.
DISCUSSION
Morphological study of 7. stigmatica from
galls and capitula on S. douglasii to date has
failed to support what otherwise might have
been interpreted as sympatric speciation oc-
curring on the same species of host plant.
For example, Goeden (1987) reported that
the somewhat rare native tephritid, 7ru-
panea conjuncta (Adams), facultatively
either galls the apical meristems or develops
gregariously feeding on florets (including
Ovaries) in capitula of its sole host plant,
Trixis californica Kellogg (Asteraceae). This
was the first published report of this facul-
tative mode of development in nonfrugi-
vorous Tephritidae (Zwolfer 1983, Freid-
berg 1984, Price et al. 1986). However, with
T. stigmatica, a member of the genus most
closely allied to 7rupanea according to Foote
(1959, 1960a, 1960b), development in ca-
pitula or galls apparently are not mutually
exclusive activities on S. douglasii at a par-
ticular site and season, 1.e. both gall for-
mation and feeding in capitula may occur
simultaneously during summer into fall.
However, gall formation apparently is the
requisite mode of development for the F,
generation on S. douglasii earlier in the year.
And, I suspect that at least part of the over-
wintered generation produced in branch
(axillary bud) galls begun during the pre-
ceding year, as well as overwintered adults
produced in S. douglasii capitula, reproduce
in summer at higher elevations in capitula
of a succession of alternate hosts, e.g. S.
integerrimus and S. triangularis Hooker
(Wasbauer 1972), on which galls apparently
are not formed (Tauber and Toschi 1965).
This part of the life history of 7. stigmatica
needs clarification.
Individual host plants were observed at
Spanish Needle Creek on March 3, 1987,
that bore current and previous year’s F, galls
as well as F, galls on dead upright stems that
terminated on branches with last year’s ca-
VOLUME 90, NUMBER |
pitula containing empty puparia. This pro-
vided evidence of three successive genera-
tions reproducing as gall formers augmented
by a capitula-infesting generation on the
same host individuals commensurate with
the seasonal development described above.
As previously noted, galls have not been
found on S. douglasii at most locations
where and when plants were examined in
the field as opportunity allowed since 1981,
indicating the absence of some unknown
environmental requisite(s) for gall forma-
tion by 7. stigmatica. This spotty incidence
ofa gall-forming insect occupying only tiny,
discrete, often distant, fractions of the total
geographic range of its native host plant also
was documented for the stem-galling moth,
Carollela beevorana Comstock (Lepidop-
tera: Cochylidae) by Goeden and Ricker
(1981). In contrast, a congener of C. bee-
vorana commonly forms galls on a related
host plant throughout a large part of south-
ern California (Goeden and Ricker 1986).
ACKNOWLEDGMENTS
My thanks to F. L. Blanc, R. F. Foote, A.
L. Norrbom, and J. D. Pinto for their help-
ful reviews of earlier drafts of this manu-
script. The technical support of D. W. Rick-
er, including the photography in Fig. 1, also
is gratefully acknowledged.
LITERATURE CITED
Berube, D. E. 1978. Larval descriptions and biology
of Tephritis dilacerata (Dip.: Tephritidae), a can-
didate for the biocontrol of Sonchus arvensis in
Canada. Entomophaga 23: 69-82.
Foote, R. H. 1959. A new North American species
of Tephritis, with some observations on its generic
position (Diptera, Tephritidae). Bull. Brooklyn
Entomol. Soc. 54: 13-17.
1960a. The genus Jephritis Latreille in the
Nearctic Region north of Mexico: descriptions of
four new species and notes on others. J. Kans.
Entomol. Soc. 33: 71-85.
1960b. A revision of the genus 7rupanea in
America north of Mexico. U.S. Dep. Agric. Tech.
Bull. 1214. 29 pp.
43
Foote. R. H. and F. L. Blanc. 1963. The fruit flies or
Tephritidae of California. Bull. Calif. Insect Surv.
7. 115 pp.
1979. New species of Tephritidae (Diptera)
from the western United States, Mexico, and Gua-
temala, with revisionary notes. Pan-Pac. Entomol.
55: 161-179.
Freidberg, A. 1984. Gall Tephritidae (Diptera), pp.
129-167. In T. N. Ananthakrishnan, ed., Biology
of Gall Insects. Oxford & IBH Publ. Co., New
Delhi, Bombay, Calcutta.
Goeden, R. D. 1987. Life history of Trupanea con-
juncta (Adams) on Trixis californica Kellogg in
southern California (Diptera: Tephritidae). Pan-
Pac. Entomol. 63: 284-291.
Goeden, R. D. and D. W. Ricker. 1981. Life history
of the gall-forming moth, Carollela beevorana
Comstock, on the ragweed, Ambrosia dumosa
(Gray) Payne, in southern California (Lepidoptera:
Cochylidae). Pan-Pac. Entomol. 57: 402-410.
1986. Phytophagous insect fauna of the des-
ert shrub Hymenoclea salsola in southern Cali-
fornia. Ann. Entomol. Soc. Am. 79: 39-47.
Munz, P. A. 1968. Supplement to a California Flora.
Univ. Calif. Press, Berkeley, Los Angeles, London.
224 pp.
1974. A Flora of Southern California. Univ.
Calif. Press, Berkeley, Los Angeles, London. 1086
pp.
Munz, P. A. and D. D. Keck. 1959. A California
Flora. Univ. Calif. Press, Berkeley, Los Angeles.
1681 pp.
Price, P. W., G. L. Waring, and G. W. Fernandes.
1986. Hypotheses on the adaptive nature of galls.
Proc. Entomol. Soc. Wash. 88: 361-363.
Quisenberry, B. F. 1951. A study of the genus 7e-
phritis Latreille in the Nearctic region north of
Mexico (Diptera: Tephritidae). J. Kans. Entomol.
Soc. 24: 56-72.
Shorthouse, J. D. 1980. Modification of the flower
heads of Sonchus arvensis (family Compositae) by
the gall former Tephritis dilacerata (order Diptera,
family Tephritidae). Can. J. Bot. 58: 1534-1540.
Tauber, M. J. and C. A. Toschi. 1965. Life history
and mating behavior of Tephritis stigmatica (Co-
quillett). Pan-Pac. Entomol. 41: 73-79.
Wasbauer, M. W. 1972. An annotated host catalog
of the fruit flies of America north of Mexico (Dip-
tera: Tephritidae). Calif. Dep. Agric. Bur. Ento-
mol. Occas. Pap. 19. 172 pp.
Zwolfer, H. 1983. Life systems and strategies of re-
source exploitation in tephritids, pp. 16-30. /n R.
Cavalloro, ed., Fruit Flies of Economic Impor-
tance. Proc. CEC/IOBC Int. Sym., Athens, Greece,
Nov. 1982, A. A. Balkema, Rotterdam.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 44-46
NOTES ON THE BIOLOGY OF TWO PHYCITINES
(LEPIDOPTERA: PYRALIDAE) ASSOCIATED WITH
TOUMEYELLA PINI (HOMOPTERA: COCCIDAE)
ON PINE
A. T. Drooz AND H. H. NEUNZIG
(AD) 705 Jefferson Dr., Cary, North Carolina 27511; (HN) Department of Entomology,
North Carolina State University, Raleigh, North Carolina 27695-7613.
Abstract. —The phycitines, Laetilia coccidivora (Comstock) and Ribua innoxia Heinrich
were reared from slash pine, Pinus e/liottii elliottii Engelmann, infested by the striped pine
scale, Toumeyella pini (King), in northern Florida. R. innoxia is new to the USA. It was
found previously only in Cuba and Puerto Rico, where it reportedly fed upon fungus on
pineapple.
We report here the new distribution and record of association of R. innoxia, and its
fall emergence and that of L. coccidivora in northern Florida. A comparison of the emer-
gence patterns of the 2 species and significant differences in the appearance of their cocoons
are presented.
Scale-infested slash pine, Pinus elliottii el-
liottii Engelmann, shoots were collected and
caged on 30 September, 1985, at a pine seed
orchard near White Springs, Florida, to in-
vestigate the natural control factors of the
striped pine scale, Toumeyella pini (King).
Two species of phycitine moths emerged,
Laetilia coccidivora (Comstock) and Ribua
innoxia Heinrich. L. coccidivora is preda-
ceous on numerous species of scale insects
(Heinrich 1956). R. innoxia has not been
reported from the USA, and information on
its biology is fragmentary. It had been found
only in Cuba and Puerto Rico and is con-
sidered to be a scavenger or mycetophage
(Heinrich 1956).
METHODS
A total of 100 scale-infested terminals
within reach of the ground were cut from a
number of pines. The foliage was trimmed
to ca. 2 cm to fit the cages and the twigs
were cut to 10 cm including the buds. The
twigs were caged individually in 5 cm
diam. x 8.5 cm long clear plastic vials with
snap lids. Water was not used with the twigs,
as the scale crawlers were about to hatch. A
1 cm hole was cut out of each lid and a piece
of fine mesh nylon was glued to cover the
inside of the hole. Thus, aeration was pro-
vided, yet scale crawlers and other insects
that might emerge were retained. The in-
sects were reared at ambient indoor tem-
peratures. The phycitines were larvae at this
time.
The cages were examined daily until no
moths appeared over a 2-week period.
Moths were removed each evening as they
emerged and were frozen until they could
be mounted and identified. The number and
species of moths emerging by date were re-
corded.
RESULTS AND DISCUSSION
Ninety-four L. coccidivora were reared
from 46 twigs, and 18 R. innoxia came from
VOLUME 90, NUMBER |
Figs. 1, 2. 1, Cocoon of Laetilia coccidivora. 2, Cocoon of Ribua innoxia.
45
46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
24 twigs. No moths were found in 39 cages.
Cocoons of the two phycitines differed
greatly. The cocoon of L. coccidivora 1s sim-
ilar to that made by many Lepidoptera that
pupate above ground: an elongate, white,
densely silked, but soft enclosure (Fig. 1).
Larvae of R. innoxia formed a fragile, mesh
or netlike cocoon composed of silk and frass
(Fig. 2).
Emergence of L. coccidivora started on 18
Oct., with peak emergence between 27 Oct.
and 9 Nov. and decreased through 17 Nov.,
with the last 2 moths appearing between
then and | Dec. Moths of R. innoxia began
to emerge on 4 Nov., more than 2 wk after
L. coccidivora. Only | or 2 moths emerged
occasionally until the last one appeared on
13)'Dec:
Holes in the integument of the scales,
caused by feeding of L. coccidivora larvae,
were common. The relationship between R.
innoxia larvae and the scales was not es-
tablished. Large amounts of “honeydew”
were produced by the scales, and this sub-
stance was infested with sooty mold, Cap-
nodium sp.
Heinrich (1956) postulated that R. in-
noxia larvae feed upon fungus; his type se-
ries were associated with fungus on pine-
apple, Ananas comosus (L.) Merrill, growing
in Cuba. Our brief observations suggests a
mycelium-feeding habit for the larvae. The
opinion that the larvae of L. coccidivora and
R. innoxia have diverse feeding habits is
supported by taxonomic studies of the adults
(Heinrich 1956). Laetilia and Ribua are not
closely related genera. Laetilia appears to
be allied to some of the cactus-feeding phy-
citines, whereas Ribua shows affinities to
some of the stored products pests, partic-
ularly the Indian meal moth, Plodia inter-
punctella (Hubner). Further work is needed
to establish with certainty the feeding habits
and host of R. innoxia.
ACKNOWLEDGMENTS
We thank Douglass R. Miller, USDA,
ARS, Systematic Entomology Lab., Belts-
ville, MD, for identification of the scale, and
Charles Lassiter, USDA, Forest Serv.,
Southeast. Forest Expt. Stn., Naval Stores
and Timber Production Lab., Olustee, FL
for help in collecting infested twigs. This 1s
paper No. 11072 of the Journal Series of
the North Carolina Agricultural Research
Service, Raleigh, North Carolina 27695-
7601.
LITERATURE CITED
Heinrich, C. 1956. American moths of the subfamily
Phycitinae. U.S. Natl. Mus. Bull. 207. 581 pp.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 47-51
OCCURRENCE OF SELECTED FLOWER HEAD INSECTS OF
CENTAUREA SOLSTITIALIS IN ITALY AND GREECE
STEPHEN L. CLEMENT AND TIZIANA MIMMOCCHI
USDA, ARS, Biological Control of Weeds Laboratory —Europe, % American Embassy,
Rome, Italy, APO New York 09794-0007; (SLC) USDA, ARS Plant Introduction Station,
59 Johnson Hall, Washington State University, Pullman, Washington 99164-6402; (TM)
Via Baldo degli Ubaldi 59, 00167 Rome, Italy.
Abstract.—A 1984 survey was conducted in the south-Italian mainland and central
Greece to locate sites where biocontrol specialists could collect insects of 6 promising
biocontrol agents of yellow starthistle (YST), Centaurea solstitialis L., for use in host
specificity tests. Four of these flower head insects were found: Urophora quadrifasciata
and Terellia sp. (Diptera: Tephritidae) in Italy, and Eustenopus hirtus and Larinus curtus
(Coleoptera: Curculionidae) in Greece. Urophora jaculata was the most abundant and
ubiquitous species, but it is not a potential biocontrol agent because larvae will not develop
in the heads of U.S. forms of YST. New and supplementary information on the distribution
of several YST flower head insects and the extent to which they attack heads in Italy and
Greece are also presented and discussed in relation to published information.
Centaurea solstitialis L. (yellow starthis-
tle [YST]) is a Eurasian winter annual or
biennial plant that has spread to the United
States where it is a weed on over 3 million
hectares in some western states (Maddox et
al. 1985, Maddox and Mayfield 1985). At-
tempts to control YST biologically in the
U.S. began in the 1960’s when a flower head
gall fly, Urophora jaculata Rondani (Dip-
tera: Tephritidae), erroneously called U. si-
runaseva (Hering) in some earlier refer-
ences, was introduced from Italy (White and
Clement 1987). Repeated efforts to estab-
lish this fly on U.S. forms of YST were un-
successful. However, weed biocontrol
workers discovered that a flower head wee-
vil from Greece (Bangasternus orientalis
(Cap.); Coleoptera: Curculionidae) will at-
tack and complete its development on U.S.
plants (Maddox and Sobhian 1987). This
weevil, first released in western U.S. in 1985,
1s now established in California (Maddox et
al. 1986).
We assumed that additional biological
control agents would be needed to supple-
ment the action of B. orientalis so in 1984
we surveyed YST in Italy and Greece to
locate populations of promising agents,
namely the tephritid flies Chaetorellia hexa-
chaeta (Loew), U. sirunaseva, U. quadrifas-
ciata (Meigen), and an undescribed Terellia
species (= 7. cf virens (Loew) in Sobhian
and Zwolfer [1985]), and the curculionid
beetles Eustenopus hirtus (Waltl) (= E. cf
abbreviatus Faust in Sobhian and Zwolfer
[1985]) and Larinus curtus Hochhut. We
targeted these six flower head species for
study because our preliminary work and un-
published reports at the USDA, ARS Bio-
logical Control of Weeds Laboratory—Eu-
rope (BCWLE), Rome, Italy, indicated they
had restricted host ranges in southern Eu-
rope. More than one species may be con-
fused under the name quadrifasciata (1. M.
White, pers. comm.) and some populations
of C. hexachaeta may be separate species
48
Table 1.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Sites surveyed in Italy and Greece, collection dates, total number of flower heads of Centaurea
solstitialis collected and percent damaged by insects at each site, 1984.
Site
Sites Surveyed
Italy
5 km E S. Giovanni Rotondo,
Promontorio del Gargano, Puglia Region
(41°45'N, 15°55'E)
9 km S S. Giovanni Rotondo
12 km E S. Giovanni Rotondo
15 km NW S. Paolo di Civitate, Puglia
(41°40'N, 15°20’E)
Castel del Monte, Puglia
(41°40'N, 16°15’E)
10 km N Rome, Lazio Region
(42°10'N, 12°15’E)
Greece
5.5 km W Agiokambos
(39°45’N, 22°45’E)
Xiniada (39°10'N, 22°20’E)
ca. 6 km E Karpenissi
(38°50'N, 22°50’E)
Hounti (38°45'N, 21°30'E)
ca. 2 km S Arta
(39°10'N, 20°55’E)
10 km N Igoumenitsa
(39°30'N, 20°15’E)
' Same plants were sampled each time.
(I. M. White, in press), but further taxo-
nomic study is needed to clarify these pos-
sible species groups.
The objective was to locate sources of in-
sects of the aforementioned species for use
in host specificity tests and to provide new
and supplementary information on the ex-
tent to which these and other species attack
YST heads in southern Europe. Our ap-
proach was to record the occurrence of each
flower head species on single host popula-
tions at several sites in the south-Italian
mainland and central Greece. Information
of this type is virtually nonexistent in the
literature (Zw6lfer 1965, ZwGlfer et al. 1971,
Sobhian and Zwoélfer 1985) on YST flower
head insects in Europe.
METHODS
Samples were collected at twelve sites
during the 1984 flowering season (Table 1).
i)
wn
No. No. % of
Plants Heads Heads
Collection Dates Sampled Collected Damaged
July 17 and Aug. 21 10! 454 37.89
July 17 and Aug. 21 20! 320) 818575
July 17 and Aug. 21 10! 273 24.54
July 17 52 155 14.02
July 17 and Aug. 21 16! 649 14.19
July 25, Aug. 14 10! 1627 18.19
and Sept. 7
Aug. 3 5 161 18.01
Aug. 3 5) 179 50.28
Aug. 4 5 105 41.90
Aug. 4 2 44 65.91
Aug. 5 10 86 ~—-:10.41
Aug. 5 6 258 19.77
Seven of the sites were sampled once during
July and August, but the literature (Sobhian
and Zwolfer 1985) and our unpublished data
indicate that the six species we were most
concerned with can be found on YST during
these two months. Therefore, we are rea-
sonably certain that our one-time collec-
tions were sufficient to establish the pres-
ence or absence of these insects at each site.
It was convenient to sample five Italian sites
more than once because other studies were
being conducted at or near these sites. Sur-
vey sites were roadside areas, embank-
ments, and open fields (<3.5 ha) along roads.
Each collection consisted of all heads in the
flowering and seed formation stages (see
Maddox 1981 for description of stages) from
five or more randomly selected plants per
site, except at one site where only two plants
were available for sampling. At five Italian
sites, the same plants were sampled two or
VOLUME 90, NUMBER 1
49
Table 2. Occurrence of flower head insects of Centaurea solstitialis at several sites in Italy and Greece, 1984.
Relative Occurrence!
Italian Sites Greek Sites
Species 1 3 4 5 6 1 2 3 4 5 6
Diptera
Tephritidae?
Urophora jaculata Rondani te Ser a Ft TE SEED OE ee (OTE Oke x
U. quadrifasciata (Mg.) * sia
Terellia sp. ia aa =i
Acanthiophyllus helianthi (Rossi) ae se ee
Chaetorellia sp. nr. C. carthami Stack. is oe et xs
Coleoptera
Curculionidae?’
Bangasternus orientalis (Capiomont) 12) |) 22
Eustenopus hirtus (Waltl) eS
* *
Larinus curtus Hochhut
Bruchidae
Bruchidius tuberculatus (Hochhut)*
Anobiidae
Lasioderma sp. nr. haemorrhoidale
(Illiger)>
Lepidoptera
Cosmopterigidae
Pyroderces argyrogrammos (Zeller)°
*
'** Abundant (>30 specimens emerged). ** Low abundance (5-29 specimens emerged). * Very low abun-
dance (1-4 specimens emerged). P = species present (see text for explanation).
> Identity of 4. helianthi was checked by comparison with specimens identified by Dr. R. H. Foote, former
Research Entomologist, Systematic Entomology Laboratory, HBIII, USDA, Beltsville, Maryland. Other te-
phritids were identified by Dr. I. M. White, C.A.B. International Institute of Entomology, London, England.
‘Identified by E. Colonnelli, Dipartimento di Biologia Animale e dell Uomo, Vaile dell Universita, Rome,
Italy.
‘Identified by Dr. M. L. Cox, C.A.B. International Institute of Entomology, London, England.
Identified by Dr. R. Madge, C.A.B. International Institute of Entomology, London, England.
° Identified by Dr. R. W. Hodges, Research Entomologist, Systematic Entomology Laboratory, IIBIII, USDA.
three times (Table 1). Samples from each
site were pooled to calculate the percentage
of heads with insect-damaged seeds.
The relative occurrence of each species
and the percentage of heads with damaged
seeds and receptacle tissues was assessed for
each site by rearing the insects and dissect-
ing all of the heads in a laboratory at the
BCWLE. A species was artibrarily rated as
abundant, low in abundance, or very low in
abundance according to the number of
emerging adults (see Table 2). Reared in-
sects were identified to species. Because less
than 10% of the eggs of B. orientalis survive
to the adult stage (Sobhian and Zwolfer
1985, Clement and Sobhian, unpub. data),
we assumed that very few, if any, adults
would be reared-out. Thus, we recorded the
presence or absence of this weevil at each
site by looking for its eggs, which are usually
laid singly on leaflets near a flower bud (Bu
1-2 stages of Maddox 1981) and are covered
by a characteristic black cap.
RESULTS AND DISCUSSION
The percentage of heads with insect-dam-
aged seeds varied from 13.75-37.89% (av-
erage of 20.43% of heads attacked) in Italy
and 10.41-65.91% (average 34.38%) in
Greece; less than 20% of the heads were
50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
damaged at 4 Italian and 3 Greek sites (Ta-
ble 1). Species packing (no. of species) per
site ranged from 2-6 (average 3.67) in Italy
and 2-5 (average 3.0) in Greece (Table 2).
In contrast, Sobhian and Zwéolfer (1985) re-
ported average levels of resource utilization
(% of heads attacked) and species packing
of 36.9% and ‘tabove” 4 species for Italy,
and 77.3% and 9.5 species for Greece, but
these higher levels were based on samples
from the south-Italian mainland and Sicily,
and northern Greece. The average levels re-
ported above for central Greece are com-
parable to the ones Sobhian and Zwéolfer
(1985) reported for Yugoslavia, Bulgaria and
Romania (35.6% and 3.4 species). Thus, the
collective evidence suggests that predispers-
al seed predation by YST flower head in-
sects is not markedly high in many areas of
southern Europe, including Greece where
Sobhian and Zwolfer (1985) and Zwéolfer
(1985) reported that average levels of re-
source utilization and species packing were
significantly higher than they were in the
western Mediterranean (Italy and France).
High rates of parasitization of several species
(Sobhian and Zwélfer 1985) might account
for the fairly low levels of resource utiliza-
tion in many areas.
In all, we found 11 species of seed pred-
ators to be associated with YST heads (Ta-
ble 2). Three of these, Acanthiophyllus he-
lianthi (Rossi) (Diptera: Tephritidae),
Lasioderma sp. nr. haemorrhoidale (Illiger)
(Coleoptera: Anobiidae) and Pyroderces ar-
gyrogrammos (Zeller) (Lepidoptera: Cos-
mopterigidae) use plants in several genera
as hosts, and 8, B. orientalis, E. hirtus, L.
curtus (Coleoptera: Curculionidae), Terellia
sp., U. quadrifasciata, U. jaculata Rondani,
Chaetorellia sp. nr. carthami Stack. (Dip-
tera: Tephritidae), and Bruchidius tubercu-
latus (Hochhut) (Coleoptera: Bruchidae) ap-
pear to be restricted to the genus Centaurea
in the field (Sobhian and Zwéolfer 1985,
Clement, unpub. data). Urophora jaculata
was the most ubiquitous and abundant
species, but this tephritid is not a candidate
biocontrol agent because it will not develop
in the heads of U.S. forms of YST (White
and Clement 1987). Chaetorellia sp. nr. car-
thami was widespread in Italy; however, this
species will form hybrids with C. carthami
Stack., a pest of cultivated safflower, so its
safety as a biocontrol agent has been ques-
tioned by biocontrol workers (Sobhian and
Zwolfer 1985). A third stenophagous species
(i.e. restricted to Centaurea spp.), B. tuber-
culatus, has been disqualified because adults
were found in the heads of cultivated saf-
flower, Carthamus tinctorius L., in northern
Greece (Sobhian and Zwé6lfer 1985). Four
of the 6 species that we set out to find were
detected; U. quadrifasciata and Terellia sp.
were represented in the guild of flower head
insects in Italy while EF. hirtus and L. curtus
were detected in Greece. None of these 4
species were abundant at any site (Table 2).
The failure of this survey to detect U.
sirunaseva and C. hexachaeta was unex-
pected because Sobhian and Zwo6lfer (1985)
reported that both species occur throughout
much of southern Europe. However, recent
taxonomic studies (White and Clement
1987, White in press) have revealed a more
restricted distribution for these tephritids.
This new information on U. sirunaseva and
C. hexachaeta, together with information
from this survey and the literature on E.
hirtus (Ter-Minasyan 1967, Fremuth 1982,
Sobhian and Zwolfer 1985) and L. curtus
(Zwolfer et al. 1971, Fremuth 1982, Sob-
hian and Zwo6lfer 1985) suggest that none
of these 4 species are rare but the 2 weevil
species are better able to exploit YST over
a much wider geographical area than are the
2 tephritid species. The seemingly broad eco-
climatic tolerances of EF. hirtus and L. curtus
would improve their chances for establish-
ment in the western U.S. where YST occurs
in markedly different climatic and vegeta-
tional zones (Maddox 1981, Maddox et al.
1985, Maddox and Mayfield 1985, Roché
et al. 1986).
In summary, this survey has enabled us
to: pinpoint sites where biocontrol special-
VOLUME 90, NUMBER |
ists might be able to collect insects of 4 po-
tential agents for use in host-specificity tests;
clarify the Palearctic distribution of several
YST flower head insects, including 6 species
that are promising biocontrol agents; and
contribute towards a better understanding
of the extent to which YST flower heads are
attacked by insects in southern Europe.
ACKNOWLEDGMENTS
We are grateful to the taxonomic spe-
cialists (listed in Table 2) who identified
specimens, I. White, D. Maddox and an
anonymous reviewer for comments on the
manuscript, and S. Craig for typing the
manuscript.
LITERATURE CITED
Fremuth, J. 1982. Cleoninae aus der Turkei und den
angrenzenden gebieten (Coleoptera: Curculioni-
dae). Fragm. Entomol., Roma 16(2): 239-258.
Maddox, D. M. 1981. Introduction, phenology, and
density of yellow starthistle in coastal, intercoast-
al, and central valley situations in California. Agric.
Res. Serv. U.S. Dept. Agric. ARR-W-20, July. 33
pp.
Maddox, D. M., A. Mayfield, and N. H. Poritz. 1985.
Distribution of yellow starthistle (Centaurea sol-
stitialis) and Russian knapweed (Centaurea re-
pens). Weed Science 33: 315-327.
Maddox, D. M. and A. Mayfield. 1985. Yellow star-
thistle infestations are on the increase. Calif. Agric.
39(1 1-12): 10-12.
Maddox, D. M. and R. Sobhian. 1987. Field exper-
iment to determine host specificity and oviposi-
tion behavior of Bangasternus orientalis and Ban-
gasternus fausti (Coleoptera: Curculionidae),
biological control candidates for yellow starthistle
51
and diffuse knapweed. Environ. Entomol. 16: 645-
648.
Maddox, D. M., R. Sobhian, D. B. Joley, A. Mayfield,
and D. Supkoff. 1986. New biocontrol agent in-
troduced for yellow starthistle. Calif. Agric. 40(1 I-
12): 4-5.
Roché, B. F., G. L. Piper, and C. J. Talbott. 1986.
Knapweeds of Washington. Coop. Ext. Bull. 1393.
Washington State Univ., Pullman, WA. 41 pp.
Sobhian, R. and H. Zwolfer. 1985. Phytophagous
insect species associated with flower heads of yel-
low starthistle (Centaurea solstitialis L.). Z. Ang.
Ent. 99: 301-321.
Ter-Minasyan, M. E. 1967. Zhuki-dolgonosiki pod-
semeistra Cleoninae fauny SSSR. Tsvetozhily 1
stebleedy (Triba Lixini). Nauka Publishers, Len-
ingrad. 141 pp.
White, I. M. and S. L. Clement. 1987. Systematic
notes on the Urophora Robineau-Desvoidy (Dip-
tera: Tephritidae) species associated with Centau-
rea solstitialis L. (Asteraceae, Cardneae) and other
Palearctic weeds adventive in North America. Proc.
Entomol. Soc. Wash. 89: 571-580.
White, I. M. In press. Taxonomic problems in the
genus Chaetorellia Hendel. /n Cavalloro, R., ed.,
Proc. CEC-IOBC Int. Symp. on Fruit Flies of Eco-
nomic Importance, 7-10 April 1987, Rome, Italy.
Balkema, Rotterdam.
Zwolfer, H., K. E. Frick, and L. A. Andres. 1971. A
study of the host plant relationships of European
members of the genus Larinus (Col.: Curculioni-
dae). Tech. Bull. Commonw. Inst. Biol. Control,
No. 14, pp. 97-143.
Zwolfer, H. 1965. Preliminary list of phytophagous
insects attacking wild Cynareae (Compositae) in
Europe. Commonw. Inst. Biol. Contr. Tech. Bull.,
No. 6, pp. 81-154.
Zwolfer, H. 1985. Insects and thistle heads: Resource
utilization and guild structure, pp. 407-416. Jn
DelFosse, E. S., ed., Proc. VI Int. Symp. Biol. Contr.
Weeds, 19-25 August 1984, Vancouver, Canada.
Agric. Can.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 52-54
UNUSUAL OVIPOSITION BEHAVIOR ON EVERGREEN AZALEA BY
THE ANDROMEDA LACE BUG STEPHANITIS TAKEYAI
(DRAKE AND MAA) (HETEROPTERA: TINGIDAE)
JOHN W. NEAL, JR.
Florist and Nursery Crops Laboratory, Horticultural Science Institute, Agricultural
Research Service, U.S. Department of Agriculture, Building 470, Beltsville, Maryland
20705.
Abstract.—The andromeda lace bug, Stephanitis takeyai (Drake and Maa), an adventive
tingid from Asia, has a feeding host range in North America of twelve species in the
Lauraceae, Salicaceae, Ericaceae and Styracaceae. Hybrid evergreen azalea (Rhododendron
sp.) is reported as a new feeding and breeding host. Females oviposit in the midrib of the
azalea leaf, a behavior different from that on its preferred host, Japanese andromeda
(Pieris japonica). A comparison is made of oviposition site behavior between the androme-
da lace bug, and S. pyrioides (Scott) the azalea lace bug.
Stenophagous lace bugs in North Amer-
ica have been assigned common names
based on the major reproductive hosts. Mi-
nor feeding hosts usually involve congeneric
species. Drake and Ruhoff (1965) report that
‘All species [of lace bugs] are rather highly
specialized in their food habits, and gen-
eration after generation live on the same
kind of plant or closely related ones.”’ Host
plants of the andromeda lace bug Stephani-
tis takeyai (Drake and Maa) (= S. globuli-
fera Matsumura), an adventive tingid from
Asia (Schread 1953), have been reported by
Drake and Ruhoff (1965), Schread (1953),
Dunbar (1974), and Wheeler (1977) and
world-wide include 12 species in Lauraceae,
Ericaceae, Salicaceae and Styracaceae. Bai-
ley’s (1951) review of New England tingids
contains a report that a few S. takeyai were
found in association with azalea lace bug,
S. pyrioides (Scott), on a single deciduous
azalea Rhododendron sp. in Connecticut.
Bailey (1974) also reported S. takeyai on
Rhododendron calendulaceum (Michx.)
Torr. Wheeler (1977) reported specimens in
the U.S. National Museum of Natural His-
tory from azalea at a nursery in Falls Church,
Virginia, June 1969.
Dunbar (1974) reported that S. takeyai
oviposits on the abaxial leaf surface, usually
along the side of the midrib of the Japanese
andromeda Pieris japonica(Thunb.) D. Don,
and that overwintering eggs are found along
the midrib but on occasion were distributed
over the undersurface of the leaf.
I observed adults of S. takeyai in low
numbers on evergreen and deciduous aza-
leas throughout Prince George’s and How-
ard counties, Maryland. These observations
suggested that azalea is used by S. takeyai
as a minor host, and prompted this study
and report of insect behavior, host fitness
and potential pest status.
MATERIALS AND METHODS
Containerized Japanese andromeda har-
boring all stages of S. takeyai were pur-
chased from a retail nursery in Burtonsville,
VOLUME 90, NUMBER |
te
Fig. 1.
shy ng “4
(A) Frass-covered eggs of S. takeyai placed lateral to the midvein of an andromeda leaf, (B) Eggs of
S. takeyai placed in the midvein of an azalea leaf, (C) Eggs S. pyrioides placed lateral to the midvein of an azalea
leaf. Arrow identifies one of several frass-covered eggs.
Maryland. Plants were transplanted to 11.6
liter (3 gal) containers, held in a heated
greenhouse, and watered as needed. Stems
of various lengths were removed periodi-
cally to a bottle of water and placed in a
gauze covered plexiglass cylindrical cage (32
cm high x 31 cm dia.) in a Sherer walk-in
rearing chamber programmed at 26.1 + 1°C,
with a photoperiod of 14:10 (L:D).
To test for host preference and acceptabil-
ity, stems of the evergreen hybrid azalea
‘Martha Hitchcock’ (Rhododendron mu-
cronatum x Shinnyo-no-tsuki) were placed
in the cylinder contiguous to the infested
andromeda cuttings. Leaves with adults and
nymphs of S. takeyai found on the azalea
during daily observations were transferred
for isolation to a similar cage containing an
uninfested second azalea cutting. Trans-
ferred adults were allowed to feed and ovi-
posit. Water was added to the bottle as
needed. Both azalea and andromeda cut-
tings were introduced weekly to the cylinder
with andromeda lace bugs, and the transfer
of adults from andromeda to azalea was
conducted over several weeks.
Leaves with eggs of S. takevai were com-
pared against leaves with eggs of S. pyrioides
removed from the same azalea cultiver used
as host plants to maintain a greenhouse col-
ony.
RESULTS
Tingids that leaf feed in an inverted po-
sition defy gravity and deposit their fecal
material on the abaxial surface at random.
Preliminary tests confirmed that by remov-
ing all frass by rinsing the leaf with warm
running water, S. takeyai and S. pyrioides
confined their egg laying either in or adja-
cent to the midrib or at major lateral veins
depending on host. Defecation by the fe-
male on the operculum leaves a prominent
mark at the egg site (Fig. 1, arrows). Stepha-
nitis takeyai fed and oviposited on azalea
cuttings. Originally some eggs on this host
54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
hatched, but most failed to produce second
instars. Washing leaves prior to hatch to
remove sticky plant exudate greatly reduced
first instar mortality; nymphs developed to
the adults when foliage was washed. The
presence of plant exudates on the cuttings
is a phenomenon due to the growing of aza-
leas in the protective greenhouse environ-
ment. Exudates are normally reduced by rain
or rendered non-sticky by the accumulation
of airborne particles.
Stephanitis takeyai oviposited only in and
along the midrib in all azalea leaves ob-
served (Fig. 1B), whereas S. pyrioides de-
posited its eggs below and lateral to the mid-
rib (Fig. 1C). Oviposition by S. takeyai on
Japanese andromeda included periodic
placement below and lateral to the midrib,
but eggs were never inserted in the midrib
(Fig. 1A). Andromeda leaves have conspic-
uous midribs, but they are not raised as in
other hosts such as azalea, possibly account-
ing for this change in oviposition behavior.
Wheeler (1977) found S. takeyai eggs in-
serted in the midvein of spicebush, Lindera
bezoin (L.) Blume, and sassafras, Sassafras
albidum (Nutt.). This ovipositional site
preference by the andromeda lace bug on
azalea and other hosts is unusual, because
when compared, the andromeda lace bug
Oviposits on andromeda lateral to the un-
raised midrib which is similar to oviposi-
tion by the azalea lace bug below and lateral
to the raised midrib on azalea. This behav-
ior by S. takeyvai on azalea raises the ques-
tion what would be the oviposition site on
andromeda if the midrib were raised.
These findings determined that an ever-
green azalea can be a suitable feeding-breed-
ing host for S. takeyai. Further, it was found
that females oviposit in and along the mid-
vein of the azalea leaf which is different than
when on andromeda. Fertilized eggs hatched
and nymphs developed to adults normally.
These results suggest that S. takeyai could
develop to be a late season threat to azalea
production. There is, however, no data to
suggest that eggs of S. takeyai overwinter
on azalea.
ACKNOWLEDGMENTS
Thanks are extended to Richard C.
Froeschner (USNM) and Thomas J. Henry
(USDA, SEL) for constructive comments
and helpful suggestions on an early draft of
the manuscript.
LITERATURE CITED
Bailey, N. S. 1951. The Tingoidea of New England
and their biology. Entomol. Am. 31: 1-133.
Bailey, N. S. 1974. Additional notes on Stephanitis
takeyai in New England (Heteroptera: Tingidae).
Psyche 81: 534-538.
Drake, C. J. and F. A. Ruhoff. 1965. Lacebugs of the
World: A catalogue (Hemiptera: Tingidae). Bull.
U.S. Natl. Mus. 243: 1-634.
Dunbar, D. M. 1974. Bionomics of the andromeda
lacebug Stephanitis takeyai, pp. 277-289. In Mem.
Conn. Entomol. Soc. 25th Ann.
Schread, J. C. 1953. Control of the andromeda lace
bug Stephanitis globulifera and the holly leaf miner
Phytomyza ilicis. Conn. Agric. Exp. Stn. Bull. 568:
1-13.
Wheeler, A. G., Jr. 1977. Spicebush and sassafras as
new North American hosts ofandromeda lace bug,
Stephanitis takeyai (Hemiptera: Tingidae). Proc.
Entomol. Soc. Wash. 79: 168-171.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 55-61
A REVIEW OF THE SOUTH PACIFIC GENUS AUSTROMEGALOMUS
ESBEN-PETERSEN (NEUROPTERA: HEMEROBIIDAE) WITH
A DESCRIPTION OF A NEW SPECIES FROM RAPA
JOHN D. OSwALD
Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York
14853-0999.
Abstract. —The hemerobiid genus 4Austromegalomus Esben-Petersen is reviewed. Aus-
tromegalomus and its type species 4. brunneus are redescribed and A. insulanus is de-
scribed as new. Figures, known distributions and a key to the two recognized species are
provided. Several shared characters of the male ectoprocts, mediuncus and parameres
suggest that the genera Austromegalomus, Conchopterella and Drepanacra are closely
related.
The genus Austromegalomus Esben-Pe-
tersen, 1935, was proposed to accommo-
date the single species 4. brunneus Esben-
Petersen which was described in the same
paper from three male specimens collected
on the South Pacific island of Tahiti. Until
now no additional specimens or species of
Austromegalomus have been recorded in the
literature. In this paper Austromegalomus
insulanus is described as new, from 20 spec-
imens collected on the island of Rapa lo-
cated approximately 1200 km (750 mi.) SSE
of Tahiti, and the genus Austromegalomus
and the male of 4. brunneus are redescribed.
As with many early hemerobiid descrip-
tions, the original descriptions of Austro-
megalomus and A. brunneus are based al-
most entirely upon venational characters.
Austromegalomus insulanus is shown here
to exhibit a wide range of intraspecific vari-
ation in a variety of forewing venational
traits and venation is judged inadequate to
confidently separate the two species. The
descriptions presented here emphasize
characters of the male genitalia.
Intraindividual, as well as interindividu-
al, variation in venational characters is
common in Austromegalomus. In tabulat-
ing the variability of several forewing ve-
national traits, both forewings of each in-
dividual were scored for each trait. Estimates
of mean forewing length were based on mea-
surements of a single forewing of each spec-
imen. Consequently, the sample sizes given
in the species descriptions for venational
traits are twice those given for estimates of
mean forewing lengths.
Austromegalomus Esben-Petersen
Austromegalomus Esben-Petersen, 1935:
139. Type species: Austromegalomus
brunneus Esben-Petersen, 1935: 140, by
original designation.
Diagnosis. — Head: Temporal sutures well
developed, marked internally by prominent
costae; epicranial suture absent; labial palp
three segmented, distal segment longest and
with an apical subsegment, palpimacula
present; maxillary palp five segmented, dis-
tal segment longest and with an apical sub-
segment.
Forewing: Length 5-9 mm, hind margin
rounded, apex broadly pointed; costal area
56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6.
the first (proximal) oblique branch of the forewing radius (diagrammatic). 4-6. Female. 4, apex of abdomen
(lateral view). 5, spermatheca and apex of bursa. 6, subgenitale (ventral view).
broad proximally, recurrent vein pectinate-
ly branched; proximal half of subcostal space
with 2 crossveins (the distal of these rarely
absent); radius with 4-10 oblique branches;
2 well developed, posteriorly convergent,
gradate series in outer half of wing.
Hindwing: Radius with 2 oblique branch-
es; Cu2 frequently, though not always,
traceable to near the posterior margin either
as a distinct or indistinct vein or a row of
setae; Outer gradate series well developed;
inner gradate series with 1-3 crossveins or
absent.
Male genitalia: Tergite nine a sclerotized
arch, lateral lobes dilated ventrally; sternite
nine in ventral view a remiform plate (Figs.
7, 13), shallowly arched in anterior view;
ectoproct elongate oval, without narrowed
projecting lobes; gonarcus with arms of
Austromegalomus insulanus. 1, venation of forewing and hindwing. 2-3, two possible states of
moderate size the greater part of which pro-
ject free into body cavity, exposed surface
limited to a narrow strip to which the medi-
uncus and epimeres are fused; mediuncus a
rigid plate dorsal to epimeres and para-
meres, bilobed proximally and distally, lon-
gitudinal midline shallowly depressed,
proximal lobes strongly divergent and fused
to gonarcus at a pair of widely separated
points on opposite sides of gonarcus bridge,
medial margins of proximal lobes and pos-
terior margin of gonarcus bridge enclosing
a triangular membranous fenestra; epimeres
a pair of elongate strips of sclerotized mem-
brane lying in the membranous sack sup-
ported dorsally by the mediuncus, fused to
gonarcus ventral to fusion of mediuncus with
gonarcus; parameres with internal end of
apophysis proxima enlarged in lateral view,
VOLUME 90, NUMBER 1
.2mm
[smn
9-12,15-18
7-8,13-14
Figs. 7-18. 7-12. Austromegalomus brunneus, male. 7, ninth sternite (ventral view). 8, apex of abdomen
(lateral view). 9, gonarcus, mediuncus and epimere (lateral view). 10, gonarcus, mediuncus and epimere (dorsal
view). 11, parameres (dorsal view). 12, parameres (lateral view). 13-18. 4. insulanus, male. 13, ninth sternite
(ventral view). 14, apex of abdomen (lateral view). 15, gonarcus, mediuncus and epimere (lateral view). 16,
gonarcus, mediuncus and epimere (dorsal view). 17, parameres (dorsal view). 18, parameres (lateral view).
external lobes narrow and linear in dorsal
view but with their apices upturned in lat-
eral view, medioventral surfaces of lobes
sclerotized, laterodorsal aspects membra-
nous.
Female genitalia: See below under A. In-
sulanus.
Natural history and immature stages. —
Unknown.
Distribution.—Known only from the
French Polynesian islands of Tahiti and
Rapa.
Etymology.—Name unexplained but al-
most certainly from the Latin “australis,”
southern, and “.Wegalomus,” a hemerobud
genus to which Esben-Petersen allied Aus-
tromegalomus. Gender: masculine.
Discussion.—Esben-Petersen diagnosed
Austromegalomus by the branching ar-
rangement of the first oblique branch of the
radius (‘basal Rs” of Esben-Petersen). In
Austromegalomus the vein track which an-
teriorly parallels the median flexion line is
nearly straight. The curvature of vein seg-
ments confluent at forks along this track are
usually somewhat asymmetric. The branch-
es originating at these forks tend to form a
linear series on the anterior side of the track
58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(Fig. 2). In an alternate state found in many
other hemerobiid genera, the vein forks
along this track are more symmetric giving
the track a more or less undulate appearance
(Fig. 3). Though these states are rather dis-
tinctive when viewed as the opposite ends
of a morphocline, intraspecific variation
within A. insulanus encompasses both states.
Too few specimens are available to ade-
quately assess the degree of intraspecific
variation of this trait in 4. brunneus. Fur-
thermore, as pointed out by Esben-Petersen
(1935) and Handschin (1955), similar
asymmetric patterns of veins along this track
are found in other hemerobiid genera (e.g.
Drepanacra and Conchopterella). For these
reasons this character cannot be used as a
synapomorphy of Austromegalomus. Due
to still unresolved questions concerning the
homologies and polarities of diagnostic
characters of the male genitalia, I have been
unable to confidently identify any synapo-
morphic characters for this taxon.
Key TO ADULT MALES OF
AUSTROMEGALOMUS
la. Emargination separating distal lobes of medi-
uncus V-shaped (Fig. 16); gonarcus bridge ar-
cuate in dorsal view (Fig. 16); epimeres short
and narrow and not subtended by an acces-
sory sclerite (Fig. 15) (Rapa) A. insulanus n. sp.
1b. Emargination separating distal lobes of medi-
uncus quadrate (Fig. 10); gonarcus bridge
quadrate (Fig. 10); epimeres long and broad
and subtended distally by a small accessory
sclerite (Fig. 9) (Tahiti) :
.. A. brunneus Esben-Petersen
Austromegalomus insulanus, NEw SPECIES
Figs. 1-6, 13-18
Diagnosis.— Diagnosed by characters in
key couplet la. The longer forewing length
and the alternating light and dark brown
segments of the forewing longitudinal veins
may also be diagnostic, though not enough
specimens of A. brunneus are available to
fully assess the potential overlap of these
characters with those found in A. insulanus.
Description. — Forewing (Fig. 1): Length
6.09-8.48 mm (x = 7.06, N = 20); longi-
tudinal veins mostly marked with alternat-
ing light and dark brown segments, though
several specimens (likely teneral) with ve-
nation nearly evenly pale; membrane hya-
line to brown, darker adjacent to dark vein
segments. Venation (Fig. 1, N = 40): Num-
ber of subcostal crossveins in proximal half
of subcostal space = 2 (39 wings), 3 (1 wing);
number of oblique radial branches proximal
to stigmal subcostal crossvein = 6 (1 wing),
7 (6 wings), 8 (16 wings), 9 (14 wings), 10
(3 wings); number of inner gradate cross-
veins anterior to cubitus = 10 (3 wings), 11
(7 wings), 12 (20 wings), 13 (8 wings), 14 (2
wings); number of outer gradate crossveins
anterior to cubitus = 11 (2 wings), 12 (2
wings), 13 (8 wings), 14 (15 wings), 15 (13
wings); number of forkings of first oblique
radial branch proximal to inner gradate se-
ries = | (7 wings), 2 (29 wings), 3 (4 wings).
Male genitalia: Apex of abdomen as in
Fig. 14. Gonarcus (Figs. 15, 16): gonarcus
bridge arcuate in dorsal view. Mediuncus
(Figs. 15, 16): distal pair of lobes contiguous
medially at their bases. Epimeres (Figs. 15,
16): short and very narrow, weakly tanned
and easily overlooked; not extending as far
posteriorly as in 4. brunneus and not sub-
tended distomedially by a pair of accessory
sclerites. Parameres (Figs. 17, 18): internal
end of apophysis proxima enlarged but not
as prominently as in 4. brunneus; apices of
external lobes tipped with a minute spine.
Female genitalia (Figs. 4, 5, 6): Gon-
apophyses laterales remiform, styli arising
dorsad of middle of sclerites; gonapophyses
posteriores present as a pair of narrow rods;
subgenitale present, attached to ventral body
wall by a short membranous tube, apex
emarginate; spermatheca composed of a
darkly tanned bulb and a pair of ducts—a
short duct joining the bulb to the bursa and
a longer convoluted duct arising from the
distal end of the bulb.
Etymology. —An adjective from the Latin
“insula,” island, in reference to the island
type locality.
VOLUME 90, NUMBER 1
Distribution.— Known only from the type
series from the South Pacific island of Rapa
(French Polynesia, Austral Islands).
Primary type material examined.— Male
holotype (USNM). Verbatim label data:
“Rapa/Anatakuri/Bay 28 XI 63,” “J. F. G.
Clarke/Thelma M. Clarke,” ““USNM Loan/
USNM Loan,” ‘“Holotype/Austromegalo-
mus/insulanus Oswald/J. D. Oswald 1987.”
Condition: Excellent, no parts missing.
Genitalia cleared and placed in a glycerin
filled microvial pinned below the specimen.
Other material examined.— 19 paratypes.
RAPA ISLAND: 2 6, Anatakuri Bay,
28.xi.1963 (Clarke) (USNM); 3 4, 2 2, Haur-
ei, 15.x.-3.xii.1963 (Clarke) (USNM); | 4,
1 2, Mau Bay, 23.x.1963 (Clarke) (USNM);
2 2, Maugaoa, 244 m & 290 m, 18.1x.-
23.x1.1963 (Clarke) (USNM); | 4, 1 2, Man-
gaoa [sic = Maugaoa] Pk., NE ridge, 305-
366 m, 6.vil.1934 (Zimmerman) (BPBM);
1 6, 2 2, Maurua, 61 m & 183 m, 25.1x.-
25.x.1963 (Clarke) (USNM); | 2, Mt. Oror-
angi, SE valley, 183-244 m, 3.vu.1934
(Zimmerman) (BPBM), 2 2, Point Teak-
aurae, 61 m, 7.x.1963 (Clarke) (USNM).
Note.— For a general account of the Clarke
Expedition to Rapa, including collecting lo-
calities and physiography, see Clarke (1971).
Austromegalomus brunneus Esben-Petersen
Figs. 7-12
Austromegalomus brunneus Esben-Peter-
sen, 1935: 140 (original description, fig-
ures): Esben-Petersen 1937: 51 (listed);
Handschin 1955: 9 (compared to Con-
chopterella).
Diagnosis.— Diagnosed by characters in
key couplet 1b. The shorter forewing length
and the uniform brown coloration of the
forewing longitudinal veins may also be di-
agnostic, though not enough specimens of
A. brunneus are available to adequately as-
sess the potential range of intraspecific vari-
ation in these characters.
Description. — Forewing: Length 5.37-
5.56 mm (x = 5.47, N = 2); longitudinal
59
veins uniformly brown, membrane also
brown. Venation (N = 4): Number of sub-
costal crossveins in proximal half of sub-
costal space = 2 (4 wings); number of oblique
radial branches proximal to stigmal sub-
costal crossvein = 4 (3 wings), 5 (1 wing);
number of inner gradate crossveins anterior
to cubitus = 8 (1 wing), 9 (1 wing), 10 (2
wings); number of outer gradate crossveins
anterior to cubitus = 11 (2 wings), 12 (2
wings); number of forkings of first oblique
radial branch proximal to inner gradate se-
ries = 3 (4 wings).
Male genitalia: Apex of abdomen as in
Fig. 8. Gonarcus (Figs. 9, 10): gonarcus
bridge quadrate in dorsal view; anterodorsal
region of gonarcus arm broader in lateral
view than in A. insu/anus. Mediuncus (Figs.
9, 10): distal pair of lobes separated medi-
ally at their bases by a space about equal to
width of each lobe. Epimeres (Fig. 9, 10):
prominent, long and broad relative to 4.
insulanus; apex of each epimere subtended
medially by a small, weakly sclerotized and
poorly delimited accessory sclerite. Para-
meres (Figs. 11, 12): internal end of apoph-
ysis proxima considerably enlarged in lat-
eral view; apices of external lobes tipped
with a minute spine.
Female: Unknown.
Etymology.—An adjective from the Latin
“brunneus,” dusky or tawny, in reference
to the brownish coloration of the body and
forewing.
Distribution.— Known ony from the type
series from the South Pacific island of Tahiti
(French Polynesia, Society Islands).
Primary type material examined.— Male
holotype (BPBM). Verbatim label data:
“Society Is./1500'/Tahiti I.,” *“Fautaua Val./
IX-11-28,° “A. M. Adamson/Collector,”
“Pacific Entomological Survey,” “TYPE
791,” “Austromegalo=/mus brunneus/é n.
sp./det. Esben-Petersen.”’ Condition: Ex-
cellent, only several small pieces of wings
missing. Genitalia cleared and placed in a
glycerin filled microvial pinned below the
specimen.
60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Other material examined.—One male
paratype (BPBM). Collection data same as
holotype. A second paratype stated in the
original description to have been retained
by Esben-Petersen has not been traced.
PHYLOGENETIC POSITION OF
AUSTROMEGALOMUS
The morphology of the male genital struc-
tures of Austromegalomus suggests that it is
closely related to the southern hemisphere
genera Drepanacra and Conchopterella,
which are known from the Australian region
and the Juan Fernandez islands respective-
ly. This evidence supports the conclusions
of Esben-Petersen (1935) and Handschin
(1955) based on venational characters. The
following three shared traits appear to sup-
port the hypothesis that these genera are
closely related:
(1) The male ectoprocts are elongate oval
without projecting narrow lobes. The ec-
toprocts of many other hemerobiid genera
are variously lobed.
(2) The mediuncus forms a rigid horizon-
tal plate which is bilobed distally and at-
tached to the gonarcus by a pair of widely
divergent proximal arms. The full distri-
bution of this state and its possible deriv-
atives within the Hemerobiidae needs ad-
ditional investigation.
(3) The parameres consist ofa prominent,
anteriorly projecting apophysis proxima and
a pair of small apical lobes. The medioven-
tral surfaces of the apical lobes are sclero-
tized, the dorsolateral surfaces membra-
nous. The parameres of many other
hemerobiid genera possess various other
dorsal and/or lateral lobes and patterns of
sclerotization.
Though one or more of the preceding
characters may in the future prove synap-
omorphic of a clade (Austromegalomus +
Conchopterella + Drepanacra), at present,
the polarities of these shared traits in rela-
tion to their homologues found in other
hemerobiid genera are not known with con-
fidence. Consequently, firm conclusions
about the relative relationships among these
three genera are presently impossible.
Currently available comparative analyses
of important hemerobiid character com-
plexes (e.g. wing venation and male geni-
talia) are in most cases insufficiently de-
tailed, with regard to hypotheses of
homologies and/or polarities, to allow con-
fident differentiation of synapomorphies and
symplesiomorphies. Consequently, it has
not been possible to fully assess the status
of some Austromegalomus character states
which might later prove to be useful indi-
cators of phylogenetic relationships.
Several factors have contributed to the
dilemma described above. First, no recent
comprehensive revision of the Hemerobi-
idae, with attention to character analysis, is
available. Second, many terms widely em-
ployed in the current nomenclature of neu-
ropterous genital structures were originally
proposed expressly as labels of convenience,
without critical investigation of the homol-
ogies of the labeled structures. Though some
of these terms have apparently been applied
to homologous structures (e.g. the gonar-
cus), others have not (e.g. the mediuncus-
arcessus). Uncritical application of existing
genitalic terms has hindered the improve-
ment of hypotheses of homology for some
genitalic structures.
Most hemerobiid genera are currently di-
agnosed, at least in part, on the basis of
distinctive combinations of male genitalic
characters. Given the importance of this
character complex, additional comparative
morphological studies are needed to clarify
the homologies and polarities of genitalic
characters. Until such analyses are under-
taken, the phylogenetic position of Austro-
megalomus, and many other hemerobiid
genera, will likely remain unclear.
ACKNOWLEDGMENTS
I thank Oliver S. Flint, Jr., of the National
Museum of Natural History (USMN),
Washington, D.C., and Gordon M. Nishida
of the Bernice P. Bishop Museum (BPBM),
VOLUME 90, NUMBER 1 61
Honolulu, Hawaii, for loaning material used | Esben-Petersen, P. 1935. Neuroptera from the So-
in this study. I also thank James K. Liebherr ciety Islands. Bull. Bernice P. Bishop Mus. 142:
d 137-142.
and Quentin D. Wheeler, Department of 1937. Check List of Neuroptera Planipennia
Entomology, Cornell University and an of Oceania. Occas. Pap. Bernice P. Bishop Mus.
anonymous reviewer for providing com- 13: 49-60.
ments on an earlier draft of this paper. Handschin, E. 1955. Los insectos de las Islas Juan
Fernandez. 15. Neuroptera. Rev. Chil. Entomol.
LITERATURE CITED 4: 3-20.
Clarke, J. F. G. 1971. The Lepidoptera of Rapa Is-
land. Smithson. Contrib. Zool. 56: 1-282.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 62-65
LECTOTYPE DESIGNATION FOR EMPIS CHICHIMECA
WHEELER AND MELANDER (DIPTERA: EMPIDIDAE)
WILLIAM J. TURNER
Department of Entomology, Washington State University, Pullman, Washington 99164.
Abstract.— A lectotype and six paralectotypes for Empis (= Lamprempis) chichimeca
Wheeler and Melander are designated from the syntype series. Diagnostic leg features of
the lectotype male are discussed and illustrated. Comments concerning the lectotype and
its presumed detached hind leg are provided.
The Neotropical genus Lamprempis
Wheeler and Melander presently includes
22 species of metallic greenish blue to black
flies with an evanescent anal wing vein and
peculiarly ornamented legs. The often di-
morphic sexes show presence or absence of
pennate hair fringes and other modifica-
tions of the legs. Several species are known
from one sex only. Little information is
available about the biology and habits for
species of Lamprempis. Smith (1975) re-
ports that one species, L. sazimae, occurs
in great numbers in the highlands of Minas
Gerais, Brazil, where it serves as an impor-
tant pollinating agent for certain Umbelli-
ferae and Eriocaulaceae growing in mead-
ows at 1300 m above sea level.
The purpose of this paper is to report the
interesting results of my study of the avail-
able syntype series for Empis chichimeca
Wheeler and Melander (1901: 368), the type
species of Lamprempis, and to designate a
lectotype for this species.
In 1981, while examining the A. L. Me-
lander types of Empis Linnaeus at the Na-
tional Museum of Natural History (USNM),
I found several syntype specimens of E. chi-
chimeca. The original series consisted of nine
specimens (two males and seven females)
collected by H. H. Smith in Amula, Guer-
rero, Mexico. I could find only three female
specimens in the USNM type collection.
Also present was the apparent right hind leg
of a male, glued to a card rectangle and la-
beled “type” in Melander’s hand. It had been
attached on its anterior side and embedded
in an unknown adhesive, but the characters
of the exposed posterior surface are easily
visible and match the description of the
species provided by Wheeler and Melander
(1901). At that time I supposed that the leg,
which possesses characters sufficient for rec-
ognizing the species, was the only portion
remaining of one male, the remainder de-
stroyed by pests or otherwise lost.
Later, in the collection of the American
Museum of Natural History (AMNH), I dis-
covered another part of the same syntype
series (one male, three females). All speci-
mens are in good condition. Interestingly, I
found that the AMNH male is intact except
for the missing right rear leg. After re-ex-
amining the USNM point-mounted leg, I
concluded that it likely represents the miss-
ing leg from the AMNH specimen.
There is no indication when the leg of the
male syntype was broken or removed from
the otherwise intact specimen. One can only
speculate why the leg was not kept with the
associated male. The detached leg, however,
possesses the diagnostic features of the
species (see Smith 1975) and it serves as an
VOLUME 90, NUMBER |
a
—
ra
Pil
Pes Pare
SS ~~
\\ wae
<i
Fig. 1.
excellent reference even without the intact
specimen itself. The leg may have permitted
Melander to have the ideal “‘reference spec-
imen” in his collection while Wheeler had
the remainder of the specimen. Because I
cannot find the other male in the syntype
series, I assume that the detached leg may
have served this function for Melander.
Because this male and especially its de-
tached leg bear the diagnostic features of the
species, and nearly intact AMNH male is
hereby designated the lectotype of Empis
(= Lamprempis) chichimeca and its de-
tached right rear leg (in the USNM) 1s sim-
ilarly marked with my red lectotype label.
I have illustrated the detached right leg (Fig.
1) along with the left hind leg (Fig. 2). The
remaining six females of the known syntype
series have been labeled paralectotypes.
The male specimen (AMNH) selected as
63
2
Right hind leg (detached) of Empis (= Lamprempis) chichimeca, lectotype male, in posterior view.
lectotype is in excellent condition, except
for the missing right rear leg, and bears the
following label data: ““Amula, Guerrero,
6000 ft., Sept., H. H. Smith/W. M. Wheeler
Collection/TYPE NO. | > AMNH [red la-
bel]/AMNH, DIZ No. 918 [white label]/
LECTOTYPE, Empis (= Lamprempis) chi-
chimeca Wheeler and Melander, des. W. J.
Turner 1988 [red label, hand written].”
The point-mounted leg (USNM) lacks a
locality label but has the following label data:
“E. chichimeca W & M TYPE [white label
in Melander’s hand]/Cotype Lamprempis
chichimeca W & M [red cotype label in Me-
lander’s hand]/A L Melander Coll. 1961
[white label with green checked margin]/
LECTOTYPE (part), Empis (= Lamprem-
pis) chichimeca Wheeler and Melander, des.
W. J. Turner 1988 [red label, hand written].”
Besides the lectotype male, I found that
64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2.
only two of the available females (AMNH)
bear the same label data. The remaining
four females (USNM, AMNH) have iden-
tical labels but were collected in August.
Only seven specimens (one male, six fe-
males) from the original syntype series of
nine specimens have been accounted for.
The location of one additional male and
female remain unknown. Because much of
the insect material described in the Biologia
Centrali Americana was subsequently de-
posited in the British Museum (Natural
History) (BM), I asked John Chainey, Cu-
rator of Diptera (BM), to check for syntype
specimens of this species. He was unable to
locate any representatives of chichimeca
under either Empis or Lamprempis in the
BM collection. Further, there was no ref-
erence made to the species in any lists of
holdings by the museum. All of the known
syntypes, now in either the USNM or
AMNH, were originally in the collections
of W. M. Wheeler or A. L. Melander re-
/
Left hind leg of Empis (= Lamprempis) chichimeca, lectotype male, in anterior view.
spectively, as indicated by the personal col-
lection labels attached to the specimens.
Only one USNM female lacks such a label
probably because it was placed in that col-
lection by the authors shortly after the
species was described.
DISCUSSION
In 1901 Empis chichimeca was described
by W. M. Wheeler and A. L. Melander, and
placed into their new subgenus Lamprempis
along with five other species from Mexico.
Coquillett (1903) elevated Lamprempis to
generic rank and designated EF. chichimeca
as the type species. Although the diagnostic
features for this species have never been
illustrated, the species is easily keyed. Smith
(1975) includes E. chichimeca in his ten-
tative key to the described species of Lam-
prempis and uses essentially the same word-
ing as in the original description by Wheeler
and Melander for describing the unique fea-
tures of the hind leg: “Hind femora pos-
VOLUME 90, NUMBER |
teroventrally with two slender finger-like
processes, with an emargination between
them; hind tibia posteriorly with a stout
scoop-shaped process truncated and flat-
tened at the extremity; hind basitarsus in-
crassate with an anterior projection tipped
with two small black spines.”
The hind legs of this species are somewhat
asymmetrical with minor differences in
structures from left to right. Similar asym-
metry can be found in the armature of the
hind legs of males of Empis (Enoplempis)
mira Bigot. In the case of E. chichimeca the
right femur appears to have a small hooked
process on the posterior surface near the
base of the larger, digitate process. Proximal
to the small hook is a low, irregular carina
with toothlike projections running oblique-
ly across the subbasal fourth of the hind
surface. Unfortunately the leg 1s embedded
in an adhesive glue matrix and the edge of
the glue follows along the carina. On the left
femur, in comparison, the small hook is
lacking as is the oblique carina. The de-
scription was likely made from the right (de-
tached) appendage as it refers to the two,
slender, fingerlike processes, probably the
thicker digitate process and the small hooked
one. I found that the similar digitate pro-
cesses located medially on the posteroven-
tral margin of both hind femora also differ
from left to right in orientation, the left one
being more linear, the right more oblique.
One will also see from the illustrations that
the general outline of each femur is different
as well.
Both hind tibiae are moderately concave
medially on both the anterior and posterior
surfaces along nearly their entire length. The
concavities appear natural and not simply
artifacts of the legs having collapsed at death.
Although the surrounding areas are heavily
bristled, the depressed spaces remain essen-
tually bare.
Smith (1975) indicates in couplet 14 of
his key that E. chichimeca has only simple
leg bristles. However, pennate bristles can
be found in two irregular rows along the
65
entire dorsal surface of each hind tibia and
flanked by fewer, less developed but still
flattened bristles. An additional five or six
pennate setae can be found at the extreme
base of the tibiae ventrally while each femur
bears a cluster on its inner and dorsal sur-
faces apically. Although pennate leg bristles
are not uncommon in females of some Em-
pidinae (e.g., Rhamphomyia species), they
are unusual in males and appear restricted
to Lamprempis.
ACKNOWLEDGMENTS
I thank Lloyd Knutson, Systematic Ento-
mology Laboratory, USDA, % National
Museum of Natural History, Washington,
D.C., and Pedro Wygodzinsky, Department
of Entomology, American Museum of Nat-
ural History, New York, New York, who
made syntype specimens of Empis (Lam-
prempis) chichimeca in their respective col-
lections available for study. Dr. Knutson
also provided work space and optical equip-
ment in his office during my sabbatical stay
in Washington, D.C. John Chainey at the
British Museum (Natural History) checked
for additional syntype material and his ef-
forts are appreciated. Thanks also to Paul
Arnaud, Jr., James Johnson and Norman
Woodley who reviewed this manuscript.
Scientific Paper No. 7622, Agricultural Re-
search Center, College of Agriculture and
Home Economics, Washington State Uni-
versity, Pullman, Washington. The work was
conducted under Project 9043.
LITERATURE CITED
Coquillett, D. W. 1903. The genera of the dipterous
family Empididae, with notes and new species.
Proc. Entomol. Soc. Wash. 5: 245-272.
Smith, K.G. V. 1975. A new species of Lamprempis
Wheeler & Melander from Brazil, with a key to
the described species of the genus (Diptera, Em-
pididae). Pap. Avulso Zool. (Sao Paulo) 29: 21-
26.
Wheeler, W. M. and A. L. Melander. 1901. Supple-
ment. Empididae, pp. 366-367. Jn Godman, F.
D. and O. Salvin, eds., Biologia Centrali Ameri-
cana. Insecta. Diptera. Vol. |. 378 pp., 6 pls. Lon-
don.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 66-75
MAYACNEPHIA SALASIT (DIPTERA: SIMULIIDAE), A NEW
BLACK FLY SPECIES FROM COSTA RICA
J. RAMiREZ-PEREZ, B. V. PETERSON, AND M. VARGAS V.
(JRP) Instituto de Biomedicina, P.O. Box 4043, Caracas, Venezuela; (BVP) Systematic
Entomology Laboratory, BBII, Agricultural Research Service, USDA, % National Mu-
seum of Natural History, NHB-168, Washington, D.C., U.S.A. 20560; (MVV) Centro de
Investigacion y Diagnostico en Parasitologia, Universidad de Costa Rica, Ciudad Uni-
versitaria Rodrigo Facio, Costa Rica.
Abstract.—The female, male, pupa and larva of Mayacnephia salasi, new species, are
described and illustrated. This genus is recorded from Costa Rica for the first time, and
is now known from western North America, Mexico, Guatemala, Costa Rica and Panama.
A key to the species of Mayacnephia known in the pupal stage is provided.
The genus Mayacnephia Wygodzinsky
and Coscar6n (1973) was established to in-
clude six Mesoamerican species that had
been placed previously in the genus Cnephia
Enderlein. Diaz Najera (1971) described
another new species in the genus Cnephia
that belongs in Mayacnephia, and J. L. Pe-
tersen (1985) described a new species from
Panama. B. V. Peterson (1981), using an
expanded concept of the genus, assigned two
species to it from western United States and
an undescribed species from Canada. The
species described below is the eleventh de-
scribed species now assigned to the genus
and the first to be reported from Costa Rica.
We describe this new species to make its
name available for biological studies cur-
rently being conducted on black flies in Cos-
ta Rica. We also provide a key to the known
pupae of Mayacnephia, including the un-
described species from Canada, and include
distributions and references to published
figures of these species.
Mayacnephia salasi Ramirez-Perez,
Peterson, and Vargas,
New SPECIES
Figs. 1-18
Female (preserved in alcohol).— General
body color dark brown. Length: body, 2.88
mm; wing, 3.48 mm.
Head: Lightly grayish pollinose. Frons
(Fig. 5) narrow, nearly parallel sided, only
slightly widening dorsally, about five times
as long as width at narrowest point, about
, width of head; slightly paler than occiput,
densely covered with long, decumbent, pale
yellow pile, and with a few black setae lat-
erally. Clypeus concolorous or slightly light-
er than frons; slightly longer than wide;
densely covered with long, ventromedially
directed, pale yellow pile interspersed with
some dark setae laterally and ventrally. Oc-
ciput densely covered with long, pale yellow
pile and with a few dark setae mid-dorsally;
postocular setae black, closely bending over
eye margin. Antenna with nine flagello-
meres; scape and pedicel pale yellowish,
contrasting with rest of flagellum which is
dark brown; pedicel and first flagellomere
larger than other antennomeres and sub-
equal in length and width; remaining flagel-
lomeres subequal in length to each other but
tapering in width distally; fine pubescence
and longer setae dark. Mandible (Fig. 3) with
33-40 serrations. Blade of maxilla (Fig. 2)
with 21-25 retrorse teeth. Palpus (Fig. 1)
dark brown to black, proximal two palpo-
meres pale brownish, distal two palpomeres
slightly lighter brownish than palpomere
three, all with black setae; palpomere five
about 3 longer than palpomere three. Sen-
VOLUME 90, NUMBER 1
sory vesicle as in Fig. 4, about '2 as long as
its segment, proximally situated, its neck
short, arising anterodorsally and extending
vertically, with an enlarged ovoid mouth.
Median proximal space of cibarium shal-
low, broadly U-shaped, and without den-
ticles; dorsolateral arms short, rather broad,
sclerotized, inner surface of each arm with
a rather extensive patch of minute setulae
arising from dark granular bases.
Thorax: Antepronotal lobe concolorous
with scutum, with dense, long, pale yellow
pile interspersed with a few dark setae. Post-
pronotum yellowish brown, distinctly paler
than scutum, covered with long, semi-erect
pale yellow pile. Scutum dark brown to
blackish brown except lateral margins which
are narrowly paler, and with a grayish pol-
linose border extending around lateral and
hind margins, posterior declivity broadly
grayish pollinose; each anterolateral corner
of scutum, adjacent to postpronotal lobes,
a paler yellowish brown color which, in pos-
terior view, extends posteriorly as a faint,
narrow, submedian line, and with a similar
faint, slender, median line or vitta that ex-
tends to posterior declivity, these lines not
visible in anterior view; scutum densely
covered with short, recumbent, pale yellow
pile that is longer along anterior and lateral
margins and still longer posteromedially,
also a few scattered dark setae present. Scu-
tellum paler brownish than scutum, lightly
grayish pollinose, densely covered with long,
pale yellow and dark setae. Postnotum only
faintly darker than scutellum, with a faint
pollinosity. Anterior half of pleuron dark
brown mottled with some paler areas, and
distinctly paler brown on posterior half that
is mottled with some darker areas; prester-
nal lobe with moderately long pale yellow
pile; anepisternal membrane distinctly paler
than rest of pleuron; mesepimeral tuft dark.
Wing (Fig. 6) membrane hyaline but with a
light brownish tinge; veins yellowish brown.
Base of C, stem vein, and other veins with
dark pile; Sc setose ventrally; R, setose dor-
sally; R,,; setose ventrally; cell bm present;
67
C with spiniform setae as long as regular
setae, apical half of R, with a few similar
spiniform setae; fringe of anal lobe and ca-
lypter pale yellow. Stem of halter and base
of knob brownish yellow, rest of knob yel-
low; stem with pale yellow pile. Legs rather
uniformly dark brown; all coxae with both
pale and dark setae but pale setae more nu-
merous on fore and midcoxae, rest of setae
on legs dark; hind basitarsus about seven
times as long as broad. Calcipala short but
distinct, broadly rounded; pedisulcus ab-
sent. Claw only slightly curving from base,
with a prominent, bluntly pointed, basal
tooth that is wider than claw and over '2 as
long.
Abdomen: Yellowish brown, basal scale
(tergite one) dark brown, with a fringe of
long, pale yellow pile; tergites broad, tergite
two widest, others decreasing in width pos-
teriorly; tergites yellowish centrally with
darker brown margins, sparsely covered with
short, pale yellow setae; pleural membrane
paler and more yellowish brown, with both
pale yellow and dark setae; sternites scarcely
distinguishable; venter of abdomen pale
brownish yellow, with mostly short, dark
setae but with some scattered pale yellow
setae. Terminalia as in Figs. 7-9. Anal lobe
(Fig. 9) narrow dorsally, broadening ven-
trally, widest at about midheight, broadly
rounded ventrally, with a slight but distinct
notch posteroventrally, not produced be-
neath cercus, moderately setose. Cercus
subrectangular, hind margin varying from
strongly rounded to nearly straight. Hypo-
gynial valves short, barely reaching to bases
of cerci; valves subtruncate posteriorly, their
medial margins lightly sclerotized; lightly
setose. Stem of genital fork (sternite nine)
(Fig. 7) long, heavily sclerotized, slightly
more than '3 longer than arms; arms short,
rather weakly sclerotized except for a short,
heavily sclerotized rodlike extension on each
side emanating from stem of genital fork;
arm with a sclerotized subtriangular tooth-
like process on anterior margin; arms rather
broadly attached to tergite nine. Spermathe-
68
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
REL
TF AAMIREZ 7e
VOLUME 90, NUMBER 1
ca (Fig. 8) kidney-shaped, moderately scler-
otized, with a faint, loose reticulate pattern
but without internal spicules; with only a
small clear area at junction with spermathe-
cal duct.
Male (preserved in alcohol).—General
body color velvety dark brown to black.
Length: body, 3.12 mm; wing, 3.18-3.45
mm.
Head: Frons and clypeus lightly grayish
pollinose, with erect, black pile. Occiput with
long, dark brown to black setae. Antenna
entirely dark brown; first flagellomere an-
gularly broadened distally, slightly longer
than pedicel: fine pubescence pale yellow,
longer setae black. Palpomere three darker
than other palpomeres, all with black pile
interspersed with a few more yellowish se-
tae; palpomere five about 3 longer than pal-
pomere three and about '2 longer than pal-
pomere four. Sensory vesicle about ' as
long as its segment; neck short, enlarging to
form a round mouth.
Thorax. Antepronotum and postprono-
tum concolorous, slightly paler brown than
scutum; with some dark setae and some pale
yellow pile having dark bases. Scutum with
a light grayish pollinosity; densely covered
with short, recumbent, pale yellow pile that
is longer anteriorly, laterally and postero-
medially. Scutellum brown, paler than scu-
tum; densely covered with long, erect, dark
setae and some decumbent, pale yellow se-
tae. Postnotum concolorous with scutellum,
lightly grayish pollinose. Pleuron brown an-
teriorly, grayish pollinose, becoming paler
brown medially and posteriorly; anepister-
nal membrane brownish yellow; mesepi-
meral tuft dark. Wing membrane hyaline
but with a distinct brownish tinge, veins
yellowish brown. Base of C, stem vein, and
other veins with dark pile; Sc lightly setose
69
ventrally; R, setose dorsally; R,,; setose
ventrally; C and about distal 73 of R, with
spiniform setae that are about as long as
regular setae; cell bm present; fringe of anal
lobe brownish yellow; fringe of calypter pale
yellow. Knob of halter brown, stem yellow
with pale yellow pile. Legs rather uniformly
dark brown, with dark pile; hind basitarsus
swollen, about 3.5 times as long as broad;
calcipala short but distinct, rounded api-
cally; pedisulcus absent.
Abdomen: Basal scale dark brown, with a
fringe of long dark pile; tergites with dark-
ened margins, paler medially, covered with
short, brown pile; sternites concolorous with
tergites, with long, dark setae. Terminalia
as in Figs. 10 and 11. Gonocoxite (Fig. 10)
subtriangular to conical, greatest length and
width nearly equal, covered with pile on all
but basal '4 to /s. Gonostylus short, about
'’» longer than greatest width at base; taper-
ing to a bluntly pointed, apical margin bear-
ing two tiny terminal spines. Body of ventral
plate of aedeagus (Fig. 11) subrectangular,
broader than long, with a short, ventrally
directed hirsute lip; in ventral view, apical
margin slightly convex and shortly pro-
duced nipplelike medially, lateral margins
slightly concave just distal to junction with
basal arms; basal arms bowed, nearly equal
in length to body of ventral plate; median
sclerite of aedeagus short, Y-shaped, stem
variably longer than arms; aedeagal mem-
brane rather densely covered with numer-
ous groups of 8-10 minute setulae arranged
in rows. Plate of endoparameral organ an
elongate subtriangular shape, moderately
sclerotized; arm moderately long, and twist-
ing.
Pupa.—Length of specimens at hand 3.5
mm. Respiratory organ (gill) (Fig. 12) 1.62
mm long; consisting of four rather short but
—
Figs. 1-11.
Mayacnephia salasi. Figs. 1-9, female. 1, Maxillary palpus. 2, Blade of maxilla showing retrorse
teeth. 3, Tip of mandible showing serrations. 4, Enlarged view of sensory organ of third palpomere. 5, Front
view of frons and ocular notches. 6, Portion of wing showing setation. 7, Genital fork (sternite 9). 8, Spermatheca.
9, Anal lobe and cercus. Figs. 10-11, male. 10, Gonocoxite and gonostylus (dorsal (inner) surface). 11, Ventral
plate of aedeagus, ventral view.
TRAMRE? PERL
15
Figs. 12-18. Mayacnephia salasi. Figs. 12-14, pupa. 12, Respiratory organ (gill). 13, Frons. 14, Abdomen
showing chaetotaxy on dorsal (d) and ventral (v) surfaces. Figs. 15-18, Larva. 15, Hypostoma. 16, Inner distal
and subapical margins of mandible showing dentation. 17, Antenna. 18, Hypostomal cleft.
70
VOLUME 90, NUMBER 1
cylindrical, inflated saclike filaments origi-
nating from a common short, rather broad
base covered with minute spicules; these
saclike filaments are rather uniform in length
and width, nearly transparent and have a
minutely granular texture that is visible only
at high magnifications. Head and thoracic
integument glabrous; antenna of male ex-
tending about '2 distance to hind margin of
head; antenna of female extending about %4
or more of distance to hind margin of head;
a single stout seta present near inner corner
of each antenna, and two or three, some-
what separated, shorter and more slender
setae present along outer margin of frons at
about midlength of antenna (Fig. 13). Dor-
sum of thorax without any trace of integu-
mental pattern; each side of thorax with
about two anterodorsal and one postero-
dorsal, and one anteroventral and one pos-
teroventral long, simple trichomes, antero-
dorsal trichomes stoutest. Chaetotaxy of
each lateral half of abdominal tergites as
follows (Fig. 14): tergite one with five or six
fine setae; tergite two with four or five fine
setae and four stouter hooks; tergites three
and four each with 2-5 minute setae and
four anteriorly directed spines along pos-
terior margin; tergite five with about seven
minute setae; tergite six with two or three
minute setae and a row of minute, poste-
riorly directed spinules along anterior mar-
gin; tergite seven with three minute setae
and an anterior row of minute spinules; ter-
gite eight with two minute setae and an an-
terior row of minute spinules; tergite nine
with a few minute spinules anteriorly, and
two long caudal spines situated on two
slightly swollen convexities, these spines
slightly curved, tips divergent, each with a
long, stout seta near base posteriorly. Chae-
totaxy of each lateral half of sternites as fol-
lows: sternite three with three or four weakly
sclerotized hooklets and one fine seta; ster-
nites four and five each with four or five
hooklets; sternite six with three well-devel-
oped hooks; sternite seven with two well-
developed hooklets, and one pale, medial
71
oval area; sternite eight with one strong
hooklet, and one fine seta; sternite nine with
two fairly strong setae in striated mem-
brane, plus a strong seta at base of caudal
spine, otherwise bare; sternites three to eight
each with a variably sized but distinct patch
of minute spinules. Striated pleural mem-
brane on each side of: segment five with two
fine setae one of which is in a platelet-like
sclerite; segment six with one hooklet and
one fine seta in a platelet-like sclerite; seg-
ment seven with one hooklet; segment eight
with one fine seta; segment nine with eight
stout hooks that may be simple, bifurcate,
or grapnel-shaped; intersegmental membra-
nous area between segment eight and nine
with three short but distinct nipple-like
bumps on each half. Cocoon a loosely
wooven, saclike structure without any def-
inite shape, and covered by detritus.
Larva (mature, with fully developed res-
piratory histoblasts).— Length 6.5-8.5 mm.
General body color pale creamy brown; in-
tersegmental lines narrow, slightly lighter
than rest of abdomen. Head capsule pale
yellowish brown; head spots pale brown but
darker than surrounding fulvus area, an-
teromedian and posteromedian spots slen-
der, elongate, the two sets of spots well sep-
arated, first and second anterolateral spots
roundish, about equal in size and distinctly
separated, posterolateral spots slightly dark-
ened and somewhat obscure; eye spots small.
Postocciput with broad gap dorsally, en-
closing small cervical sclerites. Antenna (Fig.
17) pale brownish; about 4 as long as stalk
of labral fan; proportions of segments (basal
to apical) 1:7.7:2.6. Labral fan with 25-33
(av. 29) primary rays. Hypostoma as in Fig.
15, with 13 teeth arranged in three main
groups of 4 + 3 + 4 plus a small tooth on
each side of base of median tooth; median
tooth long, subequal to longest lateral teeth
of each side; each lateral group of teeth sim-
ilar in structure to median group, consisting
of one main tooth and a smaller tooth on
each inner and outer margin, and a short,
more ventral, lateral tooth; outer lateral
72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
margins of hypostoma with 3-6 weak ser-
rations and two or three long and one or
two short hypostomal setae, with longest
seta reaching tip of longest hypostomal
tooth. Hypostomal cleft (Fig. 18) poorly de-
fined, a broad, shallow, U-shaped excava-
tion extending about ¥ distance to base of
hypostoma. Hypostomal bridge distinctly
longer (25:18) than hypostoma. Mandible
(Fig. 16) with one large apical tooth, three
stout preapical teeth followed by a series of
six or seven more seta-like teeth, and two
outer teeth; inner subapical ridge with about
16-25 fine serrations. Maxillary palpus
about 2.5 times as long as width at base.
Lateral plate of proleg extending about ' or
more length of apical segment; irregularly
subquadrate to subtriangular, greatest width
and height nearly equal; lightly sclerotized,
with about 20 very slender rod-like exten-
sions projecting distally toward bases of
hooks; circlet of apical hooks arranged in
about 17 rows. Segment eight of abdomen
with two short, broadly rounded tubercles
that extend about , to '4 depth of abdomen
below their points of attachment. Anal pa-
pillae with three simple lobes; minute rectal
setulae present lateral to anterodorsal arms
of anal sclerite. Anterodorsal arms of anal
sclerite about '2 as long as posteroventral
arms; anterior arms slender, posterior arms
considerably broader; sclerotized platelike
junction of arms bearing 8-10 fine setae,
these setae short but conspicuously longer
than rectal setulae. Posterior circlet of hooks
consisting of about 8-10 hooks in about 62-
65 rows.
Types.— Holotype, 2 (mounted on five
slides), temporary stream (#45a) located at
one side of the road between km 96 and 97,
near La Georgina on route from Cartago to
San Isidro del General, Provincia San José,
Costa Rica, November 14, 1983, C. R.
Méndez and A. Solano V.
Paratypes.—1 9°, 1 6, same data as type
except preserved in alcohol and terminalia
of both specimens mounted on slides; 1 4,
same data except November 1, 1983
(mounted on three slides); 1 6, same data
except November 6, 1983 (mounted on three
slides); 1 2, same data except November 12,
1983 (mounted on four slides); 2 99, 1 é (all
pinned), same data except October 28, 1983,
A. Solano V. and H. Mayreno; | 4(mounted
on six slides), same data except October 24,
1983, and | ¢(mounted on six slides), same
data except November 7, 1983, A. Solano
V. and H. Mayreno; 2 pupae, same data
except October 28, 1983 (one pupa mount-
ed on two slides, the other pupa mounted
on four slides) ; 30 pupae (four mounted on
slides), same data except October 28, 1983,
A. Solano V. and H. Mayreno; 13 pupae,
same data except July 14, 1986, A. Solano
V.and H. Mayreno; | larva, same data ex-
cept June 7, 1983 (mounted on five slides):
1 larva, same data except June 17, 1983
(mounted on six slides); 12 larvae (five
mounted on six slides each), same data ex-
cept July 14, 1986, A. Solano V. and H.
Mayreno.
Holotype deposited in the collection of
the U.S. National Museum of Natural His-
tory, Washington, D.C. Paratypes are de-
posited in the U.S. National Museum of
Natural History, and the entomology col-
lection of the Department of Parasitology,
University of Costa Rica.
This species is dedicated to Eng. Luis An-
gel Salas F., a distinguished acarologist, and
Professor Emeritus at the School of Agron-
omy, University of Costa Rica.
Biological notes.—All available speci-
mens of M. sa/asi came from the same tem-
porary stream (numbered 45a) at the type
locality. This small, shallow, clean-water
stream is situated at an elevation of 3150
meters in the bottom ofa deep, steep-sloped
gulley in an area partially to heavily shaded
by forest, and with a moderate amount of
vegetation on the banks. The stream is about
one meter wide, with a bed of rocky sections
and muddy areas. The water flows at a slow
to moderate rate, has a temperature that
ranges between 10-14°C, and pH values are
between 7.1 and 7.8. Larvae are present in
the stream throughout the year.
Remarks.—Wygodzinsky and Coscaron
VOLUME 90, NUMBER |
(1973) recognized two groups of species in
Mayacnephia based on the chaetotaxy of
the abdomen and the form of the respiratory
gills of the pupae. The new species described
here belongs to their ““apomorphic” group
having a reduced number of filaments and
without at least an apical filamentous por-
tion to the filaments, as well as having a
larger number of sternal setae and hooks.
Mayacnephia salasi has only four simple,
inflated, saclike filaments in the respiratory
organ, and has more numerous spines on
the sternal sclerites (sternite five with about
5+5, sternite six with about 3+ 3, and ster-
nite seven with about 2+ 2) characteristic of
this group of species. The recently described
M. fortunensis Petersen (1985), from Pan-
ama, also belongs to this group.
Mayacnephia salasi appears to be most
similar to M. grenieri (Vargas and Diaz Na-
jera) in the adult and pupal stages. The fe-
male of M. sa/asi can be most easily differ-
entiated from M. grenieri by the antenna,
which has a yellowish scape and pedicel that
contrast with the dark brown flagellum (in
grenieri the antenna is entirely yellow ex-
cept for the black first flagellomere), the hal-
ter with its yellow knob and brownish yel-
low stem (knob black, stem whitish), and
the genital fork whose arms diverge from
the stem in a broad V-shape and have rect-
angular plates that are wider than long, each
plate with a small, sclerotized, toothlike
process on its anterior margin (arms of gen-
ital fork diverge from the stem at nearly
right-angles so the posterior space between
the arms 1s broadly U-shaped, the plates are
shorter, broader and subrectangular, and
each has a prominent tooth-like process).
The male of MM. sa/asi can be distinguished
from M. grenieri by the all black antenna
(in grenieri the antenna is yellow except for
the black first flagellomere), the pale yellow,
recumbent pile of the scutum (recumbent
pile light brown), halter brown with a yellow
stem (halter entirely black), and by the shape
of the ventral plate of the aedeagus which,
in ventral view, is more rectangular with a
more broadly rounded distal margin, and
73
longer basal arms (more triangular in ven-
tral view, with narrower and more pointed
apical margin and much shorter basal arms).
Larvae of M. salasi can be distinguished
from M. grenieri by the following combi-
nation of characters: length 6.5-8.5 mm
(larvae of grenieri range from 8.0—9.0 mm),
pale creamy brown color (distinctly yellow-
ish but more opaque especially posteroven-
trally), antenna about *4 length of stalk of
labral fan (antenna as long as stalk of labral
fan), labral fan with 25-33 primary rays (40-
45 primary rays), hypostoma with 13 apical
teeth (9 apical teeth), hypostomal bridge
distinctly longer (25:18) than hypostoma
(hypostomal bridge and hypostoma nearly
equal in length), and posterior circlet with
62-65 rows of hooklets (76 rows of hooklets
in grenier!).
The species of Mayacnephia, as with some
other groups of Simulidae, are most easily
differentiated on the basis of the number
and shape of the filaments of the pupal re-
spiratory organ (gill). The following key to
the pupae of the species of Mayacnephia
will differentiate M/. sa/asi from the other
described species of the genus. The pupa of
M. osborni (Stains and Knowlton), a species
described from California, does not appear
in the key because it is unknown. We are
unable to prepare a reliable key to the other
stages of the species of Mayacnephia for
lack of specimens. Dalmat (1955) and Diaz
Najera (1962) provided keys that include
the larvae of various species now included
in Mayacnephia, and Dalmat provided a
key to the males and females of the three
species known from Guatemala.
KEY TO SPECIES OF
MAYACNEPHIA PUPAE
1. Respiratory organ with two long, swollen tu-
bular filaments arranged in the form of a V
(figs. 17-18 in Dalmat 1949; fig. 1OB in Wy-
godzinsky and Coscaron 1973). Highlands of
Guatemala, Mexico (Chiapas) aguirrei (Dalmat)
— Respiratory organ with three or more fila-
ments of varying form and arrangement 2
2. Respiratory organ with three long, swollen,
tubular filaments (fig. 20 in Diaz Najera 1962).
Mexico (Oaxaca) ....... mixensis (Diaz Najera)
74
10.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Respiratory organ with four or more filaments
OL WAT YIN) S POU oa ao rsycs cs, y's Sac eaters agshnyays cours 3
. Respiratory organ with four’ alarients ree A
Respiratory organ with six or more filaments
BH eigee cehatiat JL dss Bee 5
Filaments short, inflated and saclike with
broadly rounded tips, all arising from a com-
mon base (fig. 12 herein). Costa Rica ......
eae ic Se EI ch ae salasi n. sp.
Filaments longer, swollen but not greatly in-
flated or saclike, tubular with pointed apices,
arranged in two pairs (fig. 22 in Vargas and
Diaz Najera 1948; fig. 9A in Wygodzinsky
and Coscaron 1973). Mexico (Veracruz) .
. grenieri (Vargas and Diaz Najera)
Respiratory organ with six long, swollen, tu-
bular filaments that are rounded distally, an-
terior two filaments single, posterior two fil-
aments branching into two petiolate pairs (fig.
4 in Coleman 1953). California
stewarti (Coleman)
Respiratory organ with seven or more fila-
ments of varying form 6
Respiratory organ with seven or eight fila-
MGT: creas pecs cetitrs Haseeno ara e 7
Respiratory organ with iy l- 15 filaments 9
Respiratory organ with seven filaments 8
Respiratory organ with eight filaments (fig. 27
in Currie 1986). Western Canada Me
; .(unnamed species X)
Respiratory organ n with four swollen, tubular
filaments, three of which, in turn, give rise to
two slender filaments distally, and the fourth
which has a single slender terminal filament
(fig. 8A in Wygodzinsky and Coscaron 1973).
Highlands of Guatemala roblesi (Leon)
Respiratory organ with four swollen, tubular,
clavate filaments; anteromedial and postero-
medial filaments unbranched although the
former sometimes with a variably developed
thumblike hump on its mesal surface; antero-
lateral filament branching into two petiolate
filaments, and posterolateral filament branch-
ing into three filaments (fig. 8 in Petersen
1985). Panama _fortunensis Petersen
Respiratory organ with 11-12 filaments . 10
Respiratory organ with 14-15 filaments aris-
ing from four thickened main trunks (figs. 13-
18 in Diaz Najera 1971). Mexico (Coahuila)
_ muzquicensis (Diaz Najera)
Respiratory organ with 11 tubular filaments
that are variably swollen basally, some of the
filaments are much longer than the others (fig.
9 in Diaz Najera 1962). Mexico (Morelos)
.. atzompensis (Diaz Najera)
Respiratory organ with 11-12 long, slender,
tapering filaments and one short, medial fil-
ament, flaments branching fanlike horizon-
tally (fig. 6H in Wygodzinsky and Coscaron
1973). Guatemala, Mexico (Chiapas)
pachecolunai (Leon)
ACKNOWLEDGMENTS
We thank C. R. Méndez and A. Solano
V., field collectors, and W. Gonzalez, lab-
oratory assistant, Centro de Investigacion y
Diagnostico en Parasitologia, Universidad
de Costa Rica, for field and technical sup-
port during our investigations. Financial and
logistic support was provided for our studies
by the Vicerrectoria de Investigacion, Uni-
versidad de Costa Rica; and the Interna-
tional Committee of Migrations (CIM) pro-
vided financial support to J. Ramirez-Pérez
for travel to Costa Rica. We are grateful to
P. Malikul, laboratory technician, System-
atic Entomology Laboratory, for help in the
preparation of the plates that accompany
this paper. We thank P. H. Adler, Depart-
ment of Entomology, Clemson University,
Clemson, S.C.; W. N. Mathis, Department
of Entomology, Smithsonian Institution,
Washington, D.C.; and P. M. Marsh, N. E.
Woodley, and F. C. Thompson, Systematic
Entomology Laboratory, ARS, USDA, who
kindly read our manuscript and made help-
ful comments.
LITERATURE CITED
Coleman, R. W. 1953. A new blackfly species from
California (Diptera, Simuliidae). Proc. Entomol.
Soc. Wash. 55: 45-46.
Currie, D.C. 1986. An annotated list of and keys to
the immature black flies of Alberta (Diptera: Sim-
uliidae). Entomol. Soc. Can. Mem. 134: 1-90.
Dalmat,H.T. 1949. New species of Simultidae (Dip-
tera) from Guatemala. I. Ann. Entomol. Soc. Am.
42: 538-553.
1955. The black flies (Diptera, Simultidae)
of Guatemala and their role as vectors of oncho-
cerciasis. Smithsonian Misc. Colls. 125: 1-425, 44
Pls.
Diaz Najera, A. 1962. Claves para identificar las lar-
vas del género Cnephia y descripcion de dos nue-
vas especies. (Diptera: Simultidae). Rev. Inst. Sa-
lubr. Enferm. Trop. 22: 271-287.
1971. Descripcién de una nueva especie del
genero Cnephia del norte de Mexico. (Diptera:
Simuliidae). Rev. Invest. Salud Publica 31: 239-
247.
VOLUME 90, NUMBER 1
Petersen, J. L. 1985. Mayacnephia fortunensis (Dip-
tera: Simuliidae), a new black fly species from Pan-
ama. Proc. Entomol. Soc. Wash. 87: 80-84.
Peterson, B. V. 1981. Chapter 27, Simuliidae, pp.
355-391. In J. F. McAlpine, B. V. Peterson, G.
E. Shewell, H. J. Teskey, J. R. Vockeroth, and D.
M. Wood, eds., Manual of Nearctic Diptera, Vol.
1. Res. Br., Agr. Can. Monogr. 27.
75
Vargas, L.and A. Diaz Najera. 1948. Nuevas especies
de simulidos de Mexico y consideraciones diver-
sas sobre especies ya descritas. Rev. Inst. Salub.
Enferm. Trop. 9: 321-369.
Wygodzinsky, P. and S. Coscaron. 1973. A review
of the Mesoamerican and South American black
flies of the tribe Prosimulini (Simulinae, Simu-
liidae). Bull. Am. Mus. Nat. Hist. 151: 129-200.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 76-86
SIMULIUM (HEMICNETHA) HIEROGLYPHICUM
(DIPTERA: SIMULIIDAE), A NEW BLACK FLY SPECIES
FROM COSTA RICA
B. V. PETERSON, M. VARGAS V., AND J. RAMiREZ-PEREZ
(BVP) Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA,
% National Museum of Natural History, NHB-168, Washington, D.C., U.S.A. 20560;
(MVV) Centro de Investigacién y Diagnostico en Parasitologia, Universidad de Costa
Rica, Ciudad Universitaria Rodrigo Facio, Costa Rica; (JRP) Instituto de Biomedicina,
P.O. Box 4043, Caracas, Venezuela.
Abstract.—The female, male, pupa and larva of Simulium (Hemicnetha) hieroglyphi-
cum, new species, are described and illustrated. This species occurs in the highlands of
Costa Rica, and is most readily differentiated from all other described species of the
subgenus Hemicnetha by the large number of filaments in the respiratory organ (gill) and
the peculiar hieroglyphic-like markings on the thorax and frons of the pupa. A key to the
species of Hemicnetha known in the pupal stage is provided.
The subgenus Hemicnetha Enderlein
presently includes about 22 New World
species, mostly from the Neotropical Re-
gion. The number of species reported by
country is as follows: Mexico, 12; Guate-
mala, seven; Belize, one; El Salvador, one;
Costa Rica, two including the new species
described here; Panama, six; Colombia,
three; Venezuela, six; Trinidad and Tobago,
one; Guyana, one; Brazil, two; Bolivia, one;
and Argentina with two species (Pinto 1932,
Vargas and Diaz Najera 1951, 1957, Dal-
mat 1955, Vulcano 1967, Barreto 1969,
Field 1969, Ramirez-Pérez 1971, Ramirez-
Pérez and Vulcano 1973, Buenoetal. 1979).
Two of the species included in the above
listing, viz S. solarii Stone and S. virgatum
Coquillett, are known also from the Nearc-
tic Region (Stone 1948). Despite the num-
ber of species assigned to this subgenus, there
is no comprehensive study of the group.
Even so, pupae are known for most of the
described species, all of which differ from
the distinctive pupa of the new species de-
scribed below. This new species 1s described
to provide a name to use in work currently
being done on the black fly fauna of Costa
Rica. This is the second paper in a series
describing new species from Costa Rica; for
the first paper see Ramirez-Pérez, Peterson,
and Vargas (1988). A key to the pupae of
the described species of Hemicnetha, with
distributions and references to published
figures, is provided.
Simulium (Hemicnetha) hieroglyphicum,
New SPECIES
Figs. 1-21
Female (preserved in alcohol). — General
body color blackish brown. Length: body,
4.0 mm; wing, 4.08-4.5 mm.
Head: Lightly silvery pollinose. Frons
(Fig. 1) moderately broad, at vertex about
Y, wider than at narrowest point, distinctly
less than '2 as wide as head, and narrower
than long; covered with long, decumbent,
black pile. Clypeus concolorous or slightly
lighter than frons; slightly longer than wide;
VOLUME 90, NUMBER 1
covered with long, ventromedially directed,
black pile. Occiput silvery pollinose, dense-
ly covered with long, black pile; postocular
setae black, closely bending over eye mar-
gin. Antenna entirely dark brown to black,
with nine flagellomeres; pedicel slightly
longer than first flagellomere; fine pubes-
cence black. Mandible (Fig. 5) with 44-51
serrations. Blade of maxilla (Fig. 6) with 23-
28 retrorse teeth. Palpus (Fig. 3) with basal
two palpomeres and fifth palpomere slightly
lighter than palpomere three; palpomere five
slightly more than twice as long as palpo-
mere three; all palpomeres with black setae.
Sensory vesicle (Fig. 4) about '2 as long as
its segment, proximally situated, neck ab-
sent or very short with an enlarged, ovoid
mouth. Median proximal space of cibarium
shallow, broadly U-shaped, and with about
45 minute setulae with rounded bases in
membrane medially; dorsolateral arms
short, rather broad, sclerotized, inner sur-
faces of arms with numerous, minute, sen-
sory setulae.
Thorax: Postpronotum small; slightly
paler than scutum especially along adjoin-
ing margin; covered with long, recumbent,
golden yellow pile interspersed with some
semi-erect to erect, black setae. Scutum with
lateral margins narrowly more pale brown-
ish and with a silvery pollinose border ex-
tending around margins, posterior declivity
broadly silvery pollinose; anterolateral cor-
ners of scutum without distinct silvery spots,
but in posterior view, with three slender,
dark vittae that extend anteriorly from pos-
terior declivity, median stripe longest, lat-
eral stripes extending about '/ distance to
anterior margin; scutum densely covered
with short, recumbent, golden yellow setae
grouped in small clusters, pile longer along
anterior and lateral margins and still longer
posteromedially; anterior margin of scutum
with a number of long, dark, erect setae, and
posterior declivity with more numerous,
longer, dark setae. Scutellum yellowish
brown; densely covered with short, ap-
pressed, golden yellow setae, and numerous
Ife
long, black setae. Postnotum with dense sil-
very pollinosity. Pleuron brownish black
anteriorly, densely silvery pollinose, be-
coming paler yellowish brown medially and
posteriorly, and often mottled with dark
areas; anepisternal membrane brownish
yellow, often mottled; mesepimeral tuft of
long, black setae. Wing (Fig. 2) membrane
hyaline but with a definite brownish tinge;
veins brown. Base of costa, stem vein, and
remaining veins with black pile; Sc with nu-
merous setae ventrally except for about api-
cal 4, which is bare; R, with both setae and
spinules dorsally; R,,; setose ventrally; stem
of wing just basal to MA lightly sclerotized
with a conspicuous oval windowlike area
present; fringe of anal lobe and calypter with
black setae. Knob of halter yellowish white;
stem yellowish brown, with pale yellowish
setae. Legs with forecoxa, trochanter and
femur yellow to brownish yellow especially
on distal margins, these segments with short,
golden yellow scales plus longer black setae;
tibia mostly black with some yellow me-
dially, tarsus black, both tibia and tarsus
with black setae mixed with a few yellow
setae. Midcoxa, trochanter, basal 3 of fe-
mur, basal * of tibia, and about basal * of
basitarsus yellow, remaining portions of
midleg black; with mostly black setae. Hind
coxa brown; trochanter, basal 3 of femur,
about basal '2 of tibia, and basal *%4 to '2 of
basitarsus yellow, remainder of hindleg
black; with mostly black setae; hind basi-
tarsus swollen, about five times as long as
broad; calcipala prominent, stout, broadly
rounded apically, reaching to middle of
pedisulcus or slightly beyond; pedisulcus
moderately deep but not conspicuous. Claw
evenly curving from base, with a small but
conspicuous subbasal tooth.
Abdomen: Brownish black dorsally, be-
coming paler laterally and ventrally; basal
scale (tergite one) with fringe of long, pale
yellow pile; tergites blackish brown, with
darker hind margins, covered with short,
dark setae; tergite 10 small, subrectangular,
wider than long. Pleural membrane paler
78
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
VOLUME 90, NUMBER |
brownish yellow, with dark setae. Sternites
heavily sclerotized; sternites 2-7 subequal
in length, sparsely covered with dark setae;
posterior margin of sternite eight with long,
black setae. Terminalia as in Figs. 9-11. Anal
lobe (Fig. 10) broad, subrectangular, ante-
rior margin slightly concave medially, hind
margin nearly straight or slightly bilobed,
ventral margin nearly straight, not pro-
duced beneath cercus, densely setose. Cer-
cus (Fig. 10) rather small, subrectangular,
about two times as high as long, hind margin
broadly rounded. Hypogynial valves elon-
gate, 4 longer than greatest basal width, not
reaching hind margins of cerci; outer margin
of each valve rather deeply concave, inner
margin broadly rounded, apex rounded, with
a curved, longitudinal ridge that runs from
near apex to near inner basal margin, inner
“4 moderately sclerotized and bare, remain-
ing area densely microsetose. Stem of gen-
ital fork (sternite nine) (Fig. 9) long, heavily
sclerotized, twice as long as arms; each arm
short, expanding into a large, subrectangu-
lar plate with outer lateral margin thickened
ridgelike and inner proximal corner thick-
ened as a short, rounded process; arms
broadly attached to segment nine. Sper-
matheca (Fig. 11) globular, heavily sclero-
tized, without a pattern, but with a small
circular membranous area at junction with
spermathecal duct; inner surface of sper-
matheca rather evenly covered with nu-
merous, but well separated, minute spic-
ules.
Male.— General body color velvety brown
to black. Length: body, 4.6-6.0 mm, wing,
3.9-4.2 mm.
Head: Frons and clypeus densely silvery
pollinose, clypeus with erect, black pile. Oc-
ciput densely covered with long, black setae.
Antenna with scape, pedicel and extreme
a9
base of first flagellomere yellow, rest of fla-
gellum black; first flagellomere nearly twice
as long as pedicel; scape with tuft of dark
setae that are much longer than sparse setae
of pedicel especially those of distal margin,
fine pubescence of flagellomeres pale yel-
low, longer setae black. Palpus entirely dark
brown to black, with black pile; palpomere
four slightly but distinctly longer than three,
palpomere five about three times as long as
palpomere three. Sensory vesicle small,
about '4 as long as its segment; neck distinct,
enlarging to form a round mouth.
Thorax: Postpronotum small, yellow, with
short, recumbent, golden yellow pile inter-
spersed with a few more erect black setae.
Scutum brownish black, margins, especially
laterally and on notopleuron, slightly paler
brown, narrowly silvery pollinose along an-
terior margin but covering most of posterior
declivity; densely covered with short, re-
cumbent, golden yellow pile grouped into
small clusters, pile slightly longer laterally
and posteromedially, posterior declivity
with some long, erect, black setae. Scutel-
lum yellow, densely covered with moder-
ately long, semi-erect, golden yellow setae
interspersed with some long, erect, black se-
tae. Extreme anterior margin of postnotum
concolorous with scutellum, remainder
brownish black, densely silvery pollinose.
Pleuron dark blackish brown and densely
silvery pollinose anteriorly, becoming paler
yellowish brown medially and darker brown
posteriorly; anepisternal membrane brown-
ish yellow; mesepimeral tuft of long, black
setae. Wing membrane hyaline but with a
faint yellowish tinge; veins yellowish brown;
base of costa, stem vein, and remaining veins
with black setae; Sc with about 12 setae at
extreme base ventrally; R, with both setae
and spinules dorsally; R,,; setose ventrally;
_—
Figs. 1-11.
Simulium (H.) hieroglyphicum. Figs. 1-6, female. 1, Front view of frons and ocular notches. 2,
Portion of wing showing setation. 3, Maxillary palpus. 4, Enlarged view of sensory organ of third palpomere.
5, Tip of mandible showing serrations. 6, Blade of maxilla showing retrorse teeth. Figs. 7-8, male. 7, Gonocoxite
and gonostylus (dorsal (inner) surface). 8, Ventral plate of aedeagus, lateral view. Figs. 9-11, female. 9, Genital
fork (sternite 9). 10, Anal lobe and cercus. 11, Spermatheca.
80
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
a)
»
2
z
x
N
x
a
z
VOLUME 90, NUMBER |
stem of wing just basal to MA lightly scler-
otized with a conspicuous oval windowlike
area present; fringe of anal lobe and calypter
brownish yellow, setae often with black bas-
es. Knob of halter white, stem brownish yel-
low with pale yellow pile. Fore coxa, tro-
chanter and femur yellow tinged with brown,
especially on distal margins, tibia and tarsus
black; coxa and trochanter anteriorly with
yellow setae plus scattered black setae; tip
of femur and anterior surface of tibia with
appressed, golden yellow scales, rest of setae
black. Mid- and hind coxae dark brown; tro-
chanters dark brown mottled with patches
of yellow; mid- and hind femora brownish
yellow with dark brown apices; tibiae large-
ly dark brown on anterior face but bases
and inner surfaces yellow; about basal 2 of
first two tarsal segments yellow, their apices
dark brown, remaining portions and rest of
tarsomeres dark brown; anterior surfaces of
mid- and hind femora, tibiae and basitarsi
with appressed, golden yellow scales, rest of
setae black; hind basitarsus swollen, about
three times as long as broad; calcipala prom-
inent, broadly rounded, nearly obscuring
pedisulcus; pedisulcus deep. Claw short,
slender, with a short but conspicuous sub-
basal tooth.
Abdomen: Brownish black dorsally, be-
coming brownish yellow laterally and ven-
trally; basal scale with fringe of long, dark
setae black at base and distally paler brown-
ish to yellowish; tergites broad, all nearly
uniform in width, velvety in texture and
with black hind margins, tergites 2-7 each
with an oblique, silvery, pollinose patch lat-
erally, covered with short, black setae; ter-
gite 10 small, rectangular, longer than broad.
Pleural membrane of segments 3-6 with a
dorsal patch of long, black setae just lateral
to margin of respective tergites, segment
seven with a similar but smaller patch. Ster-
81
nites 3-8 nearly uniform in width, dark
brown on about basal *4, paler yellowish on
about distal ', with black setae. Terminalia
as in Figs. 7-8. Gonocoxite (Fig. 7) subrec-
tangular, about 3 wider than long, moder-
ately setose on about distal /2. Gonostylus
long, slightly more than three times as long
as greatest width at base; margins strongly
sinuous, inner margin with distinct bulge at
midlength, apical margin rounded, with a
single terminal spine. Ventral plate of ae-
deagus (Fig. 8) with a prominent, broadly
rounded ventral lip which, in lateral view,
resembles a medieval broadhead ax; in ven-
tral view, body slightly wider than long, lat-
eral and apical margins broadly rounded;
basal arms rather slender and_ straight,
obliquely directed outwardly with tips rath-
er strongly curving inwardly. Median scler-
ite of aedeagus with a slender, straight stem
that branches into two broad arms nearly
as long as stem, arms tapering distally, nar-
rowly rounded to slightly pointed. Plate of
endoparameral organ moderately large,
subrectangular, moderately sclerotized; arm
only slightly longer than basal plate, twist-
ing, with a series of corrugations and rather
poorly defined teeth on outer margin, apical
five or six teeth short, stout and better de-
fined than more proximal teeth; aedeagal
membrane densely covered with minute
spinules arranged in rather regular comblike
series of about three to 12 spinules per comb.
Pupa.—Length 6.3-7.0 mm. Respiratory
organ (gill) (Fig. 12) 3.0 mm long, often
reaching hind margin of thorax; consisting
of a short, rather broad base, covered with
minute spicules, and with one thick basal
branch projecting posteriorly over thorax
and dividing apically into two or more slen-
der filaments, and with two similar, thick,
closely appressed basal branches, one stout-
er than the other, which project anteriorly
—
Figs. 12-21.
Simulium (H.) hieroglyphicum. Figs. 12-15, pupa. 12, Respiratory organ (gill) (anterior branch
to the left). 13, Cocoon. 14, Portion of frons showing integumental pattern. 15, Abdomen showing chaetotaxy
on dorsal (d) and ventral (v) surfaces. Figs. 16-21, larva. 16, Inner distal and subapical margins of mandible
showing dentation. 17, Antenna. 18, Anal sclerite. 19, Anal papillae. 20, Hypostomal cleft. 21, Hypostoma.
82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
over head and apparently not dividing into
finer filaments apically nor giving rise to
other filaments along their dorsal surfaces;
dorsal surface of main posterior branch giv-
ing rise to about 60-90 relatively short, slen-
der, pale grayish white filaments, some of
which are simple, and some dividing at
varying distances from their bases into two
or more apical filaments. Frons (Fig. 14)
with a series of strong, rugosities giving rise
to short, spurlike projections; without other
granulations; antenna of both female and
male reaching only about 2 distance to hind
margin of head: a single, short, fine seta
present medial to base of each antenna.
Dorsum of thorax with strong rugosities,
some with short, spurlike projections; these
rugosities arranged in a loose reticulate pat-
tern, without other granules dorsally; two
long, simple, dorsal trichomes present on
each side of thorax. Chaetotaxy of each lat-
eral half of tergites as follows (Fig. 15): ter-
gite one with two fine setae and a closely
associated lateral patch of minute spinules;
tergite two with three fine setae, and a me-
dian patch of minute spinules; tergite three
with three small hooklets, three fine setae,
and a patch of minute spinules anterolateral
to hooklets, and a similar but smaller patch
just posterior to hooklets; tergites four and
five each with four stout, anteriorly directed
hooks near hind margin, 3-4 fine setae, and
an anterolateral patch of minute spinules;
tergites 6-8 without fine setae but with a
small anterolateral patch of minute spi-
nules; tergite nine bare, caudal spines ab-
sent. Chaetotaxy of each lateral half of ster-
nites as follows: sternites 1-3 bare; sternites
4-5 each with an anterolateral patch of mi-
nute spinules; sternites 6-7 each with two
anteriorly directed hooks, and sternite eight
with one similar hook, all three of these
sternites with a patch of minute spinules just
anterior to hooks; sternite nine with an an-
terior patch of minute spinules across its
width. Cocoon (Fig. 13) boot-shaped,
densely woven, with floor extending ante-
riorly about '2 length of flat bottom portion
of cocoon; in lateral view, anterior collar of
cocoon relatively short, slanting anterodor-
sally, with one or two small festoons or win-
dowlike openings near anterolateral margin,
which, however, are easily broken off and
lost.
Larva.—Length 13.0-13.5 mm. Body
gradually expanding posteriorly; color gray-
ish dorsally, lighter and more yellowish ven-
trally; intersegmental lines rather broad,
slightly lighter than rest of abdomen dor-
sally. Head capsule brown, darker laterally
and ventrally; head spots darker than rest
of frontoclypeal apotome, posteromedian
spot broad basally, strongly tapering distal-
ly, well separated from anteromedian spot,
this paler and not as distinct; first antero-
lateral spot not discernible, second antero-
lateral spot large, distinct, well separated
from first posterolateral spot, this smaller
and less distinct; second posterolateral spot
large but pale and diffuse; eye spots large.
Antenna (Fig. 17) slightly longer than stalk
of labral fan; proportions of segments (basal
to apical) 1:2:0.68; dorsal half of basal two
antennomeres yellowish brown, ventral half
transparent, distal antennomere entirely
yellowish brown. Labral fan with 52-59 (av.
55) primary rays. Hypostoma as in Fig. 21;
median tooth short but longer than others;
lateral teeth small, subequal in length but
distal margin convex; lateral margins of hy-
postoma with 1-3 variable but small, weak
serrations; 11-13 hypostomal setae along
each margin and with 1-3 much smaller,
more medial setae near hind margin. Hy-
postomal cleft (Fig. 20) moderately deep,
extending about 7 distance to base of hy-
postoma, a narrow inverted V-shape. Hy-
postomal bridge slightly but distinctly long-
er than hypostoma. Mandible (Fig. 16) with
3-5 apical teeth, 5-8 preapical teeth, and
inner subapical ridge with one fine but
prominent tooth. Maxillary palpus 2.5 times
as long as width at base. Lateral plate of
proleg short and broad, heavily sclerotized,
extending about /4 length of apical segment;
circlet of apical hooks in about 80 rows of
about 20 hooks each. Rectal setulae minute;
anal papillae (Fig. 19) complex, arranged in
VOLUME 90, NUMBER 1
three main groups of about 22-24-41 short,
digitiform papillae. Anal sclerite (Fig. 18)
heavily sclerotized, arms slender, narrowly
joined; anterodorsal arms about 7 as long
as posteroventral arms, anterodorsal arms
terminating in slightly enlarged, subquad-
rate to subrectangular, lightly sclerotized
plates. Posterior circlet of hooks consisting
of 50-55 hooks in 500-550 rows.
Types.— Holotype, 4 (reared with asso-
ciated pupal pelt, all preserved in alcohol),
stream (#34), upstream from bridge, Rio
Poasito, Canton Pods, Provincia Alajuela,
Costa Rica, November 4, 1986, A. Solano
V. and W. Gonzalez. The stream 1s located
23.5 km from Carrizal on the road to Poas
Volcano.
Paratypes.—1 ¢ (mounted on 5 slides),
same data except January 31, 1970; 1 2
(mounted on six slides) (reared with asso-
ciated pupal pelt mounted on two slides),
same data except July 18, 1986; 1 2 (mount-
ed on five slides), 2 64 (one male mounted
on four slides and one mounted on three
slides) (all reared with associated pupal pelts
each mounted on two slides), same data ex-
cept August 8, 1986; | @ (reared) (pinned),
1 4(pinned) and 2 44 (in alcohol), same data
except August-September 1986; 1 pupa,
same data except August 9, 1968: 19 pupae,
same data except January 1, 1970; 10 pu-
pae, same data except March 20, 1970; 5
pupae, same data except August-September
1986; 12 larvae, same data except January
31, 1970; 3 larvae, same data except March
20, 1970; 3 larvae, same data except Sep-
tember 5, 1986 (one larva mounted on six
slides; two mounted on five slides each); 58
larvae, same data except August-September
1986.
Holotype deposited in the collection of
the U.S. National Museum of Natural His-
tory, Washington, D.C. Paratypes are de-
posited in the U.S. National Museum of
Natural History, and the entomology col-
lection of the Department of Parasitology,
University of Costa Rica.
Etymology.—The specific name 1s the
singular, neuter form of the Latin adjective
83
hieroglyphicus, and refers to the hieroglyph-
ic-like markings on the dorsum of the head
capsule and thorax of the pupa.
Biological notes.—All available speci-
mens of Simulium (Hemicnetha) /iero-
glyphicum came from the same stream (#34)
of the type locality. The stream arises from
nearby slopes, which are covered with abun-
dant vegetation and large trees that shade
the area, and then passes through pasture
land at about 1940 m in elevation. The
stream is about 3 m wide, 0.3 m in depth
and has a moderate to fast cascading flow
over large boulders and smaller stones. A
small amount of trailing vegetation, mostly
torch ginger (Nicolaia elatior (Jack) Horan),
occurs along the stream banks. There 1s no
emergent vegetation in this portion of the
stream. The water 1s unpolluted and ranges
in temperature from 13 to 14°C. Larvae and
pupae were found on both rocks and trailing
vegetation. Adults were neither attracted to
nor taken biting humans.
Remarks.—There is not full agreement as
to the subgeneric assignments of a number
of species of black flies. This is especially
true for the Neotropical species, and, in fact,
Vulcano (1967) in her catalog of the black
fly species of the Americas south of the
United States, did not assign species to any
subgenera. Based on larval characters and
those of the male and female terminalia, as
described by Stone (1963), we have selected
for inclusion in the key to pupae presented
below those species that seemed to us to
belong to Hemicnetha; even so, there might
be some eventual reassignments as the Neo-
tropical species are more thoroughly stud-
ied and become better known in all their
life history stages.
There currently are no keys that include
all the adult and immature stages of all the
described species of Hemicnetha. Adults of
S. hieroglyphicum do not satisfactorily run
to any species in the keys of Dalmat (1955)
or Vargas and Diaz Najera (1957). Even
though they seem to be most similar to S.
smarti Vargas they differ in many more
characters than there are similarities. Fe-
84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
males of S. hieroglyphicum can be distin-
guished from those of all other known
species by the Sc being setose ventrally ex-
cept for about apical 5 which is bare, these
setae placed in a single row except at the
extreme base (four to five setae long) in
which they are placed in a double row; an-
terior marginal area of scutum with nu-
merous clusters of 2-3 golden yellow scale-
like setae; the three dark, slender vittae that
are visible in posterior view; and by the
shape and form of the various genital struc-
tures (see Figs. 9-11). The males can be dis-
tinguished from all other known males of
this group by the following combination of
characters: Sc bare except for about 12 setae
at extreme base; scutum without distinct
dark vittae, but clothed with numerous
groups of golden yellow, scalelike setae;
broad hind basitarsus; and the distinctive
form of the genital structures (see Figs. 7—
8).
The respiratory organ, or gill, of the pupa
of S. hieroglyphicum is distinctly different
from those of all other known species as
demonstrated by the following key. Two
species, viz muiscorum Bueno, Moncada,
and Munoz de Hoyos, and keenani Field,
are included in the key on the basis of pub-
lished descriptions as we do not have ma-
terial of these species to check for distin-
guishing characters. Although we have
specimens of pu/verulentum Knab we lack
material of S. guerrerense Vargas and Diaz
Najera and so have followed the key of Var-
gas and Diaz Najera (1957) to separate the
pupae of these two species. Consequently,
the key is not complete but it should be
useful for separating the pupae of the ma-
jority of the species included. In the liter-
ature there are only a few other keys to pu-
pae that contain two or more of the species
that we include in Hemicnetha, and these
are included under varying names and in
varying combinations of species. None of
these keys include all the species that are in
the key given below. The pupa of S. (H.)
dehnei Field, from Panama, is not known
and so does not appear in the key. We are
unable, at this time, to prepare a reliable
key to the other stages of the species of
Hemicnetha because of the lack of adequate
material.
Key to NEw WorLD SPECIES
SIMULIUM (HEMICNETHA)
PUPAE
1. Respiratory organ (gill) with six filaments .. 2
— Respiratory organ with eight or more fila-
ments .
. Two of the six filaments much shorter than
the other four. Granulations on dorsum of
thorax with spinules (Venezuela; fig. 30A, B,
J, P in Ramirez-Pérez 1983)
Pate . oviedoi Ramirez-Perez
— All filaments subequal in length. Granula-
tions on dorsum of thorax disk- or buttonlike
and without spinules .
3. Cocoon shorter and higher than in following
species, anterolateral margin of cocoon nearly
straight and set at almost a right-angle to an-
teroventral surface of collar (Brazil; figs. 23-
25 in Pinto 1932)
te
anterolateral margin of cocoon sloping and
concave, anteroventral portion of cocoon pro-
jecting liplike (Venezuela; fig. 42D, E, K in
Ramirez-Pérez 1983) ..... rivasi Ramirez-Peérez
4. Respiratory organ with eight filaments .... 5
— Respiratory organ with ten or more filaments
: Aone neereogot SOT BO ee ntroo nc & 9
5. Cocoon boot-shaped, anterodorsal margin
with well-developed festoons, these some-
times broken but traces of them usually re-
LAIN yon oprte Sak Cee eee reel 6
— Cocoon boot-shaped but anterodorsal margin
simple, or with poorly-defined festoons .... 9
6. Frons almost totally covered by small spi-
nules; thorax with small spinules ventral to
base of respiratory organ. Caudal spines pres-
ent but small (Colombia, Guyana, Brazil, Ar-
gentina; figs. 33-41 in Vulcano 1958)
SE Peal OSE re Se oes oe rubrithorax Lutz
— Frons smooth, with, at most, a few granula-
tions but without spinules; thorax without
spinules. Caudal spines absent ............ 7
7. Filaments arising from stem in two distinct
groups of four filaments each (U.S.A., Mex-
ico, Guatemala, Panama; figs. 7, 9 in Stone
1948; and 263, 270 in Vargas and Diaz Najera
OS) sas ae see cea ak virgatum Coquillett
— Filaments all arising at about the same level
from a common stem
VOLUME 90, NUMBER |
8.
Filaments slender, about 0.12 mm in width
at base; thorax less densely granulose than in
following species (Venezuela; figs. 1-4 in Ra-
mirez-Pérez and Vulcano 1973; and 10B, O,
P in Ramirez-Pérez 1983) .
Nactamenec ee. conviti Ramirez-Perez and Vulcano
Filaments broader, about 0.60 mm in width
at base; thorax more densely granulose than
in above species (Mexico, Guatemala, Costa
Rica, Panama, Venezuela; figs. 235, 237 in
Vargas and Diaz Najera 1957) ... paynei Vargas
Anterodorsal margin of cocoon with poorly
developed festoons. Thorax strongly rugose
in a somewhat reticulate pattern. All filaments
petiolate, branching some distance from base
(Mexico; figs. 181, 185, 186 in Vargas and
Diaz Najera 1957) F
Rar ee bricenoi Vargas, Martinez Palacios,
and Diaz Najera
Anterodorsal margin of cocoon without fes-
toons. Thorax granulose but without reticu-
late rugosities. All filaments branching close
together at or near the base (Mexico; figs. 198,
204 in Vargas and Diaz Najera 1957)
freemani Vargas and Diaz Najera
. Respiratory organ with ten filaments ...... 11
Respiratory organ with 12 or more filaments
. Thorax densely covered with black spinules
(Mexico; figs. 213, 216,
Diaz Najera 1957)
hinmani Vargas, Martinez Palacios
and Diaz Najera
Thorax may be granulose but without spi-
MULES nas odace aos i etawets os eel:
21 in Vargas and
. Anterior collar of cocoon raised fell Brave
level of dorsum of pupal thorax; anterior face
of collar almost vertical, its dorsolateral mar-
gins curving posteriorly downward in undu-
lating fashion (Mexico, Guatemala; figs. 273,
276 in Vargas and Diaz Najera 1957)
yepocapense Dalmat
Pontenen collar of cocoon only raised slightly
above level of dorsum of pupal thorax; an-
terior face of collar distinctly oblique, its dor-
solateral margins not undulating downward
posteriorly, but simple and at least slightly
directed anteroventrally ............ 13
. Anterodorsal margin of cocoon obliquely
slanted anteroventrally. Respiratory organ
with filaments somewhat longer and more 1so-
lated from each other (Mexico; figs. 210, 211
in Vargas and Diaz Najera 1957)
guerrerense Vargas and Diaz Najera
Anterodorsal margin of cocoon more hori-
zontal, only slightly slanting anteroventrally.
Respiratory organ with filaments somewhat
85
shorter and more closely clumped [no char-
acters are known that reliably separate pupae
of this species from pupae of the above species]
(Mexico, Guatemala, Belize, El Salvador,
Panama, Venezuela; figs. 243, 246 in Vargas
and Diaz Najera 1957) pulverulentum Knab
. Respiratory organ with 12 filaments 15
Respiratory organ with 16 or more filaments
Peseeeees 17
; reco ceniral margin (lip) Ohcacoon deeply
notched medially so that lateral portions are
produced anteriorly as two subtriangular or
spatulate processes (Mexico, Guatemala,
Panama, Trinidad and Tobago, Venezuela,
Colombia, Bolivia; figs. 226, 230 in Vargas
and Diaz Najera 1957) mexicanum Bellardi
Anteroventral margin (lip) of cocoon simple,
without a medial notch and without subtrian-
gular or spatulate processes 16
. Respiratory organ with relatively short fila-
ments branching antlerlike; filaments without
minute tubercles. Thorax without trichomes.
Abdomen with two strong caudal spines (Ar-
gentina; fig. 6A-I in Wygodzinsky 1949)
NE facets oe lahillei Paterson and Shannon
Recpretony, organ with relatively short fila-
ments but these not branching antlerlike; fil-
aments covered by minute tubercles. Thorax
with branched trichomes. Abdomen without
caudal spines (Colombia; figs. 22-26 in Bueno
etal. 1979) ...... muiscorum Bueno, Moncada,
and Munoz de Hoyos
. Respiratory organ with 15-16 filaments 18
Respiratory organ with 18 or more filaments
oe ee 20
; Anterior collar Bre cocoon ae well above
level of dorsum of pupal thorax; dorsolateral
margins of opening of cocoon curving pos-
teriorly downward in undulating fashion.
Respiratory organ with 16 filaments (Mexico,
Guatemala; figs. 189, 195 in Vargas and Diaz
Najera 1957)" neg gents earlei Vargas, Martinez,
Palacios, and Diaz Najera
Anterior collar of cocoon only raised slightly
above dorsum of pupal thorax; dorsolateral
margins of opening of cocoon not curving
posteriorly downward in undulating fashion.
Respiratory organ with 15-16 filaments ... 19
. Respiratory organ usually with 15 filaments,
14 in pairs and | single filament. Dorsum of
thorax smooth. Anterior collar of cocoon short,
raised slightly above dorsum of pupal thorax,
dorsolateral margins of opening of cocoon
straight or slightly concave; (U.S.A., Mexico;
figs. 8, 10 in Stone 1948; figs. 255, 261 in
Vargas and Diaz Najera 1957) .... solarti Stone
Respiratory organ with 16 filaments variously
86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
branching near base, some filaments branch-
ing in groups of three, some in pairs, and at
least three singles. Anterior collar of cocoon
longer (Panama; fig. 8 in Field 1969) ..
and thorax without hieroglyphic-like rugosi-
ties (Mexico, Guatemala; figs. 252, 254 in
Vargas and Diaz Najera 1957) ... smarti Vargas
— Respiratory organ with two main branches, a
posterior branch giving rise to 60-90 short,
white filaments, and an anterior branch that
divides into two filaments apically. Head and
thorax with hieroglyphic-like rugosities (Cos-
ta Rica, fig. 11 herein) . . ay : Le
Re eae a hieroglyphicum Peterson, Vargas,
and Ramirez-Peérez
ACKNOWLEDGMENTS
We thank R. Echeverri, W. Gonzalez, and
the Centro de Investigacion y Diagnostico
en Parasitologia, Universidad de Costa Rica,
for field and technical support in this in-
vestigation. Financial and logistic support
was provided for our studies by the Vicer-
rectoria de Investigacion, Universidad de
Costa Rica; and the International Commit-
tee of Migrations (CIM) provided financial
support to J. Ramirez-Pérez for travel to
Costa Rica. We are grateful to P. Malikul,
laboratory technician, Systematic Ento-
mology Laboratory, for help in the prepa-
ration of the plates that accompany this
paper. We thank J. W. Amrine, Jr., De-
partment of Entomology, West Virginia
University, Morgantown, W.V., W. N.
Mathis, Department of Entomology,
Smithsonian Institution, Washington, D.C.,
and D. C. Ferguson, A. L. Norrbom and F.
C. Thompson, Systematic Entomology
Laboratory, ARS, USDA, for reading and
commenting on the manuscript.
LITERATURE CITED
Barreto, P. 1969. The species of black flies found in
Colombia (Diptera: Simulidae). J. N.Y. Entomol.
Soc. 77: 31-35.
Bueno, M. L., L. I. Moncada, and P. Munoz de Hoyos.
1979. Simuliidae (Insecta: Diptera) de Colombia.
I. Nueva especie de Simulium (Hemicnetha). Cal-
dasia 12: 581-594.
Dalmat, H. T. 1955. The black flies (Diptera, Sim-
uliidae) of Guatemala and their role as vectors of
onchocerciasis. Smithsonian Misc. Colls. 125: 1-
425, 44 Pls.
Field, G. 1969. Studies of black flies of Panama. III.
Two new species of Simulium of the subgenus
Hemicnetha. Ann. Entomol. Soc. Am. 62: 157-
163.
Pinto, C. 1932. Simulidae da America Central e do
Sul (Diptera). 77 Reun. Soc. Argent. Patol. Reg.
Norte 60: 661-763. 1931.
Ramirez-Pérez, J. 1971. Distribucion geografica y
revision taxonomica de los simulidos (Diptera:
Nematocera) de Venezuela con descripcion de diez
especies nuevas. Acta Biol. Venez. 7: 271-371.
1983. “Los jejenes de Venezuela.” Simposio
de Oncocercosis Americana, Caicet, Puerto Ay-
acucho, 15-17 Octubre 1983; 156 pp.
Ramirez-Pérez, J., B. V. Peterson, and M. Vargas V.
1988. Mayacnephia salasi (Diptera: Simultidae),
a new black fly species from Costa Rica. Proc.
Entomol. Soc. Wash. 90: 66-75.
Ramirez-Pérez, J. and M. A. Vulcano. 1973. De-
scripcion y redescripciones de algunos simulidos
de Venezuela (Diptera: Simuliidae). Arch. Venez.
Med. Trop. Parasitol. Med. 5: 375-399.
Stone, A. 1948. Simulium virgatum Coquillett and a
new related species (Diptera: Simuliidae). Jour.
Wash. Acad. Sci. 38: 399-404.
. 1963. An annotated list of genus-group names
in the Family Simuliidae (Diptera). Agr. Res. Serv.,
U.S. Dept. Agr., Tech. Bull. 1284: 1-28.
Vargas, L. and A. Diaz Najera. 1951. Notas sobre
sistematica y morfologia de simulidos. Rev. Soc.
Mex. Hist. Nat. 12: 123-172, 17 Pls.
1957. Simulidos Mexicanos. Rev. Inst. Sa-
lub. Enfer. Trop. 17: 143-399.
Vulcano, M. A. 1958. Redescrigéo do Simulium ru-
brithorax Lutz, 1909, e descrigao do alotipo 4
(Diptera, Simuliidae). Papéis Avulsos, Dept. Zool.,
Sect. Agric., S40 Paulo 13: 227-240.
. 1967. 16 Family Simuliidae, pp. 1-44. Jn N.
Papavero, ed., A Catalogue of the Diptera of the
Americas South of the United States. Dept. Zool.,
Sect. Agric., Sao Paulo.
Wygodzinsky, P. 1949. Contribuciones al conoci-
miento de los Simuliidae Argentinos. I. Introduc-
cin. Redescripcion de Simulium lahillei Paterson
y Shannon, 1927 (Diptera). Anal. Inst. Med. Reg.,
Univ. Nacional de Tucuman 2: 303-319.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 87-90
BELASPIDIA LONGICAUDA, NEW SPECIES, THE FIRST NEARCTIC
BELASPIDIA (HYMENOPTERA: CHALCIDIDAE)
J. A. HALSTEAD
Department of Biology, California State University Fresno, Fresno, California 93740;
present address: 2110 N. Hayes, Fresno, California 93722.
Abstract. —Belaspidia longicauda, new species, is described from specimens collected
in California. Diagnostic characters for the female and male are illustrated. Belaspidia
longicauda is incorporated into existing keys to world species of Belaspidia. The genus
Belaspidia is recorded for the first time in the Nearctic Region; three other species occur
in the Palearctic Region.
Prior to this paper, the genus Belaspidia
contained only three species, all Palearctic.
While examining material for a survey of
Chalcididae in California, I found speci-
mens of a Nearctic Belaspidia. Further study
showed this to be a new species, represent-
ing the first record of the genus Belaspidia
in the Nearctic Region and the fourth species
in the genus. I take this opportunity to de-
scribe this species as it greatly contributes
to the knowledge of the zoogeography and
systematics of the family Chalcididae. Host
information is known only for Belaspidia
obscura Masi. This species is a pupal para-
sitoid of Aperona crenulella Brd. (Lepidop-
tera: Psychidae) (Boucek 1951a, Nikolskaya
1952).
Belaspidia may be distinguished from
other Chalcididae in North America by the
following characters: Hindtibia truncate
distally, two hindtibial spurs present (Hal-
tichellinae); marginal vein on anterior mar-
gin of wing, postmarginal and stigmal veins
present, distinct (Haltichellini); scutellum
with a median tooth on posterior margin,
postmarginal vein 1.7 x as long as marginal
vein, and tergite | without longitudinal ca-
rinae, coloration never green metallic.
Belaspidia longicauda Halstead,
New SPECIES
Figs. 1-9
Holotype female.— Length 4.2 mm. Black,
with orange markings. The following areas
orange: mandibles, joint between scape and
frons, joint between femora and tibiae of
fore and middle leg, apex of tibiae, tarsi
except for brown last tarsomere and claws,
sternites, hypopygidium, ventral and latero-
posterior margins of tergites, ventral margin
of ovipositor sheath where bordered by epi-
pygidium, marginal vein of forewing, and
venation of hindwing. Compound eyes and
pubescence silver, ocelli tan.
Eyes with short pubescence. Anterior
ocellus round, separated from lateral ocelli
by 1.7 its diameter. Lateral ocelli oval,
separated from margin of compound eye by
0.8 x their diameter. Labrum oval, slightly
longer than wide, sublaterally with a depres-
sion, transversely microridged, ventral
margin with a fringe of setae. Mandibles
with a few setae on outer margin, left with
2 teeth, right with 3 teeth. Differentiation
between clypeus and frons vague. Insertion
of antennae "3 the distance from the base of
88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Qa
\
aN
\s
Figs. 1-9. Belaspidia longicauda. |. Head and an-
tenna of female. 2, Pronotum, dorsal view. 3, Thorax
of female. 4, Scutellum, dorsal view. 5, Abdomen of
female. 6, Hind leg of female, small teeth on ventral
margin omitted. 7, Venation (partial) of forewing. 8,
Antenna of male. 9, Abdomen of male. Scale lines 0.5
mm.
labrum to imaginary line between ventral
margin of compound eyes (Fig. 1). Scape
reaching almost to ventral margin of ante-
rior ocellus, widest at 3 its length. Pedicel
1.5x longer than wide. Annellus 1 and 2
short, wider than long. Flagellar segments
longer than wide. Flagellum and club longer
than height of head (lateral view). Scrobe
cavity shallowly depressed, coriaceous,
without pubescence. Area between anten-
nae flat, distance 1.5 antennal socket
width, projecting into scrobe cavity as a
raised triangle whose apex lies in the center
of frons mediad of ventral margin of com-
pound eyes. Frons at antennal insertions
slightly rounded anteriorly (Fig. 1). Occiput
medially emarginate for reception of ante-
rior projection of pronotum.
Pronotum with posterior margin broadly
emarginate, anterior medial margin slightly
projected anteriorly, 5 x as wide as long (Fig.
2). Thorax convex dorsally (Fig. 3), coria-
ceous laterally, dorsally with dense, shallow,
small punctures; moderately setose. Poste-
rior margin of scutellum with a median tri-
angular projection that is centrally concave
in dorsal view (Fig. 4). Mesopleural acetab-
ulum very shallow, almost flat. Tegula tri-
angular, ventral margin straight. Meta-
pleuron and propodeum laterally with
shallow punctures, moderately setose. Mes-
episternum coriaceous, without setae. Pro-
podeum with 2 strong, laterally arching sub-
median longitudinal carinae. Remainder of
propodeum with a low areolation of carinae,
integument coriaceous. Spiracle crescent
shaped, arched anteriorly, wider at each end.
Abdomen sessile, long, 1.7 x longer than
length of head and thorax, apex accuminate,
dorsal margin flat in lateral view (Fig. 5).
Tergites with a thin shiny band on margins.
T#1-4 dorsally punctate, remainder of ab-
domen except hypopygidium coriaceous.
Hypopygidium smooth, heavily chitinized
along median. Tergites (except T#1 and
T#2-3 dorsally), epipygidium (except basal
>), and ovipositor sheath with long dense
setae. T#1 submedially with a patch of se-
tae. Petiole 3 x as high as long, rectangular,
anterior margin with a carina, laterally with
a few vague longitudinal carinae.
Legs coriaceous, with short dense pubes-
cence. Hindcoxa large, slightly longer than
wide; its greatest width equal to that of
hindfemur, dorsally with a triangular tooth
which fits into posterior margin of meta-
pleuron (Fig. 6). Hindfemur narrowly ovoid,
2x as long as wide, ventral margin with an
acute tooth near middle, ventral margin with
many minute teeth from large tooth to apex
(Fig. 6). Hindtibia anteriorly with an inner
and outer carina extending to near apex.
Forewing extending posteriorly to near
apex of epipygidium. Forewing and hind-
wing clear, densely setose. Postmarginal vein
long, 1.7 x as long as marginal vein (Fig. 7).
Stigmal vein 4 as long as marginal vein.
Hindwing with 3 hamull.
Allotype male.—Length 3.8 mm. Similar
to female but differs in compound eye col-
VOLUME 90, NUMBER 1
oration being brown, shape of antennae (Fig.
8), and shape of abdomen (Fig. 9).
Variation.—Body length of the female
paratypes varies from 3.8 to 4.3 mm. The
length and width of the scape in females
varies slightly. One paratype with scape
slightly longer and thinner than holotype,
another with scape slightly shorter and wid-
er. Two paratypes with length of abdomen
shorter than holotype, 1.4 x as long as head
and thorax.
Specimens examined.—Holotype, 2°,
United States, California, Tulare Co., Ash
Mountain, Kaweah Powerstation #3, VIII-
19-1982, R. D. Haines, from Halstead col-
lection, deposited in National Museum of
Natural History, Washington, D.C.
(USNM). Allotype, 4, United States, Cali-
fornia, Los Angeles Co., Tanbark Flat, VI-
23-1950, on Eriogonum, from California
Insect Survey, University of California,
Berkeley (CIS) collection, deposited in
USNM. Paratypes.— California, Fresno Co.,
Coalinga, Coalinga Mineral Springs Road,
V-21-1982, R. F. Gill (1 2, from Halstead
collection to California Academy of Sci-
ences, San Francisco). Sacramento Co., 10
mi. NE Folsom, V-11-1960, on Eriodictyon
californicum, M.S. Wasbauer (1 2, Califor-
nia State Collection of Arthropods, Cali-
fornia Department of Food and Agriculture,
Sacramento). San Bernardino Co., Cronise
Valley, IV-29-1956, on Prosopis, M. S.
Wasbauer (1 2, CIS). San Bernardino Co.,
Sheep Creek, V-27-1973, E. M. and J. C.
Hall (1 2, University of California, River-
side (UCR)). Riverside Co., Gavilan, V-8,
12-1950, on Eriogonum fasciculatum, Tim-
berlake (2 2°, UCR).
Habitat.—The holotype was collected
from a five mile long hydroelectric flume
that runs through foothill woodland and
chamise chaparral habitats at an elevation
of 660 m (2200 ft). The paratype from Coa-
linga was collected in chaparral-foothill
woodland habitat at an elevation of about
550 m (1800 ft), probably from Eriogonum
fasciculatum (Gill, pers. comm.).
89
Etymology.—The species epithet, a noun
in apposition, 1s Latin meaning “long tail.”
The name refers to the ovipositor sheath of
the female.
DISCUSSION AND COMPARATIVE
COMMENTS
The four species in the genus Belaspidia
are: obscura Masi from Central and South-
ern Europe (Bouéek 195 1a, Peck et al. 1964),
Syria, Turkey (Bouéek 1951b, 1956) Cri-
mea, Transcaucasus, Central Asia, and Iran
(Nikolskaya 1952 [under the synonym nig-
ra Masi]); masit Nikolskaya from Central
Asia (Nikolskaya 1952); meridionalis Stef-
fan from France (Steffan 1951la, 1951b) and
longicauda n.sp. from California, USA.
The type species of Belaspidia (Masi 1916)
is obscura, described from females. Belas-
pidia nigra was described from a male spec-
imen (Masi 1927). Nikolskaya (1952) treat-
ed the Belaspidia of the United Soviet
Socialist Republic; described masii and rec-
ognized obscura and nigra. Steffan (195 1a)
described meridionalis from Toulon, France.
He treated the Belaspidia of France; rec-
ognized meridionalis, obscura, and nigra
(Steffan 1951b). Nikolskaya (1960) recog-
nized three world species: obscura, masii,
and meridionalis with nigra synonymized
under obscura by Bouéek (1951b). Bouéek
(1951b, 1956) indicated that meridionalis
might be a variety of obscura.
In Nikolskaya (1952), longicauda differs
from masii in that the antennae are longer
than head height, the funicular segments are
longer than wide, the first 2 abdominal ter-
gites have a thin, shiny band on posterior
margin; and the ovipositor projects poste-
riorly from the epipygidium by the same
length as T#1 dorsally. As with masii, lon-
gicauda has the hindfemur 2 as wide as
long. In Steffan (1951b), Jongicauda differs
from meridionalis in that the left mandible
is bidentate. The head and antennal char-
acters are similar for both species. In both
keys, /ongicauda differs from obscura in that
the thorax is slightly, not highly convex, the
90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pronotum is 5 times as wide as long versus
3 times, and the wings are colorless versus
slightly darkened.
ACKNOWLEDGMENTS
I thank Z. Boucek, Commonwealth In-
stitute of Entomology, London, for material
of Belaspidia obscura and for examining and
confirming the taxonomic placement of B.
longicauda, D. J. Burdick and K. J. Wood-
wick, both of California State University
Fresno, Fresno, for the use of laboratory
facilities and equipment, D. J. Burdick and
N. J. Smith, Fresno County Agricultural
Commissioner’s Office, Fresno, California,
R. D. Haines, Tulare County Agricultural
Commissioner’s Office, Visalia, California,
for editorial comments on this paper, L. E.
Caltagiorone, University of California,
Berkeley, for permission to deposit the al-
lotype male in the USNM; and the several
institutions and their personnel for the op-
portunity to have examined their material.
LITERATURE CITED
Bouéek, Z. 1951a. The first revision of the European
species of the family chalcididae (Hymenoptera).
Acta Entomol. Mus. Natl. Pragae 27 (Supplemen-
tum 1): 108 pp.
1951b. Results of the Zoological Scientific
Expedition of the National Museum in Praha to
Turkey, Hymenoptera I, Chalcidoidea (first part).
Acta Entomol. Mus. Natl. Pragae 383: No. 7, 47-
Si:
. 1956. A contribution to the knowledge of the
Chalcididae, Leucospididae and Eucharitidae
(Hymenoptera, Chalcidoidea) of the Near East.
Bull. Res. Counc. Israel, Jerusalem 5B: 227-259.
Masi, L. 1916. Calcididi del Giglio. I. Ann. Mus.
Civ. Genova 47: 54-122.
1927. Diagnosi di una n. sp. di Belaspidia.
Boll. Soc. Entomol. Ital. 59: 82-83.
Nikolskaya, M. N. 1952. The chalcid fauna of the
U.S.S.R. (Chalcidoidea). Zool. Inst. Akad. Nauk
SSR, Moscow No. 44, 240 pp.
1960. Fauna USSR: Hymenoptera, Vol. VII
(No. 5). Chalcidoids, families Chalcididae and
Leucospidae. Zool. Inst. Akad. Nauk SSR, Mos-
cow (n.s.) No. 76, 221 pp.
Peck, O., Boucek, Z., and Hoffer, A. 1964. Keys to
Chalcidoidea of Czechoslovakia (Insecta: Hyme-
noptera). Mem. Entomol. Soc. Can. 34: 170 pp.
Steffan, J. R. 195la. Remarques sur queiques genres
d’Haltichellinae (Hymenopteres: Chalcididae).
Bull. Mus. Natl. Hist. Nat. Paris. 23(4): 375-380.
1951b. Les espéces francaises d’ Haltichel-
linae (Hymenopteres: Chalcididae). Feuille Nat.
(n.s.) 6(9-10): 81-85.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 91-96
THE CULTUS DECISUS COMPLEX OF EASTERN NORTH
AMERICA (PLECOPTERA: PERLODIDAE)
BILL P. STARK, STANLEY W. SzCzYTKO AND
Boris C. KONDRATIEFF
(BPS) Dept. of Biology, Mississippi College, Clinton, Mississippi 39058; (SWS) College
of Natural Resources, Univ. of Wisconsin, Stevens Point, Wisconsin 54481; (BCK) Dept.
of Entomology, Colorado State Univ., Fort Collins, Colorado 80523.
Abstract.—The Cultus decisus complex of eastern North America is reviewed, and C.
verticalis (Banks) and C. decisus isolatus (Banks) are removed from synonymy with C.
decisus decisus (Walker). The three taxa have distinctive eggs, but no differences were
observed in adults of C. decisus decisus and C. decisus isolatus.
Since Ricker’s (1952) study, Cultus de-
cisus (Walker) has been regarded as a vari-
able species which ranges from southern
Canada, along the Appalachians, to north-
ern Georgia. The most recent taxonomic
treatment (Hitchcock 1974) includes Perla
verticalis Banks and Isoperla isolata Banks
as synonyms, but our study of available types
and other specimens suggests these names
represent valid taxa. C. decisus has also been
incorrectly applied to specimens from Great
Whale River (Ricker et al. 1968), which are
more similar to C. aestivalis (Needham and
Claassen), and by P. P. Harper (unpub. rec-
ord) to a similar series collected at James
Bay. Careful study of the western Nearctic
Cultus species is needed to clarify the status
of these specimens.
Specimens used in this study are depos-
ited in the British Museum of Natural His-
tory (BMNH), the Canadian National Col-
lection (CNC), the Illinois Natural History
Survey (INHS), the Museum of Compara-
tive Zoology (MCZ), the Royal Ontario Mu-
seum (ROM), the United States National
Museum of Natural History (USNM), the
Virginia Polytechnic Institute and State
University collection (VPI), and in the pri-
vate collections of R. F. Kirchner (RFK)
and the authors (BPS, SWS, BCK).
Cultus decisus decisus
(Walker)
Perla decisa Walker (1852). Holotype é, St.
Martin’s Falls, Albany River, Ontario
(BMNH).
Diploperla decisa: Ricker (1944).
Tsogenus (Cultus) decisus: Ricker (1952), in
part.
Cultus decisus: hes (1966), in part.
Male genitalia.—Lobe on sternum 7 well
developed, no lobe on sternum 8. Epiproct
with weakly sclerotized anterior and pos-
terior bands; mesal section membranous
without spinules or setae. Lateral stylets
long, slender and slightly curved ventrad.
Membranous cowl with a pair of postero-
lateral spiculate lobes and weakly sclero-
tized paragenital plates. Hemitergal lobes
typical of Diploperlini (Fig. 1).
Female genitalia.—Subgenital plate
broadly triangular, reaching beyond mid-
point of sternum 9.
Egg. — Dorsal surface and lid covered with
irregular, hexagonal follicle cell impressions
92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-3.
terminalia lateral. 3. C. decisus isolatus female sterna 8-9.
(FCIs). FCI walls thick and smooth, con-
cave floors contain ca. 30 minute aeropyles.
Sharp, smooth keel extends from posterior
third to near lid (Fig. 4). Lid offset by ir-
regular suture which becomes less distinct
posteriorly; lateral margins smooth (Fig. 4).
Ventral surface flat and covered with FCIs.
FCI walls consist of rows of shallow pits.
Lip smooth, anchor without specialized su-
perstructure. Micropyles ventral, slightly
anterior to equator (Fig. 5).
Specimens examined.— MICHIGAN:
Benzie Co., Platte River, Honor, 27-V-39,
T. H. Frison and H. H. Ross, 2 6, 1 2? (INHS).
C. decisus genitalia. 1. C. decisus decisus epiproct complex, lateral. 2. C. decisus isolatus male
Cheboygan Co., Sturgeon River, 3-VII-38,
J. W. Leonard, 1 2 (USNM). Mecosta Co.,
Muskegon River, Big Rapids, 22-V-36, T.
H. Frison and H. H. Ross, 2 6, 5 2 (USNM,
INHS). Otsego Co., Pigeon River, 23-VI-
36, J. W. Leonard, | 6, 2 2 (USNM, INHS).
ONTARIO: Albany River, St. Martin’s
Falls, 1 6 (Holotype, BMNH).
Cultus decisus isolatus (Banks),
New Status
Isoperla isolata Banks (1920). Holotype 4,
Hot Springs, (Madison County) North
Carolina (MCZ #10,824).
Figs. 4-7. C. decisus eggs. 4. C. decisus decisus, Otsego Co., Michigan, dorsal aspect, 380x. 5. C. decisus
decisus ventral aspect, 300 =. 6. C. decisus isolatus, Montgomery Co., Virginia, dorsal aspect, 300 =. 7. C. decisus
isolatus ventral aspect, 300 x (Ld—Lid, Lp—Lip, K—Keel, M—Micropyle, A— Anchor).
93
94
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pret =
Pete
SUETTIEG Ey
ye cere
st
Figs. 8-11.
C. verticalis genitalia. 8. Epiproct complex, lateral. 9. Epiproct complex, dorsoanterior aspect.
10. Male sterna 7-8. 11. Female sterna 8-9.
VOLUME 90, NUMBER 1
95
Figs. 12-14.
14. Dorsoanterior end, 500 =.
Perla decisa: Needham and Claassen (1925),
in part.
Cultus decisus: Mlies (1966), in part.
Male genitalia.—Indistinguishable from
C. decisus decisus (Fig. 2).
Female genitalia. —Indistinguishable from
C. decisus decisus (Fig. 3).
Egg.—Similar to C. decisus decisus but
dorsal FCIs enclose ca. 10-15 aeropyles, and
keel extends from lid to posterior margin
(Figs. 6-7).
Specimens examined.— GEORGIA: Fan-
nin Co., Toccoa River, Hwy 60, 19-V-85,
B. C. Kondratieff, 1 6, 12 2 (VPI). NORTH
CAROLINA: Madison Co., Hot Springs, 1
6 (Holotype, MCZ). VIRGINIA: Carroll Co.,
New River, Rt. 606, 19-V-80/22-IV-81, B.
C. Kondratieff, 2 6, 3 2 (VPI). Montgomery
Co., Little River, Rt. 787, 24-V-83, B. C.
Kondratieff, 1 2 (VPI).
Comments.— Reliable characters for dis-
tinguishing adults of C. decisus decisus from
C. decisus isolatus were not observed. Be-
cause of the range disjunction and the dis-
tinctive eggs (see Figs. 4 and 6), we are giv-
ing these populations subspecific status.
Hopefully, gravid females with associated
males and nymphs from intermediate lo-
C. verticalis eggs, Giles Co., Virginia. 12. Dorsal aspect, 270 x. 13. Ventrolateral aspect, 270 =.
calities will soon be available to permit test-
ing of this classification.
Cultus verticalis (Banks),
New Stratus
Perla verticalis Banks (1920). Holotype 8,
Franconia, (Grafton Co.) New Hamp-
shire (MCZ #10,815).
Diploperla verticalis: Frison (1942).
Tsogenus (Cultus) decisus: Ricker (1952), in
part.
Cultus decisus: Mlies (1966), in part.
Male genitalia.— Lobe on sternum 7 well
developed, lobe on sternum 8 small (Fig.
10). Epiproct relatively more sclerotized
than in C. decisus, and with small serrations
along anterior sclerite. Lateral stylets, cowl
and hemitergal lobes typical of genus (Figs.
8-9).
Female genitalia.—Lateral margins of
subgenital plate more convex and plate rel-
atively longer than in C. decisus (Fig. 11).
Egg.— Dorsal surface covered with faint
FCIs and scattered globular bodies. FCI
walls are shallow furrows, floors without
aeropyles. Keel smooth with a few globular
bodies clustered at either end (Figs. 12, 14).
Ventral surface covered with FCIs similar
96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
to those on dorsum. Micropyles ventral,
slightly anterior to equator (Fig. 13).
Specimens examined.—NEW HAMP-
SHIRE: Grafton Co., Franconia, Slosson, |
2 (Holotype, MCZ). NORTH CAROLINA:
Haywood Co., Cataloochee Creek, Great
Smoky Mountains National Park, 16-V-83/
3-V-85, B. C. Kondratieff, 16 4, 8 2 (BCK).
Swain Co., Oconoluftee River, Cherokee,
2-VI-78, B. Stark and K. W. Stewart, 1 é
(BPS). Deep Creek, Deep Creek Camp-
ground, 19-V-70, G. Wiggins and T. Ya-
mamoto, | 6 (ROM). PENNSYLVANIA:
Franklin, 23-VI-67, S. Gilmore, 1 ¢
(USNM). QUEBEC: Brome, 1-VI-36, G. S.
Walley, 1 ¢ (CNC). TENNESSEE: Sevier
Co., Little River, Great Smoky Mountains
National Park, 5-V-79, B. Stark, 2 6, 2
(BPS). VERMONT: Bennington Co., Bat-
tenkill, Rt. 313, 25-V-80, A. C. Graham, |
8(SWS). VIRGINIA: Giles Co., Little Ston-
ey Creek, Rt. 460, 7-17-V-80, B. C. Kon-
dratieff, 7 6, 2 2 (VPI). Big Stoney Creek,
Rt. 635, 26-V-78, D. Gray and D. Minnick,
19 6 (VPI). Stony Creek, 28-V-79, E. Bel-
linger, 1 6 (VPI). Patrick Co., Dan River,
Rt. 601, 16-V-81, Geivard, | (VPI). Smyth
Co., Big Laurel Creek, 28-V-78, G. T. and
M. W. Voreh, | 4, 1 2 (RFK). WEST VIR-
GINIA: Pocahontas Co., Williams River,
Day Run, 30-VI-82, W. Mathis and O. S.
Flint, 2 ¢ (USNM).
Comments.—The pinned female holo-
type had no eggs, but the subgenital plate is
consistent with our figures from a Giles Co.,
Virginia, specimen (Fig. 11) and the other
specimens listed above. Genitalic illustra-
tions given by Needham and Claassen (1925)
(as Perla verticalis), Hitchcock (1974) (as C.
decisus) and Kondratieff and Voshell (1982)
(as C. decisus) are of this species.
KEY TO CULTUS DECISUS
ComPLEX ADULTS
1. Anterior sclerite of epiproct armed basally with
small serrations (Figs. 8, 9); sternum 8 with
small lobe (Fig. 10); female subgenital plate
parabolic, reaching posterior margin of ster-
num 9 (Fig. 11); chorionic FCIs not enclosing
aeropyles (Figs. 12-14) C. verticalis
— Anterior sclerite of male epiproct without ser-
rations (Figs. 1, 2); sternum 8 without lobe;
female subgenital plate triangular, not reaching
posterior margin of sternum 9 (Fig. 3); cho-
rionic FCIs enclose aeropyles (Figs. 4,6) .... 2
Aeropyles of chorionic FCIs densely packed,
small, ca. 30 or more in each FCI (Fig. 4);
dorsal keel of egg not extending onto lid (Fig.
4): known from Great Lakes region
eh SO oR ooe C. decisus decisus
-— Aeropyles of chorionic FCIs larger and more
widely spaced, ca. 15 or less in each FCI (Fig.
6); dorsal keel extends onto base of lid (Fig. 6);
known from southern Appalachians ........
Se cdc neh Me eat C. decisus isolatus
i)
ACKNOWLEDGMENTS
We are grateful to O. S. Flint, P. Harper,
J. E. H. Martin, R. F. Kirchner, C. Vogt, J.
R. Voshell, D. Webb and G. Wiggins for
arranging the loan of specimens. D. L. Lentz
and S. Faison assisted in SEM micrograph
preparation. This study was supported by
NSF grant #BSR 840755.
LITERATURE CITED
Banks, N. 1920. New neuropteroid insects. Bull. Mus.
Comp. Zool. 64: 299-362.
Frison, T. H. 1942. Studies of North American Ple-
coptera, with special reference of the fauna of II-
linois. Bull. Ill. Nat. Hist. Surv. 22: 235-355.
Hitchcock, S. W. 1974. Guide to the insects of Con-
necticut. Part VII. The Plecoptera or stoneflies of
Connecticut. Bull. St. Geol. Nat. Hist. Surv. Conn.
107: 1-262.
Illies, J. 1966. Katalog der rezenten Plecoptera. Das
Tierreich, 82. Walter de Gruyter and Co., Berlin.
632 pp.
Kondratieff, B. C. and J. R. Voshell, Jr. 1982. The
Perlodinae of Virginia, USA (Plecoptera: Perlo-
didae). Proc. Entomol. Soc. Wash. 84: 761-774.
Needham, J. G. and P. W. Claassen. 1925. A mono-
graph of the Plecoptera or stoneflies of America
north of Mexico. Thomas Say Found. Entomol.
Soc. Amer. 2: 1-397.
Ricker, W. E. 1944. Some Plecoptera from the far
North. Canad. Entomol. 76: 174-185.
1952. Systematic studies in Plecoptera. Ind.
Univ. Pub. Sci. Ser. 18: 1-200.
Ricker, W. E., R. Malouin, P. Harper, and H. H. Ross.
1968. Distribution of Quebec stoneflies (Plecop-
tera). Nat. Can. 95: 1085-1123.
Walker, F. 1852. Catalogue of the specimens of neu-
ropterous insects in the collection of the British
Museum. Part I. Phryganides-Perlides. London:
British Museum. 192 pp.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, p. 97
NOTE
Neotype Designation for Raptoheptagenia cruentata
(Walsh) (Ephemeroptera: Heptageniudae)
Individuals of predaceous flatheaded
mayfly larvae in North America that have
been tentatively known as Anepeorus
McDunnough (Burks, B. D. 1953. Ill. Nat.
Hist. Surv. Bull. 26: 1-216) were recently
reared through to adults of Heptagenia
cruentata Walsh by Whiting and Lehmkuhl
(1987. Canad. Entomol. 119: 405-407) in
Saskatchewan. Those authors, while cor-
rectly associating the larvae, erected the ge-
nus Raptoheptagenia (= Heptagenia cruen-
tata), recognizing the highly unusual larvae.
These larvae have evidently been referable
to either species of Anepeorus —the eastern
A. simplex (Walsh) or the western A. rusticus
McDunnough—because the range of R.
cruentata includes both areas, and Mc-
Cafferty and Provonsha (1985. Gr. Lakes
Entomol. 18: 1-6) found that Saskatchewan
larvae presumed to be A. rusticus and east-
ern larvae presumed to be 4. simplex were
morphologically identical.
The original types of Heptagenia cruen-
tata were destroyed in the Chicago fire of
1871 (Burks 1953). Although the single male
adult and single female adult now residing
in the Harvard University Museum of
Comparative Zoology were identified as H.
cruentata by Walsh, they are not available
as lectotypes because they were collected a
year after the published description (Walsh,
B. D. 1863. Proc. Entomol. Soc. Phila. 2:
167-272). Thus, the name has remained a
nomen dubium, with the descriptions of
McDunnough (1924. Canad. Entomol. 56:
90-98) (as Heptagenia reversalis), Traver
(1935. In Needham, J. G., J. R. Traver, and
Y. C. Hsu. The Biology of Mayflies. Com-
stock Publ. Co., Ithaca, NY. 759 pp.), and
Burks (1953) providing the bases for iden-
tification. I therefore designate a neotype for
Raptoheptagenia cruentata (Walsh) as fol-
lows: Larva: Indiana, Martin County, East
Fork White River at Hindustan Falls Public
Fishing Site, VII-15-1982, A. V. Provonsha
and V. Van Allen; in alcohol with “‘neo-
type” label; deposited in the type collection
of the Purdue Entomological Research Col-
lection, West Lafayette, Indiana. Since the
generic concept is based on the distinctive
larval morphology, and in light of the recent
reared association, a larva is appropriate as
the neotype specimen of the type species of
the monospecific Raptoheptagenia. Adults
of R. cruentata have also been taken at the
neotype locality.
This paper is Purdue Experiment Station
Journal No. 11251.
W. P. McCafferty, Department of Ento-
mology, Purdue University, West Lafayette,
Indiana 47907.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 98-100
NEW SYNONYMS PERTAINING TO CHELIFERA AND GENERIC
KEY FOR NORTH AMERICAN HEMERODROMIINAE
(DIPTERA: EMPIDIDAE)
JOHN F. MACDONALD
Department of Entomology, Purdue University, W. Lafayette, Indiana 47907.
Abstract. —Empididae previously placed in Thanategia Melander are shown to be species
of Chelifera Macquart: Chelifera defecta (Loew) n. comb.; Chelifera recurvata (Melander)
n. comb.; and Chelifera stuprator (Melander) n. comb. Males with identical terminalia
possessed significant variation in the expression of crossvein dm-cu that renders it invalid
as a character of generic importance. Therefore, Thanategia is considered here a new
junior synonym of Chelifera. Also, Chelifera knutsoni Lavallee is shown to be a junior
synonym of Chelifera defecta (= T. defecta (Loew)). A revised key to the genera of North
American Hemerodromiinae is provided.
Aquatic dance flies (Diptera: Empididae)
in the subfamily Hemerodromiuinae were last
revised by Melander (1947). New revision-
ary studies of Nearctic Hemerodromiinae
now underway reveal that certain genera and
species should be placed in synonymy.
Thanategia was described as a subgenus of
Chelifera by Melander (1928: 263; type
species, Hemerodromia defecta Loew, 1862:
210) to accommodate those species that
possessed an open cell dm (= “incomplete
discal cell” or “discal cell fused with third
posterior cell’’) due to the absence of cross-
vein dm-cu (= “apical crossvein’’). In other
respects, the three species placed in that
subgenus, Thanategia defecta (Loew),
Thanategia recurvata Melander, and Than-
ategia stuprator Melander, fit the concept of
Chelifera. Without an explanation, Melan-
der (1947) later elevated Thanategia to ge-
neric status.
Examination of all specimens labelled as
Thanategia and Chelifera in the Canadian
National Collection (CNC), Ottawa, re-
vealed that males of 7. recurvata were iden-
tical to males placed in the CNC and la-
belled as Chelifera “new sp. nr. scrotifera.”
Examination of wing venation revealed that
all specimens identified as 7hanategia either
completely or partially lacked crossvein dm-
cu in one wing or both wings, whereas those
identified as Chelifera possessed a complete
dm-cu in both wings; thus, all specimens
keyed according to Melander (1928, 1947).
However, in other morphological features,
including all details of male terminalia,
males of “Chelifera new sp. near scrotifera”
were identical to males of Thanategia re-
curvata. The same study found a male placed
in the CNC as Chelifera “new sp. nr. bank-
si’ with terminalia exactly fitting the de-
scription of 7. defecta and identical to a
paratype male of Chelifera knutsoni Lavy-
allee, plus a paratype female of C. knutsoni
with incomplete expression of dm-cu in one
wing resulting in cell dm being “open” api-
cally.
Subsequent examination of holotype
males of 7. recurvata and T. stuprator and
additional paratype males and females of
Chelifera knutsoni together with study of
two series of specimens obtained in loans
from the University of New Hampshire and
Washington State University eventually
VOLUME 90, NUMBER 1
clarified the males and facilitated associa-
tion of conspecific males and females.
General analysis of venation of all avail-
able males (N = 8) with terminalia matching
the very distinctive terminalia of recurvata
(Melander 1947: 260; fig. 34) revealed that
three (including the holotype) possessed
‘“Thanategia venation,’ three possessed
“Chelifera venation,” and two could key to
Thanategia since cell dm was open in at
least one wing. Of the four associated fe-
males, two possessed “Chelifera venation”
and two (including the allotype) possessed
an open cell dm in at least one wing. This
variation in dm-cu indicates that “7. re-
curvata’’ simply represents individuals
lacking development of dm-cu, and there-
fore this species is transferred to the genus
Chelifera: Chelifera recurvata (Melander)
(= Thanategia recurvata Melander), New
Combination.
Study of a series of males (N = 8) with
terminalia matching the description of 7.
defecta (Melander 1947: 259, fig. 32) (col-
lected July—Oct., 1985 by D. S. Chandler
near Wonalancet, New Hampshire), togeth-
er with examination of two paratype males
of C. knutsoni (collected Macon Co. North
Carolina by A. G. Lavallee (1975)) and
comparison of the above mentioned two
males in the CNC, clarified another ques-
tion. Every aspect of male terminalia among
the above males was identical; however,
these males varied significantly in the
expression of dm-cu which, based on Me-
lander’s 1947 key, would lead to identifi-
cation of half as 7. defecta (1.e. dm-cu com-
pletely or partially absent) and half as C.
knutsoni (i.e. dm-cu complete in both wings).
A similar result would occur when keying
associated females. Examination of C. knut-
soni paratypes (two males and two females),
all possessing a complete dm-cu in both
wings, places them in the genus Chelifera.
The comparison, however, of all males con-
sidered above with the general description
and specific features of male terminalia of
T. defecta results in the following nomen-
99
clatorial revision: Chelifera defecta (Loew)
[= Thanategia defecta (Loew)] New Com-
bination (= Chelifera knutsoni Lavallee) New
Synonym.
The study of 7. stuprator was puzzling for
two reasons. First, all specimens first ex-
amined (four males, including the holotype,
and six females) fit the description of this
species (Melander 1947: 259; fig. 33) and
Melander’s concept of Thanategia based on
venation. Second, in addition to lacking
crossvein dm-cu diagnostic for Thanategia,
all specimens possessed a simple R,,; and
thus did not possess the branched R, , ; typ-
ical of Chelifera. Melander’s (1947) descrip-
tion of the holotype male indicated that R,
was “vestigial,” but my examination re-
vealed that R, was completely lacking in the
right wing and was expressed as an incom-
plete, approximately 2 mm, rudimentary
crossvein arising from the costa in the left
wing. Subsequent acquisition of a series of
T. stuprator (collected 12-13-VIII-1977 by
W. J. Turner in Mt. Ranier National Park)
substantiated the suspected variability in
venation. All three sfuprator males and 25/
27 associated females lacked dm-cu in both
wings, but two females possessed a partial
dm-cu in both wings. Variability pertaining
to R, was as follows: the three males pos-
sessed either complete or partially complete
R, forks in either one or both wings; 11
females expressed no development of R, in
either wing; one female had a complete R,
in the right wing (completely lacking in the
left wing); and 14 females possessed a par-
tial R, in both wings (six females) or only
the right wing (four females) or only the left
wing (four females). Clearly, R, is a trait
subject to individual variability. As regards
dm-cu, its absence may be typical of stu-
prator, but it is not of generic importance
and this species is transferred to the genus
Chelifera: Chelifera stuprator (Melander)
(= Thanategia stuprator Melander) New
Combination.
Variation in crossvein dm-cu also exists
in other species of Chelifera. For example,
100
occasional specimens of C. obsoleta (Loew)
from Indiana and Georgia, C. ensifera Me-
lander from Oregon, C. calaga Lavallee from
Utah, and C. cirrata Melander from Wash-
ington state, either lacked or possessed a
partial crossvein dm-cu in one wing or both
wings, and would “run to” 7hanategia in
existing keys.
Significant variation in the expression of
crossvein dm-cu revealed in the three species
formerly placed in the genus 7hanategia, as
well as in several other species of Chelifera,
is adequate evidence for placing 7hanategia
Melander as a junior synonym of Chelifera
Macquart, New Synonym, as has been shown
here by the correct generic placement of the
type species and the other two described
species of the former genus.
An additional consequence of this study
is the revision of the last portion of Melan-
der’s (1947) key and the related portion of
Steyskal and Knutson’s (1981) key to North
American genera of Hemerodromiinae,
which is here reproduced in its entirety with
the needed modifications stemming from
this study, as well as a few additional minor
changes:
1. Antenna with an arista, more than twice as long
as basal flagellomere; mesoscutum with several
pairs of well developed setae; laterotergite with
setae; male terminalia more or less reflexed
over abdomen, with terminal processes pro-
jecting anteriorly ... PRS. AES cM A 2
1’. Antenna with a stylus, chores than basal flag-
ellomere; mesoscutum without well developed
setae (a pair of supra-alars may exist); later-
otergite bare; male terminalia erect or pro-
jecting posteriorly 3}
2. Celldm closed apically (crossvein 1dm- -cu pres-
ent) Chelipoda (Chelipoda) Macquart
2’. Cell dm open apically (crossvein dm-cu ab-
SOMt) tts yy oes cia 206.5 Chelipoda (Phyllodromia)
(some authors treat Phyl/lodromia as a separate
genus, Phyllodromia Zetterstedt)
3. Anal cell absent and CuA, not developed; R,
ending before mid-wing; Sc fused with C close
to wing base; crossvein h absent .
Ene GaSe EOD oe Hemerodromia Meigen
3h Anal ‘ell present, or at least CuA, strongly
developed; R, ending at or beyond mid-wing;
Sc distinctly free of C, but evanescent apically;
crossvein h present
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
4. Veins M, and M, not petiolate (1.e. without a
common stem); cell dm fused basally with cell
bm; front femur relatively slender and typi-
cally lacking strong setae beneath
peLeiA! Neoplasta Coquillett
4’. Veins M, and M, petiolate (i.e. with acommon
stem); cell dm variable; front femur swollen
and possessing strong setae beneath ......... 5
5. Cells bm and dm fused (i.e. crossvein bm-cu
Absentee | dae ae Metachela Coquillett
‘. Cells bm and dm separate (i.e. crossvein bm-
cu present); crossvein dm-cu usually present,
but occasionally partially or totally lacking,
opening cell dm apically (the two wings may
differ in this regard on the same specimen)
Chelifera Macquart (Includes Thanategia
defecta, T. recurvata, and T. stuprator)
wn
ACKNOWLEDGMENTS
The following institutions loaned the ma-
terial upon which this work is based: D. S.
Chandler, University of New Hampshire,
Durham; L. Knutson, IIBIII, U.S. Depart-
ment of Agriculture (U.S. National Mu-
seum of Natural History, Washington, D.C.):
R. T. Schuh, American Museum of Natural
History, New York; H. J. Teskey, Biosys-
tematics Research Centre, Ottawa (Cana-
dian National Collection); and W. J. Turn-
er, Washington State University (James
Entomological Collection).
Sincere appreciation is extended to H. J.
Teskey and W. J. Turner for their generous
assistance during the study, and to W. P.
McCafferty for critically reading the manu-
script. This paper is Purdue Agricultural Ex-
periment Station journal No. 11,247.
LITERATURE CITED
Lavallee, A. G. 1975. New Nearctic Chelifera (Dip-
tera: Empididae). J. Georgia. Entomol. Soc. 10:
95-98.
Melander, A. L. 1928. Diptera, Fam. Empididae.
Fascicle 185, pp. 1-434, plates 1-8. Jn P. Wyts-
man, ed., Genera Insectorum. Bruxelles.
1947. Synopsis of the Hemerodromiinae
(Diptera: Empididae). J. New York Entomol. Soc.
55: 237-273.
Steyskal, G.C.,and L. V. Knutson. 1981. Empididae.
47, pp. 607-624. In McAlpine, J. F. et al., eds.
Manual of Nearctic Diptera. Res. Br., Agric. Can-
ada Monogr. 27(1): 1-674.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 101-105
A NEW SPECIES OF DONACEUS CRESSON (DIPTERA: EPHYDRIDAE)
FROM MALAYSIA
RICHARD S. ZACK AND ROBERT W. SITES
(RSZ) Department of Entomology, Washington State University, Pullman, Washington
99164-6432: (RWS), Department of Entomology, Texas Tech University, Lubbock, Texas
79409.
Abstract. —Donaceus azhari, new species, is described from Perak State, Peninsular
Malaysia. It is compared to its sole congener, D. nigronotatus Cresson.
The shore fly fauna of Malaysia 1s poorly
known, as it is for most of the Oriental Re-
gion. Cogan and Wirth (1977) record 16
species from Peninsular Malaysia (Malaya)
and only three in Sarawak and Sabah (Ma-
laysian Borneo). The number of species in
this area will most likely prove to be much
greater. During November of 1986, one of
us (RSZ) spent a month in Peninsular and
East Malaysia. During that period several
new species and material which will expand
the distributional ranges of other Oriental
ephydrids was discovered. This is the first
report concerning the malaysian material.
Herein we describe the second species in
Donaceus.
Genus Donaceus Cresson
Donaceus Cresson, 1943: 5 (type species:
Donaceus nigronotatus Cresson, by orig-
inal designation and monotypy).
Diagnosis. —Small shore flies (1.50-2.40
mm) similar to //ythea Haliday and Zeros
Cresson but distinguished by a variety of
subtle features.
Head: Microtomentose, produced for-
ward, appearing oversized for body; face flat
to facial prominence then with a slight tu-
berculose development, with 3 large facial
setae, equally spaced, uppermost at level just
below facial prominence; genal bristle sub-
equal to facials; antennae normal, plumose,
with 6-7 aristal hairs; proclinate and recli-
nate orbital setae well developed; inner and
outer vertical setae well developed; ocellar
setae large, situated between posterior ocel-
li; eyes micropubescent.
Thorax: Thoracic chaetotaxy well devel-
oped; scutum with 3 pair of dorsocentral
setae (1+2), 3-6 pair of acrostichal setae,
prescutellar and intra-alar setae strong; scu-
tellum with large lateral and apical setae,
lateral margins with or without black, vel-
vety areas; notopleuron with posterior seta
2x diameter of anterior seta and 4 as far
from notopleural suture; wing distinctive,
with clear rounded spots in a fuscous field,
vein R,.,, joins costa beyond middle of wing;
halter yellow.
Abdomen: Densely microtomentose,
ground color black; male genitalia with
epandrium not fused dorsally, cerci elon-
gate, surstyli not heavily sclerotized, var-
iously lobed and bristled.
Distribution. —Southeast Asia (Malaysia,
Thailand, Japan, Taiwan), Hawaii, Austra-
lia, and New Zealand.
Remarks.—Cresson (1943) erected the
tribe Ilytheini for three closely related gen-
era including //ythea Haliday, and the new
genera Zeros and Donaceus. Donaceus 1s
separated from J/ythea and Zeros by a num-
ber of subtle characters the most obvious
102
0.2mm
Fig. 1. Donaceus nigronotatus. Male external gen-
italia.
being the presence of 3 dorsocentral bristles
(1+2) as opposed to 2 (1+1) in the latter
genera. In addition, vein R, , , joins the cos-
ta at a point between those found in //ythea
and Zeros. Facial similarities also seem to
suggest a close relationship with Zeros. The
distribution of Donaceus also implies a close
relationship with Zeros as both are found
throughout the Oriental Region, Donaceus
being limited to this area, while //ythea has
an Oriental presence only in Japan.
It is possible that upon closer examina-
tion Donaceus may not prove to merit a
generic ranking and will be considered a
subgenus or other grouping within //ythea.
However, the question will only be resolved
once the tribe is studied on a world basis.
Thus, for the present, we have chosen to
keep Donaceus as a valid genus.
KEY TO SPECIES OF
DONACEUS
1. Coxa and femur pale, golden; scutellum with
black, velvety patches on lateral angles; size
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
larger, 1.90-2.40 mm (widespread) .........
SNE ae IS Ee ot D. nigronotatus Cresson
— Coxaand femur dark, black; scutellum without
black, velvety patches on lateral angles; size
smaller, 1.50-1.93 mm (Peninsular Malaysia)
D. azhari, new species
Donaceus nigronotatus Cresson
Fig. |
Donaceus nigronotatus Cresson, 1943: 5.
Diagnosis.—A small shore fly, length 2.0-
2.40 mm.
Description.— Head: Microtomentose,
subshiny, golden, rarely becoming some-
what dusky; genal area concolorus with face.
Second antennal segment pale, golden, con-
trasting with third segment which 1s velvety
gray to brown. Frons slightly contrasting
with face, darker, concolorous in both
ground color and microtomentosity with
scutum. Mouthparts, including palpi, con-
colorus with face. Eye height/eye width ratio
1:0.83-1:0.88; eye height/genal height ratio
1:0.30-1:0.34.
Thorax.—Thoracic chaetotaxy well de-
veloped. Scutum with 3 pair of dorsocentral
setae (1 +2), and 3 pair of strong acrostichal
setae; microtomentose, dark golden brown,
ground color black, with rather diffuse
markings at bases of setae but lacking def-
inite blotches and pattern. Scutellum con-
colorous with scutum, in posterior view, lat-
eral angles with black, velvety patches.
Pleural areas golden, contrasting with dark
coloration of the scutum, concolorous with
face. Legs concolorous with pleural areas,
golden. Wing with clear, rounded spots in
a fuscous field. Halter yellow.
Abdomen.—Microtomentose, gray with
a slight tinge of green, ground color black.
Male genitalia as in Fig. 1.
Type material.— The holotype (Academy
of Natural Sciences of Philadelphia, type
number 6650) was examined. The specimen
is a female, minuten mounted and, with the
exception of the missing head, is in good
condition. It is labeled Takai 1907.V.3/
FORMOSA Sauter/845/Type 6650 Don-
aceus nigronotatus Cress. A male paratype
VOLUME 90, NUMBER 1
' 0.7mm {
Donaceus azhari. Wing.
Fig. 2.
and a female could not be located in the
ANSP Collection.
Distribution. — Cresson (1943) described
the species from three specimens taken in
Formosa (Taiwan, The Republic of China).
Since then the species has been recorded
from Japan (Miyagi 1977), Hawai (Hardy
and Delfinado 1980), Thailand, Australia,
and New Zealand (Cogan and Wirth 1977).
Specimens examined.—The holotype
from Formosa and 12 specimens from Oahu,
Maui, and Kauai, Hawaii.
Remarks.—Both Miyagi (1977) and
Hardy and Delfinado (1980) presented de-
scriptions and figured the genitalia. In Ha-
wail, Miyagi (1977) found the species to
range from sea level to 4000 ft. in elevation
and to occur in a variety of aquatic habitats
including the margins of ponds, swamps,
reservoirs, and streams.
Donaceus azhari Zack and Sites,
New SPECIES
Figs. 2-3
Diagnosis.—A small shore fly, length 1.50
to 1.93 mm. It is distinguished from its sole
congener D. nigronotatus Cresson by the
smaller size, overall darker appearance of
the thoracic pleura and legs, the lack of lat-
eral velvet patches on the scutellum and the
distinctive male terminalia.
Description. — Head: Head microtomen-
tose, produced forward. Face light golden
brown, in profile straight to facial promi-
k 0.2mm |
Donaceus azhari. Male external genitalia.
Fig. 3.
nence and then with only a slight tubercu-
lose development, face receding to antennal
bases; with 3 large, equally spaced, facial
setae, uppermost dorsally inclined and at a
level just below facial prominence, bottom
2 mesoventrally inclined, all 3 follow con-
tour of eye in arrangement; 3—4 small, dor-
sally inclined setae lateral to larger facials.
Genal area concolorous with face: genal
bristle subequal to larger facials; situated
forward, near eye margin; 3-4 small, dark
setae situated along posterior genal margin.
Antenna with second segment ochraceous
to dark golden brown, upper angle with 2
larger setae; flagellum dark, golden brown
with numerous fine hairs; arista dark, plu-
mose, 6-7 aristal hairs which are 2-4 length
of main trunk. Frons concolorous with face:
proclinate and reclinate orbital setae well
developed, with a second, small proclinate
orbital seta situated equidistant between the
larger setae; inner and outer vertical setae
well developed, subequal to facials; ocellar
setae larger, subequal to facials, situated be-
tween the posterior ocelli; 2 pair of small,
postocellar setae; 6-8 dark, small postocular
setae. Mouthparts, including palpi, micro-
tomentose, concolorous with face. Eyes mi-
104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
cropubescent. Eye height/eye width ratio 1:
0.83-1:0.86; eye height/genal height ratio 1:
0.30-1:0.33.
Thorax.—Thoracic chaetotaxy well de-
veloped. Ground color black; microtomen-
tose, dark golden brown, often with a slight
greenish to copper tinge. Scutum with 3 pair
of well developed dorsocentral setae, 1 +2;
5-6 pair of strong acrostichal setae, irreg-
ularly placed in 2 rows, prescutellars strong:
intra-alar setae strong, posterior pair espe-
cially so, subequal to dorsocentrals; nu-
merous, irregularly-placed setae in the in-
tra-alar and supra-alar areas. Scutum
somewhat patterned, blotched with dark
brown to black at base of each seta, often
with dark brown stripes between the dor-
socentral and acrostichal rows of setae. Scu-
tellum concolorous with scutum; large lat-
eral and apical scutellar setae, a pair of small,
hair-like subapical scutellar setae; in pos-
terior view, apical, and to some extent lat-
eral margins ochraceous, without black, vel-
vety areas. Pleural areas microtomentose,
concolorous with scutum; notopleuron with
posterior notopleural seta approximately 2
diameter of anterior seta and 4 x as far from
notopleural suture; anepisternum with |
large and 1-2 smaller setae, with numerous
hairs throughout; katepisternum with a large,
central seta and 3—4 smaller, hair-like setae.
Coxa concolorous with pleural areas; femur
dark, becoming paler, ochraceous distally
(at knee), profemur with a series of strong,
ventral setae which become longer and
thicker distally; tibia lighter, variegated,
mesotibia with a large, dark, apical tibial
spine; tarsi ochraceous. Wing distinctive
(Fig. 2) with clear rounded spots in a fuscous
field. Halter yellow.
Abdomen: Microtomentose, gray with a
slight greenish tinge, ground color black; a
row of small, dark setae along the posterior
margin of each tergite, other setae posi-
tioned in rows, more common in lateral
areas. Male with ochraceous spot on dorsal
tip of abdomen; male genitalia as in Fig. 3.
Type material.—Holotype 4, allotype 8,
and 5 paratypes (3 29, 2 43) labeled-Malay-
sia: Perak; MARDI-Hilir Perak, 16 mi W
Telok Anson. 25 November 1986. R. S. Zack
collector. All specimens are paper-point
mounted. The holotype and allotype are de-
posited in the James Entomological Collec-
tion at Washington State University. A male
and female paratype are deposited in the
National Museum of Natural History
(USNM). The remaining paratypes are in
the senior author’s collection.
Distribution.— Known only from the type
locality.
Etymology.—The species 1s named in
honor of our friend and colleague, Azhar
Ismail, an entomologist with the Cocoa and
Coconut Research Division of the Malaysia
Agriculture and Development Institute
(MARDI). Mr. Azhar served as the senior
author’s host during his trip to Malaysia.
Remarks. — Donaceus azhariis easily sep-
arated from its sole congener D. nigrono-
tatus by its smaller size (1.50-1.93 mm as
opposed to 2.0-2.40 mm), the much darker
microtomentosity of the thorax, the darker
legs, and the lack of velvety black lateral
margins on the scutellum.
The type series of D. azhari was collected
on the grounds of the Malaysian Agriculture
and Development Institute (MARDI), Hilir
Perak Station. The flies were taken along
the moist soil banks of a small, approxi-
mately 2 m wide, rain-fed drainage ditch.
The ditch is very susceptible to flooding.
Numerous other shore flies were also col-
lected at this site.
ACKNOWLEDGMENTS
We thank Donald Azuma and Ruth Grif-
fith of the Academy of Natural Sciences of
Philadelphia (ANSP) for making the type
specimen of Donaceus nigronotatus avail-
able to us. E. P. Catts, W. J. Turner, R. D.
Akre, L. Chandler, R. T. Ervin, and J. C.
Cokendolpher reviewed the manuscript.
This is Scientific Paper Number 7801, Ag-
ricultural Research Center, College of Ag-
riculture and Home Economics, Washing-
VOLUME 90, NUMBER 1
ton State University, Pullman, Washington
99164. The work was conducted under proj-
ect 9043. TTU Contribution T-10-183,
College of Agricultural Sciences, Texas Tech
University, Lubbock, Texas 79409.
LITERATURE CITED
Cresson, E. T., Jr. 1943. The species of the tribe
Ilytheini (Diptera: Ephydridae: Notiphilinae).
Trans. Am. Entomol. Soc. 69: 1-16.
Cogan, B. H. and W. W. Wirth. 1977. Family Ephyd-
ridae, pp. 321-339. Jn Delfinado, M. D. and D.
105
E. Hardy, eds., A Catalog of the Diptera of the
Onental Region. Volume III Suborder Cyclorrha-
pha (excluding Division Aschiza). Univ. Press Ha-
waul, Honolulu. 853 pp.
Hardy, D. E. and M. D. Delfinado. 1980. Insects of
Hawai, Volume 13, Diptera: Cyclorrhapha III,
Series Schizophora Section Acalypterae, Exclusive
of Family Drosophilidae. Univ. Press Hawaii, Ho-
nolulu. 451 pp.
Miyagi, I. 1977. Fauna Japonica-Ephydridae (Insec-
ta: Diptera). Keigaku Publ. Co., Tokyo. 113 pp. +
49 pls.
NOTE
A Sesiid Host Record for Prerocormus chasmodops
(Hymenoptera: Ichneumonidae)
The ichneumonid, Pterocormus chas-
modops (Heinrich), has not previously been
associated with any host. During the sum-
mer of 1986, pupae of the raspberry crown
borer, Pennisetia marginata (Harris) (Lep-
idoptera: Sesiidae), were collected in Wayne
County, Ohio, from the crowns of cultivat-
ed Rubus spp. as part of biological control
research dealing with this bramble pest.
From 30 field collected pupae, 22 P. mar-
ginata adults emerged, 3 died from a dis-
ease, and 5 ichneumonids emerged from the
remaining five pupae, | per pupa. These
specimens were subsequently identified as
Pterocormus chasmodops (Heinrich) (Hy-
menoptera: Ichneumonidae). The pub-
lished host range of Prerocormus chasmo-
dops is Quebec, Maine, New Hampshire,
New York, Ontario, Michigan, Minnesota
and Manitoba (Krombein et al. 1979. Cat-
alog of Hymenoptera in America North of
Mexico, Vol. I. pg. 521). This is the first
record of this species from Ohio. Bracon
bembeciae (Walley) (Hymenoptera: Bracon-
idae) is the only other species which has
been reared from Pennisetia marginata.
We thank Dr. Robert W. Carlson of the
Systematic Entomology Laboratory, the
Biosystematics and Beneficial Insects Insti-
tute, USDA, Beltsville, Maryland, for iden-
tifying Prerocormus chasmodops.
Daniel M. Pavuk and Roger N. Williams,
Department of Entomology, Ohio State
University, Ohio Agricultural Research and
Development Center, Wooster, Ohio 44691.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 106-121
STUDIES ON THE SYSTEMATICS OF THE SHORE-FLY
TRIBE DAGINI (DIPTERA: EPHYDRIDAE)
WAYNE N. MATHIS AND TADEUSZ ZATWARNICKI
(WNM) Department of Entomology, NHB 169, Smithsonian Institution, Washington,
D.C. 20560; (TZ) Department of Zoology, Academy of Agriculture, ul. Cybulskiego 20,
50-205, Wroclaw, Poland.
Abstract. —S1x new species of the tribe Dagini are described in the genera Psilephydra
Hendel (Aaskiensis, nepalensis, lyneborgi, iridescens) and Dagus Cresson (splanglerorum
and dominicanus). Revised keys are provided for all known species of both genera and
for the genera of the tribe. Psilephydra is rediagnosed and divided into two species groups
(the fluvialis and cyanoprosopa groups), and the phylogeny of the taxa treated is discussed.
A catalog to all taxa of Dagini is presented, incuding the new species and new distributional
data.
The purpose of this paper is to present
additional information on the shore-fly tribe
Dagini. Mathis (1982) proposed Dagini as
a tribe within the subfamily Ephydrinae.
Initially the tribe comprised four genera and
11 species as follows (number of species in-
dicated in parenthesis): Dagus Cresson (1),
Psilephydra Hendel (2), Physemops Cresson
(6), and Diedrops (Mathis and Wirth (2). In
three subsequent papers, Mathis (1983,
1984) and Mathis and Hogue (1986) de-
scribed four additional species in the genera
Dagus and Diedrops. In the latter paper, the
third-instar larva and puparium of Diedrops
roldanorum were described, the first im-
matures known for the tribe. Since publi-
cation of these papers, numerous additional
specimens of Dagini have been made avail-
able to us, including several that represent
new species in the genera Psilephydra and
Dagus. Descriptions of these new species are
presented here. The addition of new species
to Psilephydra also necessitates some
changes to the characterization of that ge-
nus. In addition to the descriptions, we pre-
sent revised keys to the genera of Dagini
and to the species of the genera with new
species and a catalog of the tribe.
In this paper we essentially follow the
methods and format presented in the above-
cited papers. Those works should be con-
sulted for additional details and perspec-
tive.
Two head and two venational ratios are
used commonly in the descriptions and are
defined here for the convenience of the user
(ratios are based on measurement of three
specimens if available).
Eye-to-cheek ratio: Genal height (im-
mediately below the eye)/eye height.
Eye width-to-face length ratio: Face length
(in profile from anterior margin of eye to
anterior margin of face)/eye width (greatest
horizontal distance along plane of eye).
Costal vein ratio: The straight line dis-
tance between the apices of R,,, and R,, ;/
distance between the apices of R, and R,,;.
M vein ratio: The straight line distance
along M basad of crossvein dm-cu/distance
distal to crossvein dm-cu.
Acronyms used in the text to indicate de-
positories of specimens are as follows: BPBM
VOLUME 90, NUMBER 1
(Bernice P. Bishop Museum, Honolulu, Ha-
wail); HNHM (Hungarian Natural History
Museum, Budapest, Hungary); ZMC (Zoo-
logical Museum, Copenhagen, Denmark);
USNM (National Museum of Natural His-
tory, Smithsonian Institution, Washington,
DIE):
KEY TO THE GENERA AND SPECIES
GROUPS OF DAGINI
1. Pulvilli lacking; postpronotum with | to a few
setulae Dagus Cresson
— Pulvilli present, conspicuous; postpronotum
bare sO at phar 2
2. Distance fiend apices ory veins ims A yand Ros
short, less than half distance between veins R,, ;
and M; gena high, equal to or greater than eye
height; genal seta well developed and conspic-
uous; prescutellar acrostichal setae well devel-
oped; propleuron setulose; 5th tarsomere with
dorsoapical process extended beyond base of
tarsal claws . Diedrops Mathis and Wirth
— Distance between apices of veins R,,,and R,,;
subequal to that between veins R,,, and M;
gena short, usually not more than ' eye height;
genal seta, if present, weakly developed and
inconspicuous; prescutellar acrostichal setae not
evident; propleuron without setulae; 5th tar-
somere not as above ....... 3
3. Two to 3 large, postsutural dorsocentral setae;
arista mostly bare, at most with small hairs
(their lengths less than aristal width at base)
along basal 14 (Psilephydra Hendel) : 4
— One large, postsutural dorsocentrai seta in-
serted near scutellum; arista pectinate or mac-
ropubescent along at least basal 74 (Physemops
G@résson)) .:-2.-
4. Anterior notopleural seta weakly developed:
much smaller than the posterior seta; fore fe-
mur with posteroventral row of short, spine-
IKI ET Cetra reno neers the cyanoprosopa group
— Anterior notopleural seta well developed, sub-
equal in length to posterior seta; fore femur
unarmed, lacking spine-like setae
NMS ogra syne cies .the fluvialis group
5. Halter capitellum black; ocellar bristles lack-
ing; arista long, over twice combined length of
first 3 antennal segments; vein CuA, along pos-
terior margin of discal cell bowed posteriorly
rae eres the nemorosus group
- Halter capitellum pale, usually yellowish; ocel-
lar setae present, conspicuous; arista shorter,
rarely not over twice combined length of first
3 antennal segments; vein CuA, along posterior
margin of discal cell straight ... the panops group
nn
107
Genus Psilephydra Hendel
Psilephydra Hendel 1914: 99. Type species:
Psilephydra cyanoprosopa Hendel 1914,
by original description (see catalog sec-
tion, p. 120, for a more complete synon-
ymy).
Diagnosis.— Moderately small to medi-
um-sized shore flies, length 2.0 to 3.4 mm.
Head: Frons wide (width-to-length ratio
2.3-2.7), fronto-orbital setae 2-4, often mi-
nute; ocellar setae present; both inner and
outer vertical setae present. Arista moder-
ately long, length nearly twice to three times
length of Ist flagellomere, apex virtually
bare, basal 73 with some dorsal, minute set-
ulae or nearly bare; Ist flagellomere longer
than pedicel but not twice length of latter;
face generally shield-like, either shallowly
and uniformly protrudent over entire height
or with lower * slightly but distinctly more
protrudent (best seen in profile), sparsely
covered by setae and densely microtomen-
tose with coloration metallic silvery to
bronzish; eye-to-cheek ratio 0.30-0.66: ge-
nal setae usually present. Palpus elongate,
dark colored.
Thorax: Mesonotum and pleura subshin-
ing to shining; anterior notopleural seta sub-
equal or much smaller than posterior seta;
dorsocentral setae 2-5; prescutellar acros-
tichal setae lacking; postalar seta |, well de-
veloped; scutellar setae with anterior pair
usually smaller, although length variable as
compared to apical pair; propleuron bare;
katepisternal seta subequal or smaller in
length than posterior anepisternal seta. Hal-
ter whitish to yellowish. Wing hyaline or
uniformly lightly darkened; costal vein ratio
0.16-0.23; M vein ratio 0.51-0.73. Dis-
tance between veins R,,, and R,,,; about
equal to that between veins R,,; and M.
Legs blackish; fore femur thickened, with or
without posteroventral row of short, spine-
like setae.
Male genitalia: Epandrium shield-like,
forming cercal activity dorsally; surstyli
either distinct or apparently fused to ventral
108
margin; gonite in lateral view with slender,
ventral and anterior projections; aedeagal
apodeme long and very slender; aedeagus
either about as wide as long or longer than
wide.
Discussion.— We have divided Psilephy-
dra into the fluvialis and cyanoprosopa
groups, as characterized in the above key or
following diagnoses. Eventually, each may
be recognized as a separate genus because
of the many differences between them (see
species group diagnoses). Although our key
to the species of the genus includes all known
species, we are providing descriptions for
the new species only.
Key TO SPECIES OF PSILEPHYDRA
HENDEL
1. Anterior notopleural seta weak, much smaller
than the posterior seta; fore femur with pos-
teroventral row of short, spine-like setae
— Anterior notopleural seta well developed, sub-
equal to posterior seta; fore femur unarmed,
lacking spine-like setae mua:
Mesonotum dark blue; trochanters dark: eye-
to-cheek ratio more than 0.5 (India)
: P. lyneborgi Zatwarnicki, new species
— Mesonotum dark brown; trochanters pale, yel-
lowish to orangish; eye-to-cheek ratio less than
OLS) crea JUdoH GoORR GRID DEeoE aucun edoce 3
3. First flagellomere pale; “scutellum flat: wings
uniformly darkened; costa not thickened (Thai-
land) _.P. iridescens Zatwarnicki, new species
— First flagellomere blackish, concolorous with
scape and pedicel; scutellum convex; wings
hyaline; costa thickened (Taiwan)
ane P. cyanoprosopa Hendel
4. Face distactly bicolored and protrudent on
ventral 2, dorsal surface of protrusion metallic
blue, ventral portion along oral margin densely
microtomentose, whitish gray (Nepal) ..
oe _ P. kaskiensis Mathis, new species
- Face either unicolorous or with colorational
changes gradual, anterior surface of ventral '/2
nearly flat
. Dorsocentral setae 5, Gnterior seta presutural:
overall length 2.9 to 3.4 mm (Nepal) .
_ P. nepalensis Mathis, new species
- Dorsocentral setae 3, all postsutural; overall
length 2 to 2.6 mm (Japan and Ryukyu Islands)
operas P. fluvialis (Miyagi)
The fluvialis Group
Diagnosis.—Genal seta well developed,
conspicuous; anterior notopleural seta well
Nm
to
wn
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
developed, subequal to posterior seta, pos-
terior seta inserted at elevated level com-
pared with anterior seta; katepisternal seta
well developed, subequal in size to anepi-
sternal seta; fore femur unarmed, lacking
spine-like setae.
Species included.—Psilephydra_fluvialis
(Miyagi), P. kaskiensis Mathis, and P. ne-
palensis Mathis.
Psilephydra kaskiensis Mathis,
NEw SPECIES
Figs. 1-2
Description. — Moderately small shore
flies, length 2.50 to 2.90 mm.
Head: Fronto-orbits and mesofrons bare,
shining, with bronzish brown metallic lus-
ter; parafrons slightly duller, with sparse
microtomentum; fronto-orbital setae 3,
posterior 2 larger and subequal in size. Aris-
ta with minute hairs, length slightly more
than 3 x length of Ist flagellomere. Face dis-
tinctly bicolored and protrudent on ventral
>, dorsal surface or protrusion metallic blue,
ventral portion along oral margin densely
microtomentose, whitish gray. Eye-to-cheek
ratio 0.30; genal seta well developed and
conspicuous.
Thorax: Mesonotum and pleural sclerites
mostly shining, with dark brown, metallic
luster, only propleuron densely microto-
mentose, with grayish coloration. Anterior
notopleural seta well developed, subequal
to posterior seta, posterior seta inserted at
distinctly higher level than anterior seta;
dorsocentral setae 3, 2 larger posterior setae
(including posteriormost, slightly laterally
displaced seta) and a smaller anterior seta,
anterior seta either sutural or postsutural;
basolateral scutellar seta moderately long,
about '2 length of apical seta; katepisternal
seta well developed, subequal in size to pos-
terior anepisternal seta. Costal vein ratio
0.16; M vein ratio 0.73. Legs, including tro-
chanters, entirely blackish; fore femur un-
armed, lacking spine-like setae; setulae on
hind coxal strap variable (present on ho-
lotype).
Abdomen: Male genitalia as in Figs. 1-2
VOLUME 90, NUMBER |
109
Figs. 1-4. Psilephydra kaskiensis. 1, Male genitalia, posterior view. 2, Male genitalia, lateral view. Psilephydra
nepalensis. 3, Male genitalia, posterior view. 4, Male genitalia, lateral view.
epandrium in posterior view about as high
as wide, dorsal margin rounded, ventral
margin somewhat truncate; surstyli inserted
medially at ventral margin of epandrium,
each a ventrally projected, linear, more or
less sinuate process that is foot-like and set-
ulose apically; gonite in lateral view a
2-pronged process, basal 74 of ventral pro-
cess gradually enlarging from base to apex,
thereafter forming a hook-like apex with
rounded emargination of anterior surface,
anterior process narrowly linear and more
or less parallel sided, joined anteriorly with
similar process from opposite side; aedeagal
apodeme a broadly formed plate-like pro-
cess lying between anterior gonal processes
and base of aedeagus; aedeagus apparently
lacking or greatly reduced, mostly membra-
nous; aedeagal apodeme reduced, a well-
sclerotized V- to Y-shaped structure that is
attached to hypandrium.
Type material.— The holotype male is la-
110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
beled ‘“SNEPAL.Kaski Dist[rict] Chomrung,
Sinuwa[,] 2250 m, 22 Oct 1985[,] Wayne
N. Mathis.” The allotype female and three
additional paratypes (2¢, 12; USNM) bear
the same label data as the holotype. The
holotype is double mounted (minute nadel
in plastic elastomere block), is in good con-
dition (abdomen removed, dissected, pre-
served in glycerine in an attached micro-
vial), and is deposited in the National
Museum of Natural History (USNM),
Smithsonian Institution.
Distribution. —This species is known only
from the type series from Nepal.
Etymology.—The specific epithet, kas-
kiensis, alludes to the district in Nepal where
this species was collected.
Remarks.—This species, although simi-
lar to P. nepalensis and P. fluvialis, 1s readily
distinguished from its congeners by the dis-
tinctly bicolored and ventrally protrudent
face, shining thorax, especially the pleural
area, smaller size, and structures of the male
genitalia.
In a few external features, this species is
quite similar to those of the genus Dagus,
the nominate genus of the tribe Dagini (see
discussion under that genus below). The
protrudent and rounded lower face (best seen
in profile) is especially like Dagus.
Psilephydra nepalensis Mathis,
New SPECIES
Figs. 3-4
Description.— Moderately small to me-
dium-sized shore flies, length 2.90 to 3.40
mm.
Head: Frons subshining, moderately in-
vested with microtomentum; mesofrons
undifferentiated; fronto-orbital setae 3, pos-
terior 2 larger and subequal in size. Arista
with minute hairs, length slightly more than
3x length of Ist flagellomere. Facial pro-
trusion occupying most of face, protrusion
not limited to ventral ': face generally mi-
crotomentose, becoming more densely so
ventrally; facial coloration dark gray dor-
sally to whitish gray ventrally, colorational
change gradual. Eye-to-cheek ratio 0.40; ge-
nal seta well developed, conspicuous.
Thorax: Mesonotum moderately invest-
ed with microtomentum, subshining, dark
brown; pleural sclerites from anepisternum
ventrad more densely microtomentose than
mesonotum, microtomentum gray. Ante-
rior notopleural seta well developed, sub-
equal to posterior seta, posterior seta in-
serted at distinctly higher level than anterior
seta; dorsocentral setae 5, 3 larger posterior
setae (including posteriormost, slightly lat-
erally displaced seta) and 2 smaller anterior
setae, anterior setae either sutural or pre-
sutural; basolateral scutellar seta moderate-
ly long, about '2 length of apical seta; kat-
episternal seta well developed, subequal in
size to posterior anepisternal seta. Costal
vein ratio 0.16; M vein ratio 0.58. Legs,
including trochanters, entirely dark colored,
blackish; fore femur unarmed, lacking spine-
like setae; hind coxal strap bearing | ventral
setula.
Abdomen: Male genitalia (Figs. 3-4) as
follows: epandrium and surstyli in posterior
view more or less rectangular; surstyli evi-
dent as broadly formed lobes that are fused
to the ventral margin of the epandrium;
gonite in lateral view a large inverted
U-shaped, well-sclerotized process, the an-
teroventral arm hook-like; aedeagus rough-
ly rectangular, anterior margin concave with
pointed angles ventrally; aedeagal apodeme
poorly sclerotized, a Y-shaped process.
Type material.—The holotype male is la-
beled ““NEPAL.Kaski Dist[rict] Chomrung,
Sinuwa[,] 2250 m, 22 Oct 1985[,] Wayne
N. Mathis.” The allotype female and one
male paratype (USNM) bear the same label
data as the holotype. The holotype is double
mounted (minute nadel in plastic elasto-
mere block), is in good condition, and is
deposited in the National Museum of Nat-
ural History (USNM), Smithsonian Insti-
tution.
Distribution. — This species is known only
from the type series from Nepal.
Etymlogy.—The specific epithet, nepa-
VOLUME 90, NUMBER 1 111
Figs. 5-9. Psilephydra iridescens. 5, Male genitalia, posterior view. 6, Gonite and aedeagal apodeme, lateral
view. Psilephydra lyneborgi. 7, Male genitalia, posterior view. 8, Gonite, lateral view. 9, Aedeagal apodeme and
aedeagus, lateral view.
112
lensis, alludes to the country where this
species was collected.
Remarks.—This species is similar to P.
fluvialis and to a lesser extent to P. kas-
kiensis but may be distinguished from both
and other congeners by its larger size, facial
coloration, number of large dorsocentral se-
tae, and structures of the male genitalia.
The cyanoprosopa Group
Diagnosis.—Genal seta lacking or re-
duced; anterior notopleural seta weakly de-
veloped, much smaller than posterior seta:
fore femur bearing short, spine-like setae
along posteroventral margin.
Species included.—Psilephydra cyano-
prosopa Hendel, P. lyneborgi Zatwarnicki,
and P. iridescens Zatwarnicki.
Psilephydra iridescens Zatwarnicki,
New SPECIES
Figs. 5-6
Description.— Medium-sized shore flies,
length 3.0 mm.
Head: Frons subshining, dark brown:
mesofrons undifferentiated; fronto-orbital
setae 2, minute. First flagellomere pale, only
darkened above; arista with minute hairs
above, length slightly more than 3 x length
of Ist flagellomere. Facial protrusion oc-
cupying most of face, face dull black with
generally metallic silvery microtomentum,
gradually becoming metallic bronzish me-
dially; eye-to-cheek ratio 0.46; genal setae
poorly developed, but evident. Palpus black.
Thorax: Mesonotum moderately invest-
ed with microtomentum, subshining, dark
brown; pleura from anepisternum ventrad
more densely microtomentose than meso-
notum, microtomentum silvery gray; an-
terior notopleural seta minute, posterior seta
well developed, subequal to anepisternal
seta, posterior seta inserted at same level as
anterior seta; dorsocentral setae 2, poste-
riormost seta slightly displaced laterally, seta
well developed, length of anterior seta about
Y. posterior seta. Scutellum flat, broadly
rounded with small projection; basolateral
scutellar seta minute, about 1.4 length of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
apical seta. Halter yellowish. Wing uni-
formly darkened; costal vein ratio 0.21; M
vein ratio 0.57. Legs dark colored, blackish;
tarsi pale, darkened apically; trochanters and
apices of tibiae brown; fore femur bearing
posteroventral row of 4, short, spine-like
setae.
Abdomen: Terga concolorous with meso-
notum, invested laterally with whitish mi-
crotomentum; dorsal surface of terga 2-5
sparsely pitted. Male genitalia (Figs. S—6) as
follows: epandrium in posterior view as an
inverted U, anteroventral margin truncate;
surstyli evident as lobes with process in the
middle of its anterior margin; gonite in pos-
terior view with anterior process elongate,
posteroventral process apically rounded,
arched anteriorly, and directed ventrally;
aedeagal apodeme in posterior view
Y-shaped, in lateral view C-shaped, weakly
broader centrally; aedeagus in posterior view
roughly ovate, rounded posteroapically, an-
terior apex gradually tapered.
Type material.—The holotype male is la-
beled ““THAILAND: S. Banna. Nakhon{,]
108 m.[,] V-5-10-[19]'58/T. C. Maa Collec-
tor[,] Nr. 406.” The holotype is glued to a
small paper rectangle, is in good condition
(apex of abdomen removed, dissected, and
in an attached microvial), and is deposited
in the Bernice P. Bishop Museum, Hono-
lulu, Hawaii.
Distribution. — This species is only known
from the holotype from Thailand.
Etymology.—The specific epithet, irides-
cens, alludes to the shimmering coloration
of the face of this species.
Remarks.—This species is similar to P.
cyanoprosopa but may be distinguished from
it and other congeners by its pale Ist flag-
ellomere, darkened wings, and structures of
the male genitalia.
Psilephydra lyneborgi Zatwarnicki,
NEw SPECIES
Figs. 7-9
Description. — Medium-sized shore flies,
length 3.0 to 3.30 mm.
Head: Frons dark brown; fronto-orbits
VOLUME 90, NUMBER 1!
and mesofrons bare, shining; parafrons
slighty duller, invested with sparse micro-
tomentum. Fronto-orbital setae 3, minute,
posterior 2 larger and subequal in size. Aris-
ta with minute hairs above, length slightly
more than 3 Ist flagellomere. Facial pro-
trusion occupying most of face; face dull
black with metallic silvery blue microto-
mentum, becoming metallic bronzish dor-
sally; eye-to-cheek ratio 0.66, genal seta
poorly developed, but evident. Palpus black.
Thorax: Mesonotum mostly dark, shin-
ing, covered with sparse, dark brown micro-
tomentum; pleura with dense, silvery-gray-
ish microtomentum, posterior notopleural
seta well developed, its length more than
twice length of anterior seta, posterior no-
topleural bristle inserted at about same level
as anterior seta; dorsocentral setae 2, pos-
teriormost seta slightly displaced laterally,
well developed, length about twice that of
anterior setae. Scutellum flat, broadly
rounded with distinct projection; basolater-
al scutellar seta minute; apical scutellar seta
well developed, subequal to posteriormost
dorsocentral seta. Length of anepisternal seta
about 1.5 length of posterior notopleural
seta. Wing hyaline; costal vein ratio 0.17;
M vein ratio 0.54. Halter yellowish white.
Legs dark colored, blackish, only metatar-
sus yellowish; fore femur with posteroven-
tral row of 7, short, spine-like setae.
Abdomen: Terga concolorous with meso-
notum, invested laterally with whitish mi-
crotomentum, dorsal surface of terga 2-5
densely pitted. Male genitalia (Figs. 7-9) as
follows: epandrium in posterior view as an
inverted U, with arms wider toward ante-
rior margins; surstyli fused to broad an-
teroventral margin of epandrium, hemi-
spherical in shape, dorsal margin concave;
gonite in posterior view weakly S-shaped,
apex conspicuously wide and obtuse, ante-
rior process in lateral view with broad and
rounded apex, posteroventral margin with
sharp process; aedeagal apodeme band-like,
bent doubly in lateral view, Y-shaped in
posterior view; aedeagus in posterior view
longer than broad, basal half with folds lat-
13
erally, anterior margin tapered gradually
with obtuse apex, ventral margin in lateral
view rounded, dorsal margin roughly
creased, anterior margin truncate, antero-
ventral apex forming nose-like process.
Type material.— The holotype male is la-
beled “S. India: Karnataka. Kemmangudi,
1200-1500 m[,] 11-16. x1 1977[,] Zool. Mus.
Copenhagen Exp.” The allotype female and
one additional paratype female (ZMC) are
labeled “India (Uttar Pradesh)[,] Dehra Dun
Valley, c. 700 mf[,] 4.-13. vii 1978[,] Co-
penhagen Zool. Mus. Exp.” The holotype is
double mounted (minute nadel in plastic
elastomere block), is in good condition (ab-
domen removed, dissected, and in attached
microvial), and is deposited in the Zoolog-
ical Museum in Copenhagen.
Distribution. — This species 1s known only
from the type series from India.
Etymology.—The specific epithet, /vne-
borgi, is a genitive patronym to honor Dr.
L. Lyneborg, who has generously supported
our work on shore flies.
Remarks.—This species is similar to P.
cyanoprosopa but may be distinguished from
it and other congeners by the relatively high
gena, dark blue mesonotal color, scutellar
protrusion, and structures of the male gen-
italia.
Genus Dagus Cresson
Dagus Cresson 1935: 345. Type species:
Ephyadra rostrata Cresson 1918, by orig-
inal designation.— Wirth 1968: 24 (neo-
tropical catalog).—Mathis 1982: 20-23
(review), 1983:717-726 (revision).
Phylogenetic considerations.—The dis-
covery and study of two new species in Da-
gus and four new species in Psi/ephydra has
provided additional information concern-
ing the phylogeny of these taxa. Mathis
(1983) suggested previously that Dagus was
the sister group of Physemops, primarily
based on the elevated insertion of the pos-
terior notopleural seta. That character state
was then known only to species in these two
genera. Two of the new species of Psilephy-
114
dra, P. nepalensis and P. kaskiensis, also
have an elevated insertion of that seta, and,
moreover, the face of P. kaskiensis is pro-
trudent in a similar way to specimens of
Dagus. Discovery of these character states
casts considerable doubt on the sister-group
relationship between Dagus and Physemops
and indicates that such a relationship may
exist between Dagus and Psilephydra, es-
pecially between the fluvialis group of the
latter genus.
Although the sister group to Dagus 1s not
clearly demonstrated, the monophyly of
Dagus is not questioned in view of the evi-
dence. Character evidence to establish this
hypothesis was elaborated by Mathis (1983:
718), and we add to the characters he listed
the unique condition of the epandrium,
which does not extend around the dorsal
margins of the cerci. Typically, the dorsal
margin of the epandrium forms a connec-
tion around the cerci, forming an oval to
circular opening in which the cerci and anal
opening are situated. We must also note that
the third character listed by Mathis, the
ventral protrusion of the lower half of the
face, is considerably weakened, as that char-
acter, which was used to substantiate the
monophyly of Dagus, also occurs in Psile-
phydra (P. kaskiensis).
Among taxa clearly belonging to Dagus,
the two new species described here are
closely related, and together they form a
monophyletic lineage, i.e. they are sister
species. Character evidence to support this
hypothesis is as follows (autapomorphic
characters): fore tibia with a preapical, ven-
tral tuft of long setae; and tarsomeres 4 and
5 of foreleg and tarsomere 5 of hind leg with
large, ventral, scale-like setae.
Discussion. — The diagnosis of Dagus that
Mathis (1983) presented remains accurate
and is not repeated here. The addition of
two new species, however, necessitates a re-
vised key, presented below, to facilitate the
identification of species
Key TO SPECIES OF DAGuS
1. Arista long, 3-4 length of Ist flagellomere,
conspicuously haired, length of longer hairs
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
much greater than aristal width at base
D. trichocerus Mathis
— Arista shorter, at most 2-3 x length of Ist flag-
ellomere, hairs barely evident, length less than
aristalrwidthvatbase’ so. s.. 0+ anes ene 2
Posterior notopleural seta inserted at about
same level as anterior seta; genal seta well de-
veloped, subequal in size to anterior fronto-
orbital seta .. D. dominicanus Mathis, new species
— Posterior notopleural seta inserted conspicu-
ously above level of anterior seta, usualy 2-3 x
higher; genal seta weak, usually conspicuously
smaller than anterior fronto-orbital seta
3. Gena short, about 3 eye height; specimens short,
less than 2.25mm ...... D. rostratus (Williston)
— Gena high, '2 or more of eye height; specimens
longer, usually greater than 2.25 mm
4. Fore tibia with preapical tuft of long setae ven-
trally; facial setae generally weak, those along
oral margin especially so, usually less than
length of anterior fronto-orbital seta ........
(age D. spanglerorum Mathis, new species
— Fore tibia lacking preapical tuft of long setae
ventrally; facial setae generally well developed,
those along oral margin and usually 1-2 setae
at lateral margin of facial prominence longer
than anterior fronto-orbital seta
to
D. wirthi Mathis
Dagus spanglerorum Mathis,
New SPECIES
Figs. 10-12
Description. — Medium-sized shore flies,
length 3.2 to 4.0 mm.
Head: Frons entirely densely microto-
mentose, appearing velvety, blackish brown
to black. Arista bearing minute hairs, barely
evident, aristal length nearly 3 times length
of Ist flagellomere. Facial protrusion in lat-
eral view with anterodorsal surface less
acutely angulate with oral margin; dorsal '2
of facial protrusion with bare, shining areas
just below antennae dark, metallic blue,
otherwise face micromentose, dark brown;
facial setae generally weakly developed, es-
pecially setae along oral margin, these
smaller than ocellar setae; eye-to-cheek ra-
tio 0.47; eye width-to-face length ratio 0.66;
genal seta weakly developed, smaller than
anterior fronto-orbital seta.
Thorax: Thoracic chaetotaxy moderately
well developed. Dorsocentral setae with 4
larger setae, all postsutural, and 2-3 smaller
VOLUME 90, NUMBER 1
Figs. 10-12.
(cerci and epandrium), lateral view. 12, Internal male genitalia, lateral view.
presutural setae; postsutural intra-alar setae
with 3-5 generally small setae; posterior no-
topleural seta inserted at distinctly more el-
evated position than anterior seta, nearly
twice distance from ventral notopleural
margin than anterior seta; hind coxal strap
bare. Fore tibia with ventral, preapical tuft
of well-developed setae; tarsomeres 4 and
5 of foreleg and tarsomere 5 of hind leg
bearing large, scale-like setae along ventral
surface. Costal vein index 0.13; M vein in-
dex 0.70.
Abdomen: Male genitalia (Figs. 10-12) as
follows: epandrium, in posterior view,
roughly triangular, ventral apex conspicu-
ously cleft, projection on either side of cleft
mucronate; lateral view of epandrium near-
ly bullet shaped with dorsal margin some-
what truncate and remainder tapered grad-
ually to pointed ventral apex; gonite shorter
than aedeagus, roughly triangular, longer
115
Dagus spanglerorum. 10, Male genitalia (cerci and epandrium), posterior view. | 1, Male genitalia
than wide, dorsal margin uneven, with a
shallow projection about '4 distance from
base.
Type material.— The holotype male is la-
beled “DOMINICAN REPUBLIC[.] La
Vega Province[:] Constanza (3.5 Km S)[,] 9
Nov 1984, sweeping P. & P. Spangler & R.
Faitoute.”” The allotype female and 104
paratypes (326, 732: USNM) bear the same
label data as the holotype. Other paratypes
are as follows: DOMINICAN REPUBLIC.
La Vega Province: Constanza (8.5 km S), 9
Nov 1984, P. and P. Spangler, R. Faitoute
(138, 389: USNM); Constanza (12 km S) 9
Nov 1984, P. and P. Spangler, R. Faitoute
(12: USNM). The holotype is double
mounted (minute nadel in plastic elasto-
mere block), is in good condition, and is
deposited in the National Museum of Nat-
ural History (USNM), Smithsonian Insti-
tution.
116
Fig. 13.
Distribution. — This species is known only
from the Dominican Republic (Greater An-
tilles: Hispaniola).
Etymology.—The specific epithet, span-
glerorum, 18 a genitive, pleural patronym to
recognize the collecting efforts of Paul and
Phyllis Spangler, who, along with Robin
Faitoute, collected the known specimens of
this species as well as numerous other species
of Ephydridae.
Remarks.—This species is most easily
distinguished from its congeners, especially
D. dominicanus, by its larger size (smaller
than D. dominicanus but larger than other
congeners), the preapical, ventral tuft of long
setae on the fore tibia (also present in D.
dominicanus), the nearly bare arista, the re-
duced genal seta, the elevated insertion of
the posterior notopleural seta, and the char-
acters of the male genitalia.
Dagus dominicanus Mathis,
New SPECIES
Figs. 13-19
Description.— Medium-sized to large
shore flies, length 3.8 to 5.25 mm.
Head: Frons entirely densely microto-
mentose, appearing velvety, blackish brown
to black. Arista bearing minute hairs, barely
evident, aristal length nearly 3 times length
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
0.1mm
Dagus dominicanus. 13, Cephalopharyngeal skeleton of third-instar larva, lateral view.
of Ist flagellomere. Facial protrusion in lat-
eral view with anterodorsal surface less
acutely angulate with oral margin; dorsal '2
of facial protrusion with bare, shining area
just below antennae dark, metallic blue to
brown, becoming microtomentose ventral-
ly, dark brown to gray along oral margin;
facial setae generally well developed, espe-
cially setae along oral margin, these sub-
equal to ocellar setae; eye-to-cheek ratio
0.55; eye width-to-face length ratio 0.64;
genal seta well developed, subequal to an-
terior fronto-orbital seta.
Thorax: Thoracic chaetotaxy well devel-
oped. Dorsocentral setae with 5 larger setae,
anterior pair presutural to sutural, and 2-3
smaller presutural setae; postsutural in-
tra-alar setae with 3-5 moderately well-de-
veloped setae, all smaller than largest dor-
socentral setae; posterior notopleural seta
inserted only slightly above level of anterior
seta; hind coxal strap bare. Fore tibia with
ventral, preapical tuft of well-developed se-
tae; tarsomeres 4 and 5 of foreleg and tar-
somere 5 of hind leg bearing large, scale-
like setae along ventral surface. Costal vein
index 0.09; M vein index 0.88.
Abdomen: Male genitalia (Figs. 18-19) as
follows: epandrium, in posterior view,
roughly rectangular but with very wide me-
117
VOLUME 90, NUMBER |
7.
a fyabbbe
Seyiet 79F CEEL CG
\
Roery »?
79 9) WY EL
Ye da,
~My
SY aro) Fy v YOY UY wy WPL Se
got Y
Yow hr Wy
Supa
WY
Ws:
Dorsal View. 16, Lateral view. 17, En-
Ventral view. 15
Dagus dominicanus, puparium. 14
Figs. 14-17.
largement of 3rd welt (see arrow on Fig. 16).
118
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 18-19. Dagus dominicanus. 18, Male genitalia (cerci and epandrium), posterior view. 19, Male genitalia
(cerci, epandrium, and internal genitalia), lateral view.
dian cleft ventrally, producing 2 large, lat-
eral, broadly rounded projections and 2
much smaller, medial, pointed projections
within larger cleft, medial margins of larger
processes and lateral margin of inner pro-
jections densely setulose; lateral view of
epandrium somewhat bullet shaped but with
ventral margin broadly rounded; gonite
about as long as aedeagus, basal portion
roughly rectangular and with a long, narrow,
digitiform process extended apically; ae-
deagus gently curved and tapered to round-
ed tip.
Cephalopharyngeal skeleton of third-
instar larva (Fig. 13): Mandible lacking;
hypopharynx in lateral view roughly rec-
tangular but with knob-like process antero-
dorsally and posteroventral angle truncate;
ocular depression poorly developed; dorsal
cornu evenly tapered to apical point: ventral
cornu spatulate.
Puparium (Figs. 14-17): Dimensions:
length 4.45-4.70 mm, width 1.65-1.75 mm;
height 1.5 mm. Shape: generally oval in
ventral or dorsal views (Figs. 14, 15) with
7 ventrolateral, rounded welts forming a
crenulate lateral margin, each welt fringed
with short setulae (welts probably used for
locomotion); retreated margins between
welts extended dorsally as shallow furrows
that become weaker dorsally; in lateral view
dome-shaped (Fig. 16); dorsum gently and
evenly rounded, venter nearly flat; anterior
spiracle with 4 small papilla-like projec-
tions: respiratory tubes essentially fused to-
gether externally.
Type material.—The holotype male is la-
beled “DOMINICAN REPUBLIC[.] La
Vega Province[:] Constanza (12Km S) 9
November 1984[,] P. Spangler & R. Fai-
toute.’ The allotype female bares the same
label data as the holotype. Other paratypes
are as follows: DOMINICAN REPUBLIC:
La Vega Province: Constanza (10 km S), 10
VOLUME 90, NUMBER 1
Nov 1984, P. Spangler, R. Faitoute (4¢, 12;
USNM); Jarabacoa, 13 Nov 1984, P. and
P. Spangler, R. Faitoute (12; USNM). The
holotype is double mounted (minute nadel
in plastic elastomere block), is in good con-
dition, and is deposited in the National Mu-
seum of Natural History (USNM), Smith-
sonian Institution.
Distribution.— All known specimens of
this species are from the Dominican Re-
public (Greater Antilles: Hispaniola).
Etymology.—The specific epithet, do-
minicanus, alludes to the Dominican Re-
public, the country from which the species
was collected.
Remarks.—This species is most easily
distinguished from its congeners, especially
D. spanglerorum, by its larger size (it is the
largest species thus far known in the genus),
the preapical, ventral tuft of long setae on
the fore tibia (also present in D. spangle-
rorum), the nearly bare arista, the well-de-
veloped genal seta, the insertion of the pos-
terior notopleural seta at nearly the same
level as the anterior seta, and the characters
of the male genitalia.
The third-instar larvae and puparia were
collected 10 km south of Constanza (9 No-
vember 1984) by P. Spangler and R. Fai-
toute. The habitat was a steep hillside with
a seepage area in a shallow, V-shaped
depression just above the road cut. Some of
the area had a covering of algae.
Catalog of Genera and Species of Dagini
Tribe DAGINI Mathis 1982
Genus DAGUS Cresson
DAGUS Cresson 1935: 345. Type species:
Ephydra rostrata Cresson 1918, orig.
des.— Wirth 1968: 24 [neotropical cata-
log].—Mathis 1982: 20-23 [review],
Mathis 1983: 717-726 [revision].
dominicanus Mathis 1988: 116.
Type locality). DOMINICAN REPUB-
LIC. La Vega: Constanza (12 km S).
Distribution: Dominican Republic.
rostratus Cresson 1918: 66 (Ephydra), 1935:
119
346 [combination, designated as type
species of Dagus].— Wirth 1968: 28 [neo-
tropical catalog].—Mathis 1982: 21-23
[review, lectotype designation]; 1983:
720-722 [revision].
pygmaea Williston 1896: 402 (Ephydra)
[preoccupied, Haliday 1833].
Type locality: WEST INDIES: Saint Vin-
cent: Perseverance Valley.
Distribution: West Indies (Cuba, Domin-
ica, Jamaica, Saint Vincent) and Mexico
south through Guatemala and Costa Rica
to Venezuela and Brazil.
spanglerorum Mathis 1988: 114.
Type locality; DOMINICAN REPUB-
LIC. La Vega: Constanza (3.5 km S).
Distribution: Dominican Republic.
trichocerus Mathis 1983: 724-725.
Type locality: CUBA. Pinar del Rio: So-
roa.
Distribution: Cuba, Dominican Republic.
wirthi Mathis 1983: 722-724.
Type locality: JAMAICA, Port Parish.
Distribution: Jamaica.
Genus DIEDROPS Mathis and Wirth
DIEDROPS Mathis and Wirth 1976: 126.
Type species: Diedrops aenigma Mathis
and Wirth 1976, orig. des.— Mathis 1982:
6-9 [review].—Mathis 1984: 349-353
{key, notes].
aenigma Mathis and Wirth 1976: 129.—
Mathis 1982: 7-8[review].
Type locality; MEXICO. Michoacan:
Puerto Morillos.
Distribution: Mexico (Michoacan, Sina-
loa).
hitchcocki Mathis and Wirth 1976: 129.—
Mathis 1982: 8-10 [review].
Type locality: PERU. Moquegua: Yacan-
go.
Distribution: Peru.
roldanorum Mathis and Hogue 1986: 23-
26.
120
Type locality: COLOMBIA. Tolima: Bo-
queron (3 km W).
Distribution: Colombia.
steineri Mathis 1984: 351-352.
Type locality: PANAMA. Chiriqui: Bam-
bito (Rio Chiriqui Viejo, 1770 m).
Distribution: Central America (Costa Rica
to Panama).
Genus PH YSEMOPS Cresson
PHYSEMOPS Cresson 1934: 211. Type
species: Psilephydra nemorosa Cresson
1914, orig. des.—Wirth 1968: 20 [neo-
tropical catalog], 1970: 170-177 [re-
key and discussion], 1982: 10-20 [re-
view].
The nemorosus Group
azul Wirth 1970: 172-173.—Mathis 1982:
14 [review].
Type locality: MEXICO. Oaxaca: Valle
Nacional.
Distribution: Mexico (Oaxaca).
nemorosus Cresson 1914: 244 (Psilephy-
dra).—1918: 64 [review, figure of head];
1934: 211 [combination, designated as
type species of Physemops].— Wirth 1968:
20 [neotropical catalog], 1970: 174-175
[review].— Mathis 1982: 14-15 [review].
Type locality: COSTA RICA. Juan Vinas.
Distribution: Circumcaribbean and South
America. Mexico (Oaxaca) and the West In-
dies (Dominica) south through Central
America (El Salvador, Honduras, Nicara-
gua, Costa Rica, Panama) to Ecuador
(Chimborazo) and Brazil (Sao Paulo).
wheeleri Wirth 1970: 176.—Mathis 1982:
16-18 [review].
Type locality: PANAMA. Canal Zone: Las
Cruces Trail.
Distribution: Panama south to Ecuador
(Santo Domingo de los Colorados).
The panops Group
fairchildi Wirth 1970: 173.—Mathis 1982:
18 [review].
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Type locality; PANAMA. Panama: Cerro
Capana.
Distribution: Panama south to Colombia
(Vicinity of Bogota and Medellin).
maldonadoi Wirth 1970: 173-174.—Math-
is 1982: 19-20 [review].
Type locality; PUERTO RICO. Yauco-
Lares Road (km 29).
Distribution: Puerto Rico.
panops Wirth 1970: 175-176.— Mathis
1982: 20 [review].
Type locality: HAITI.
Distribution: Haiti.
Genus PSTILEPHYDRA Hendel
Psilephydra Hendel 1914: 99. Type species:
Psilephydra cyanoprosopa Hendel 1914,
orig. des.—Cresson 1918: 63 [diagnosis,
subfamilial placement].— Mathis and
Wirth 1976: 128 [comparison with Die-
drops].—Cogan and Wirth 1977: 338
[Oriental catalog].—Mathis 1982: 24-28
[review].
The cyanoprosopa Group
cyanoprosopa Hendel 1914: 100.—Cresson
1934: 211 [list].—Cogan and Wirth 1977:
334 [Oriental catalog].— Mathis 1982: 25-
27 [review].
Type locality: TAIWAN. Hoozan.
Distribution: Taiwan.
lyneborgi Zatwarnicki 1988: 112.
Type locality: INDIA. Karnataka: Kem-
mangudi (1200-1500 m).
Distribution: India.
iridescens Zatwarnicki 1988: 112.
Type locality) THAILAND. S. Banna
Nakhon (108 m).
Distribution: Thailand.
The fluvialis Group
fluvialis Miyagi 1977: 88 (Lamproscatel-
la).—Mathis 1982: 27-28 [review].
Type locality; JAPAN. Shikoku Island:
Nametoko, Ehime-ken.
VOLUME 90, NUMBER |
Distribution: Japan (Honshu, Shikoku)
and the Ryukyu Islands (Okinawa-honto).
kaskiensis Mathis 1988: 108.
Type locality; NEPAL. Kaski: Chomrung
(Sinuwa, 2250 m).
Distribution: Nepal.
nepalensis Mathis 1988: 110.
Type locality: NEPAL. Kaski: Chomrung
(Sinuwa, 2250 m).
Distribution: Nepal.
ACKNOWLEDGMENTS
We are grateful to the following curators
and their respective institutions for making
material available to us: N. Evenhius
(BPBM) and L. Lyneborg (ZMC). We also
thank Elaine R. S. Hodges for rendering the
illustrations of the immatures of Dagus
spanglerorum and George L. Venable for
inking the illustrations of Psilephydra kas-
kiensis, P. nepalensis, Dagus spanglerorum,
and D. dominicanus. For critically review-
ing a draft of this paper, we thank Don R.
Harris and R. V. Peterson. Support for field
work in Nepal was provided through a grant
from the Research Opportunity Fund
(Smithsonian Institution).
LITERATURE CITED
Cogan, B. H. and W. W. Wirth. 1977. Family Ephyd-
ridae, pp. 321-339. In M. D. Delfinado and D. E.
Hardy, eds., A Catalog of the Diptera of the Ori-
ental Region, Volume III. Suborder Cyclorrhapha
(excluding Division Aschiza). Honolulu: Univ.
Press of Hawaii.
Cresson, E. T., Jr. 1914. Descriptions of new genera
and species of the dipterous family Ephydridae. I.
Entomol. News 25(6): 241-250.
. 1918. Costa Rican Diptera collected by Phil-
121
ip P. Calvert, Ph.D., 1909-1910. Paper 3. A report
on the Ephydridae. Trans. Amer. Entomol. Soc.
44: 39-68.
. 1934. Descriptions of new genera and species
of the dipterous family Ephydridae. XI. Trans.
Amer. Entomol. Soc. 60: 199-222.
. 1935. Descriptions of new genera and species
of the dipterous family Ephydridae. Trans. Amer.
Entomol. Soc. 61: 345-372.
Hendel, F. 1914. H. Sauter’s Formosa-Ausbeute:
Acalyptrate Musciden (Dipt), III. Suppl. Entomol.
3: 90-117.
Mathis, W. N. 1977. Key to the neotropical genera
of Parydrinae with a revision of the genus Ele/eides
Cresson (Diptera: Ephydridae). Proc. Bio. Soc.
Wash. 90(3): 553-565.
1982. Studies of Ephydrinae (Diptera:
Ephydridae), VI: Review of the tribe Dagini.
Smithson. Cont. Zool. 345: 1-30.
1983. A revision of the genus Dagus Cresson
(Diptera: Ephydridae). Proc. Entomol. Soc. Wash.
85(4): 717-726.
1984. Notes on the shore fly genus Diedrops
(Diptera: Ephydridae). Proc. Entomol. Soc. Wash.
86(2): 349-353.
Mathis, W. N. and C. L. Hogue. 1986. Description
of a new species of the shore fly genus Diedrops
(Diptera: Ephydridae) from Colombia. Cont. Sci.
(Nat. Hist. Mus. Los Angeles County) 377: 21-26.
Mathis, W. N. and W. W. Wirth. 1976. A new neo-
tropical shore fly genus with two new species (Dip-
tera: Ephydridae). Pan-Pac. Entomol. 52(2): 126-
132.
Miyagi, I. 1977. Ephydridae (Insecta: Diptera). Jn
Fauna Japonica. 113 pp.
Williston, S. W. 1896. On the Diptera of St. Vincent
(West Indies). Trans. Entomol. Soc. London 1896:
253-446.
Wirth, W. W. 1968. Family Ephydridae, 77: 1-43.
In Papavero, N., ed., A Catalogue of the Diptera
of the Americas South of the United States. Sao
Paulo: Departmento de Zoologia, Secretaria da
Agricultura, Sao Paulo.
1970. A revision of the neotropical genus
Physemops Cresson (Diptera: Ephydridae). Pan-
Pac. Entomol. 46(3): 170-177.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 122-123
Book REVIEW
Historical Perspective and Current World
Status of the Tomato Russet Mite (Acari:
Eriophyidae). By Thomas M. Perring and
Charles A. Farrar. Entomological Society
of America, Miscellaneous Publication
Number 63, 4603 Calvert Road, College
Park, MD 20740. 14 pp. including | fig-
ure and 5 tables. 1986. $7.50 (non-mem-
bers) $6.75 (book dealers and subscrip-
tion agents); $4.50 (ESA members).
Thomas Perring and Charles Farrar have
writen an excellent review of the Tomato
Russet Mite, Aculops lycopersici (Massee),
an importrant pest on solanaceous crops
throughout many parts of the world. This
information was compiled from a review of
the existing literature on this mite and pre-
sented in the following categories: taxono-
my, morphology, life history and biology,
host range, host-plant interactions, and con-
trol (both biological and chemical).
According to the authors, there was con-
siderable controversy regarding the taxon-
omy and morphology of this mite until 1966
when Aculops lycopersici (Massee) became
the accepted name.
A concise review is given on the synon-
ymy and generic names of the species cre-
ated by Tryon (1917), Massee (1973), Lamb
(1953), and Keifer (1940, 1944, 1959, and
1966). Like others, the authors have mis-
spelled Tryon.
The morphology section is documented
by references and contains a comparison
made by Keifer of specimens described by
Massee (1937) and Keifer (1940). To some
readers, this comparison may serve as an
indication of two distinct species, rather than
variations found within a species. It is not
clear whether Keifer studied Massee’s slide-
mounted specimens or if illustrations were
used in the study. Keifer and the authors
failed to mention that these different ob-
servations could be attributed to different
optical and measuring equipment, mite
clearing agents, and mounting media.
The book contains useful documented
references and pertinent information on life
history and biology, host range, host-plant
interactions, and natural enemies for bio-
logical control.
Among the most valuable features in the
book are the five tables. The style and in-
formation compiled by the authors are com-
mendable. Table 1 presents the worldwide
distribution of 4. /ycopersici, with the per-
tinent dates, locations, authors, and refer-
ences. The earliest known date for the mite
is 1892 where it was first found in Florida
and the latest date 1895 where it was re-
ported from Mexico. Although the authors
cited the locations from original references,
many of the names for countries have
changed and the present correct names
should have been placed in parentheses be-
side the old name. Examples of the old and
present names are: Ceylon (= Sri Lanka)
and Persia (= Iran).
Table 2 gives the distribution of the mite
in the United States with the same heading
as Table 1.
Table 3 is an excellent adjunct resource
that will be useful in identifying the mite.
This table lists the common and scientific
names, dates, and references for 25 plants
known to be hosts of A. /ycopersici.
Table 4 is a list of 75 materials tested for
the control of 4. /ycopersici and includes the
common and trade names, efficacy, date,
and references for each substance.
Table 5 is composed of citations from the
Cooperative Economic Insect Reports and
Cooperative Plant Pest Reports that contain
information on A. /ycopersici. The year, vol-
ume, number, and pages are included for
the years 1952 through 1977.
Unfortunately, the front cover of this book
is inappropriate. The cover has a scanning
electron micrograph of unidentified setae.
VOLUME 90, NUMBER 1
This type of setae has not been reported or
illustrated for any member of the erio-
phyoidea. Surely a picture of any eriophyid
mite on the front cover would lend more
credence to the book.
This book is an excellent synthesis of the
present knowledge on the Tomato Russet
Mite and can be used especially by acarol-
123
ogist and biological control and integrated
pest management researchers.
Robert L. Smiley, Systematic Entomol-
ogy Laboratory, BBII, Agricultural Re-
search Service, USDA, Beltsville, Maryland
20705.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, p. 124
Book REVIEW
The Bombyliidae of Deep Canyon. Parts
Iand II. By Ali B. Tabet and Jack C. Hall.
Al-Fateh University Publications, Trip-
oli, S. P. L.A. J. 1984, Part I, A Phenology
Study of the Bombylidae of Deep Can-
yon, pages [1]-63, figs. 1-17, tabs. I-VI.
Part II, pages [1]-176, figs. 1-63 (Key to
genus Bombylius, pp. 19-21, written by
Neil Evenhuis).
This publication on the Bombyliidae of
the Philip L. Boyd Deep Canyon Desert Re-
search Center at Palm Desert, California, a
facility of the University of California, Riv-
erside, is the sixth major publication re-
sulting from studies at this center. The first,
Mammals of Deep Canyon by R. Mark
Ryan, 1968, was followed by Ants of Deep
Canyon by George C. Wheeler and Jeanette
Wheeler, 1973; Deep Canyon, a Desert Wil-
derness for Science, edited by Irwin P. Ting
and Bill Jennings, 1976; Plants of Deep
Canyon and the Central Coachella Valley,
California by Jan G. Zabriskie, 1979; and
Birds of Southern California’s Deep Can-
yon, by W. W. Weathers, 1983.
Published copies of Part I of The Bom-
byliidae of Deep Canyon were received from
Libya by Mr. Hall, at Riverside, California,
in 1984, while copies of Part II were re-
ceived in March of 1987. It has not been
possible, as of this writing, to ascertain the
exact date of publication of the second part.
Part I, A Phenology Study of the Bom-
byliidae of Deep Canyon, summarizes an
intensive year-long sampling of the Bom-
byliidae of a localized area within the Col-
orado Desert of southern California. Sev-
enty percent of the genera and 20 percent
of the species of Bombyliidae known from
North America are reported from this
13,000 acre (80 square mile) area. Species
composition and seasonal distribution of
mainly two areas of varying elevation in
Deep Canyon, both primarily in the Lower
Sonoran Zone, with habitats such as the
chaparral and coniferous forest areas oc-
curring at higher elevations in the canyon
remain undocumented.
Part II discusses the 186 species of Bom-
byliidae (in 41 genera) collected in the Deep
Canyon study area. A key to the subfamilies
and genera of Bombyliidae of California oc-
cupies pages 7-17. Fifteen species are new
and named as: Mythicomyia candida (p. 43),
M. torta (pp. 49-50), M. vernalis (pp. 50-
51), Oligodranes nigricans (pp. 61-63),
Desmatomyia proboscidea (pp. 64-66),
Aphoebantus albopilosus (pp. 83-85), A.
bilineatus (pp. 85-87), A. leucospilus (pp.
89-92), A. melanomerinyx (pp. 92-95), A.
nigropilosus (pp. 95-97), A. oxypetes (pp.
98-100), A. xanthoscelus (pp. 102-105),
Chrysanthrax albicomus (pp. 135-136), C.
partita (pp. 139-141), C. petalonyx (pp. 141-
143), Neodiplocampta caliginosa (pp. 154—
156). Primary types are deposited 1n the col-
lection of the California Academy of Sci-
ences, San Francisco, with the exception of
one species, Aphoebantus xanthocelus, that
is deposited with the University of Califor-
nia, Davis.
Mr. Jack C. Hall has a limited number of
copies of this publication that are available
for distribution. He may be contacted at the
Division of Biological Control, University
of California, Riverside, California, 92507,
by those so interested.
Paul H. Arnaud, Jr., California Academy
of Sciences, Golden Gate Park, San Fran-
cisco, California 94118.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 125-126
Book REVIEW
Manual of Nearctic Diptera, vol. 2. J. F.
McAlpine, Ed. Research Branch, Agri-
culture Canada, Ottawa. 1987, vi + 658
pp. $68.35.
The long-awaited publication of volume
2 of the widely acclaimed Manual of Nearc-
tic Diptera should be cause for celebration
by North American Dipterists, as it replaces
the long outdated, though memorable, book
by C. H. Curran, Families and Genera of
North American Diptera, published in 1934.
As stated in the preface **.. . the main pur-
pose of the Manual of Nearctic Diptera is
to provide a modern, well-illustrated, easily
interpretable means for identifying the fam-
ilies and genera of two-winged flies of
American north of Mexico.” This goal has
been admirably fulfilled!
Volume | of the projected 3 volume work
gave background coverage, discussed ter-
minology, presented a key to the Nearctic
families, and included keys to the genera of
24 families of Nematocera and 19 families
of lower Brachycera. The second volume is
largely devoted to the families of Musco-
morpha (the so-called higher flies), but also
includes corrections and addenda for vol-
ume | and an index for volume 2. Five
families of Aschiza and 60 families of Schi-
zomorpha are recognized, with the taxo-
nomic arrangement of families following the
classification scheme presented in volume
1. The coverage for each family includes a
synopsis, adult description, larval descrip-
tion (not included for all families), biology
and behavior, classification and distribu-
tion, key to genera, and references. For a
few families descriptions of the egg and pu-
parium are also given. A total of 23 different
experts authored the family chapters, thus
assuring overall quality. There are 1912 ex-
cellent illustrations arranged in 287 plates,
with all but 62 of the drawings contributed
by Ralph Idema, the same individual who
illustrated volume 1. Undoubtedly, one of
the most important attributes of volumes
of the Manual is the uniformly high quality
of the figures, with nearly all of the salient
key references being illustrated. I encoun-
tered little difficulty with the generic keys
for three acalyptrate families poorly known
to me (Agromyzidae, Milichidae, Sphaer-
oceridae) and three families with which I
have had considerable experience (Otitidae,
Sciomyzidae, Ephydridae) and felt confi-
dent that I had correctly identifed the genus
involved. I did not test the keys to genera
of the calyptrate families. The number of
Nearctic species, a Summary statement on
geographic distribution, and reference to
important revisions are given for each ge-
nus. Sadly, keys to larvae were included for
only 8 of the 65 families.
Although quality varied somewhat from
family to family, my overall impression 1s
that the authors have covered the subjects
assigned to them ina very satisfactory man-
ner. The habitus sketches that head each
family chapter were very well executed. The
synopses and descriptions of adults were
quite well done, but the coverage of natural
history and morphology of the immature
stages was a bit weak in places. This latter
comment is not meant as a criticism of the
authors involved, but is a reflection of our
poor knowledge of the immature stages of
several families, particularly those of the
Acalyptratae. The larval stages were illus-
trated in volume | in connection with the
family key. Thus, there are relatively few
illustrations of immature stages in this vol-
ume. The natural history discussions in-
cluded summaries of larval feeding habits,
comments on habitat distribution, and ref-
erences to important papers giving infor-
mation on biology and morphology of the
immature stages. Sections dealing with clas-
sification and distribution included infor-
mation on the phylogenetic placement of
126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
the family, its geographic distribution,
number of species in the world and Nearctic
Region, and comments on fossil species.
There were surprisingly few typographical
errors in the text, and the figure labels were
well done, although a few broken letters were
noted. A minor criticism is that some of the
figures are so light that patterns are poorly
differentiated. Another complaint is that the
binding seems surprisingly poor—my copy
of volume | is already showing deteriora-
tion.
It is emotionally exciting and profession-
ally satisfying that dipterists finally have a
modern, well-illustrated key to the genera
of Nearctic families, as interest in the order
undoubtedly will be stimulated now that
some of the taxonomic headaches have been
overcome. The recent publication of cata-
logs dealing with various faunal regions, the
continual appearance of family and generic
revisions, and the publication of this man-
ual all indicate that the world of Diptera is
alive and well. Now, attention should be
turned to an updating of Hennig’s volumes
on the larval stages.
B. A. Foote, Department of Biological
Sciences, Kent State University, Kent, Ohio
44242.
PROC. ENTOMOL. SOC. WASH.
90(1), 1988, pp. 127-128
Book REVIEW
Sphingidae Mundi (Hawkmoths of the
World). By Bernard D’Abrera. E. W.
Classey Ltd., Faringdon, United King-
dom [1987] 1986, 226 pages, 80 colored
plates, large format. Price: £97.50 (ap-
proximately $145.00).
More than 1000 sphingid species are 1l-
lustrated by superb colored photographs and
extremely good color separations. Compar-
ison of the plates with several specimens
shows the colors to be quite accurate. The
greatest contrast in quality is on page 117
where Sataspes tagalica is a very good rep-
resentation of metallic-reflecting scales;
however, the numerous species of Cepho-
nodes are incorrect. Most species of Cepho-
nodes have a pale yellowish-green cast to
the wing membrane; these appear bluish gray
with yellowish-gray overtones in the pho-
tographs. This result may reflect the method
used to back light the specimens when the
photographs were made. The background
has been replaced with a uniform color on
all plates. A four-plate appendix has illus-
trations of 40 species not represented in the
British Museum (N.H.).
A more appropriate title might be Sphin-
gidae of the World as Represented by the
Collection of the British Museum (N.H.) and
as Arranged by Alan Hayes. Additional, but
not inordinate, time would have been re-
quired to obtain specimens of the several
species of which the British Museum does
not have examples but which Alan Hayes
included in his draft checklist provided
D’Abrera to form the basis of the arrange-
ment of the illustrations. The very brief in-
troduction includes description of the ab-
breviations used throughout the text and on
the plates, a labeled example of fore- and
hindwing venation, a labeled diagram of a
pretarsal segment, and a generalized lateral
view of the head. The labels for forewing
venation (p. 7) are incorrect. Starting with
the posterior margin, the basally forked vein
is 1[A+1B, the first vein arising from the
cell is vein 2, then all the numbers should
be increased by one until the last numbered
vein is 12. A systematic list of valid genera
(with the exception of Neococytius, which
is not treated by name elsewhere in the text)
precedes the full work. A general treatment
for a genus includes the author and date of
publication of the type species; a discussion
of distribution, number of species, charac-
ters purporting to discriminate the taxon,
larvae, and pupae; and host list. Species are
briefly treated with the scientific name, au-
thor, date, and World List abbreviation of
original description, geographic distribu-
tion, comments about the species including
differentiating features and larval hosts.
Throughout, synonyms of species and gen-
era rarely are given. When subspecific names
are used, the original publications and dates
are not cited.
This book cries for critical editing. Start-
ing on page 12, the family Sphingidae is
attributed to Samouelle, 1819; the subfam-
ily Sphinginae to Latreille, ?1805. Accord-
ing to the International Code of Zoological
Nomenclature the author of a family-group
name is the person who first proposed the
name in a higher category sense. Thus,
Sphingidae should be attributed to Latreille.
In the discussion of Agrius (p. 12) the au-
thors of the rearing records are Szent-Ivany
and Carver, but in the references section the
authors are cited as Carver and Szent Ivany
(without hyphen). Under Coelonia fulvi-
notata (p. 14) D’Abrera dismisses Carcas-
son’s synonymy of fu/vinotata under C.
solani without explanation. Curious incon-
sistencies are his treating Weganoton scri-
bae as a valid species, yet in the discussion
he says “I consider this to be no more than
a local race of analis Felder.”” On page 12
proboscis is misspelled as probscis. The ci-
tation under Megacorma obliqua (p. 12)
128
should be List Spec. Lep. Ins. BM—not MM.
On page 24 for Manduca kuschei the place
name is Venadio, not Venodio. On page 40
Sphinx lanceolata (incorrectly attributed to
Boisduval, 1870, instead of R. Felder, 1868)
is treated as a valid name with Sphinx leu-
cophaeta (correctly leucophaeata) Clemens
a junior synonym. However, /eucophaeata
was described in 1859 and 1s the valid name
of the species. Also on page 40 the author
of vancouverensis is Edwards, not Edward.
On plate [page 123] E. damasi should be E.
adamsi. On pages 156, 157 the text correctly
gives floridensis as the valid name, whereas
the plate uses nessus.
This illustrated, synoptic checklist of
Sphingidae should be accompanied by a text
with the names, authors, and dates of all
taxa. Readers must be warned that many of
the species are best determined by genital
characters and that the higher classification
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
uses them extensively. Many species are
highly variable; attempts to identify many
of them without knowledge of their origin
and genital characters may result in incor-
rect identification. D’Abrera treats all spe-
cific epithets as nouns in apposition, a prac-
tice that will cause some readers concern;
and, parentheses are never used for authors’
names.
Despite the numerous shortcomings, I
know that I will use this work extensively
to make preliminary identifications and as
a guide to the British Museum (N.H.) col-
lection—the premier collection of world
Sphingidae.
Ronald W. Hodges, Systematic Ento-
mology Laboratory, Agricultural Research
Service, USDA, % NHB 168, National Mu-
seum of Natural History, Washington, DC
20560.
INSTRUCTION TO AUTHORS FOR
PREPARATION OF MANUSCRIPTS
GENERAL POLICY
Publication in the Proceedings is gener-
ally reserved for members. Manuscripts
should be in English and not be so lengthy
that they would exceed 15 printed pages in-
cluding illustrations (two typewritten pages
are approximately equivalent to a printed
page.) Manuscripts are peer-reviewed be-
fore they are accepted. Acceptance of manu-
scripts is the responsibility of the Editor.
Papers are published in the order they are
received rather than in order of date of ac-
ceptance. This eliminates possible bias due
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CONTENTS
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TURNER, W. J.—Lectotype designation for Empis chichimeca Wheeler and Melander (Diptera:
Empididae)
WHEELER. A. G., Jr., and E. R. HOEBEKE—Apterona helix (Lepidoptera: Psychidae), a
Palearctic bagworm moth in North America: New distribution records, seasonal history,
and host plants
ZACK, R. S. and R. W. SITES—A new species of Donaceus Cresson (Diptera: Ephydridae)
from Malaysia
NOTES
McCAFFERTY, W. P.—Neotype designation for Raptoheptagenia cruentata (Walsh) (Ephem-
eroptera: Heptageniidae)
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BOOK REVIEWS
ARNAUD, P. H., Jr.— The Bombyliidae of Deep Canyon
FOOTE, B. A.— Manual of Nearctic Diptera, Volume 2
HODGES, R. W.—Sphingidae Mundi (Hawkmoths of the World)
SMILEY, R. L.— Historical Perspective and Current World Status of the Tomato Russet Mite
(Acari: Eriophyidae)
INSTRUCTIONS TO AUTHORS
| OL. 90 APRIL 1988 NO. 2
Li (ISSN 0013-8797)
PROCEEDINGS.
of the
ENTOMOLOGICAL SOCIETY
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CONTENTS
BOLDT, P. E., W. WOODS, and T. O. ROBBINS—Phytophagous insect fauna of Baccharis
sarothroides Gray (Asteraceae) in Arizona and New Mexico .....................---.-. 207
CHEMSAK, J. A. and C. FELLER—New species of Cerambycidae from Twin Cays, Belize
(Woleoplera)y sere baie Meee PU TEs cLeRE ER BO PPh ee Re AER eee eet 179
DAVIS, D. R. and E. G. MILSTREY — Description and biology of Acrolophus pholeter, (Lepi-
doptera: Tineidae), a new moth commensal from gopher tortoise burrows in Florida .... 164
HANSON, P. E. and J. C. MILLER—Notes on the biology of Caenocephus aldrichi Bradley
fiiyimenapteras Cephidae)) Mik tO PP AE TON ie ha BS ee A 204
HARMON, J. D. and M. H. ROSS—Effects of malathion and diazinon exposure on female
German cockroaches (Dictyoptera: Blattellidae) and their oothecae .................... 248
HUANG, Y. M.— Aedes (Stegomyia) josiahae, a new species of the simipsoni subgroup (Diptera:
Gath ord PN NM PSA SE a ARR ee TN Ey Oe PE OL 155
JOHNSON, N. F.—Species of Australian Telenominae (Hymenoptera: Scelionidae) of A. P.
DWoddiandvAWAN Giraulty PEASE OPE ee PE WR eee et CE Fakir ne
JOHNSON, N. F. and F. BIN—Telenomus species (Hymenoptera: Scelionidae) associated with
thejegesiof Zyeaemidae (Lepidoptera) Wie eS. VRP hep ele PB ee ee VE le 244
MARSHALL, S. A. and D. J. S. MONTAGNES-— Ceroptera longicauda, a second North Amer-
ican species in the kleptoparasitic genus Ceropfera Macquart (Diptera: Sphaeroceridae) .. 189
NEAL, J. W., JR. and K. M. GOTT—Evidence for multivoltinism in Prodiplosis platani Gagné
(Diptera: Cecidomyiidae), a leaf curl midge of American sycamore .................... 201
PALMER, W. A. and F. D. BENNETT—The phytophagous insect fauna associated with Bac-
charis halimifolia L. in the eastern United States ...........0.0..0.00 0250000 c cee eee ee 216
QUICKE, D. L. J.—Digonogastra: The correct name for Nearctic Jphiaulax of authors (Hy-
menopleraweracooidae)) AMER ES Ppt ro Po ahi eka Ee A 196
ROBBINS, R. K.—Comparative morphology of the butterfly foreleg coxa and trochanter (Lep-
idoptera) and its systematics implications ............0 20)... e eee ee ele ee eee Mh tes}
STAINES, C. L., JR.—A review of the species of Acritispa Uhmann (Coleoptera: Chrysomelidae,
LEWIS) 0) UTE) MC EET TMT TRE DA 9 09 DP Tr a 193
(Continued on back cover)
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ENTOMOLOGICAL SOCIETY
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PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 133-154
COMPARATIVE MORPHOLOGY OF THE BUTTERFLY FORELEG
COXA AND TROCHANTER (LEPIDOPTERA) AND
ITS SYSTEMATIC IMPLICATIONS
ROBERT K. ROBBINS
Department of Entomology, NHB STOP 127, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.
Abstract.—1 describe and illustrate five qualitatively distinct forms of butterfly foreleg
coxa, trochanter, and basal femur, and give the distribution of each type by sex for the
butterfly families. I code this variation in a character matrix of four characters with 9
character states, from which I derive a most parsimonious cladogram with four mono-
phyletic groups: (1) Styginae (Riodinidae), (2) Hamearinae (in part) + Styginae + Riodini-
nae + Euselasiinae (Riodinidae), (3) Lipteninae + Poritinae + Liphyrinae + Miletinae +
Curetinae (Lycaenidae), and (4) Riodinidae + Libytheidae + Nymphalidae. The second
and third groups have not been recognized previously as monophyletic. The fourth sup-
ports previous results based on other leg characters, but is inconsistent with most published
phylogenies to the butterfly families. Contrary to previous reports, the forecoxa of male
Styx infernalis (Riodinidae: Styginae) extends beyond the trochanter, but this extension
is smaller than in most other riodinids. I also show that the male forelegs of Curetis
(Lycaenidae: Curetinae) and Riodinidae are qualitatively different, a result that does not
support the hypothesis that these two taxa are sister groups.
Key Words:
For more than 125 years, classification of
the butterfly families has relied heavily on
foreleg characters, particularly those of the
tarsus (Bates 1861, Ford 1945), but mor-
phology of the male foreleg coxa and tro-
chanter has also been used in butterfly higher
classification (cf. Borror et al. 1981 for an
introduction to insect leg morphology).
Godman and Salvin (1879-1901) discov-
ered that the male forecoxa of riodinids ex-
tends beyond its articulation with the tro-
chanter, and Stichel (1910-1911) and
Ehrlich (1958b) characterized the Riodini-
dae (Ehrlich’s Riodininae), in part, by this
structure. Ehrlich also erected a new mono-
basic “subfamily” — of rank equal to the Ly-
caenidae (his Lycaeninae) and Riodinidae—
for Styx infernalis Staudinger because its
leg characters, butterfly, cladogram
male forecoxa does not extend spinelike be-
yond the trochanter and because it differs
from riodinids in a few other structures.
Scott (1985) proposed that Curetis (a genus
that Ehrlich had considered to be a lycaenid)
and Riodinidae (his Riodininae without
Styx) are sister groups because both have
the male foreleg coxa extending beyond the
trochanter.
The few published figures of foreleg coxae
and trochanters lack detail (e.g. Ehrlich
1958a, b, Scott 1986), and I propose to solve
this problem with the use of a scanning elec-
tron microscope (SEM). It is clearly impor-
tant that the morphology of these structures
be well documented if they are to be used
in constructing familial classifications of the
butterflies. The first purpose of this paper
134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
is to describe and illustrate the foreleg coxa,
trochanter, and basal femur of males and
females from the different butterfly families.
The second purpose of this paper is to
assess the morphologic and phylogenetic
hypotheses of Ehrlich (1958b) and Scott
(1985). Specifically, I (1) check Ehrlich’s
statement that the male foreleg coxa of Styx
does not extend beyond its articulation with
the trochanter and (2) assess Scott’s pro-
posal that Curetis and the Riodinidae are
sister groups, based in part on the obser-
vation that in both taxa the male forecoxa
extends beyond the trochanter.
The third purpose of this paper is to use
variation of the foreleg coxa and trochanter
among higher taxa to further our under-
standing of butterfly phylogeny. I code this
variation in a character matrix, derive a most
parsimonious cladogram, determine wheth-
er it 1S consistent with published phyloge-
nies (Ehrlich 1958b, Kristensen 1976, Scott
1985), and assess the monophyly of some
higher taxa.
MATERIALS AND METHODS
Because foreleg coxae are difficult to re-
move from dried specimens without break-
age, In most cases I wetted the whole body
(after removing the wings) in 80% ethanol,
soaked it in 10% potassium hydroxide at
room temperature for 24-48 hours, and
transferred it to 80% ethanol. I then re-
moved both forelegs, and brushed and
scraped off as many scales as possible with
forceps and a brush with stout bristles. In
some cases where scales were particularly
hard to remove, I transferred the legs to
acetone, which helped to loosen the scales.
At this point I examined specimens with a
binocular stereomicroscope, which is often-
times sufficient to determine structures.
For examination with an SEM, I soaked
foreleg coxa, trochanter, and femur prepa-
rations in absolute ethanol for 5-10 min-
utes, and mounted them on stubs in various
aspects. I mounted some laterally so that
they presented either an outside or inside
lateral aspect, others as an upright triangle,
which provided a posterior aspect in addi-
tion to both lateral aspects, and still others
as parts of segments to show particular
structures. I glued the specimens at the or-
igin of the coxa and/or at the distal end of
the femur, and the stubs were coated with
carbon and gold.
RESULTS
There are five qualitatively distinct forms
of foreleg coxa, trochanter, and femur; all
five occur in males while two are found in
females. The foreleg coxa, trochanter, and
femur in butterflies have a complex three-
dimensional morphology that is difficult to
communicate on a two-dimensional printed
page. I describe the first leg type in detail
using pictures from inside lateral, posterior,
outside lateral, and anterior aspects, and note
some of the major morphological “‘land-
marks” and shapes. I then describe the other
leg types by focusing on how they differ from
the first one. I illustrate specimens repre-
senting diverse taxonomic groups to show
some of the quantitative variation within
each foreleg type. Under this description, I
list genera by family in which I found it.
Because distribution of the different foreleg
types differs in the sexes, I list distributions
in males and females separately. If I ex-
amined more than one specimen of one sex
in a genus, then I place an asterisk (*) after
the generic name.
The familial classification follows Ehrlich
(1958b) except for the Lycaenidae and
Riodinidae, for which I follow Eliot (1973)
and Harvey (1987), respectively. Harvey di-
vided the Riodinidae into the subfamilies
Styginae, Corrachiinae, Hamearinae, Eu-
selasiinae, and Riodininae. The Corrachiin-
ae contains a single rare species that I have
not had an opportunity to examine.
Type I
Morphology.—Foreleg coxa: A tapering
tubular structure that is shaped very differ-
ently than the midleg or hindleg coxa. Ehr-
lich (1958a) reported that the coxa 1s grooved
VOLUME 90, NUMBER 2
laterally in the monarch (Danaus plexippus
Linnaeus), an observation that I believe to
be incorrect. I list each morphological struc-
ture by letter, and use that letter to designate
it in the figures.
The foreleg coxa has a pair of posterior
pointing mid- to ventro-lateral processes
that articulate with the trochanter. The hinge
formed between these processes and the tro-
chanter allows leg movement along the lon-
gitudinal plane. (A) One process is on the
inner lateral side (Figs. 1-4) and (B) the oth-
er on the outer lateral side (Figs. 5-8).
(C) There are two rod-like “tendons”
within the coxa that attach distally to the
trochanter, one dorsally, the other ventrally
(not illustrated). When the coxa and tro-
chanter are separated, the tendons usually
remain attached to the trochanter. They are
best seen with transmitted light under a bin-
ocular stereomicroscope.
Foreleg trochanter: A complexly curved
three-dimensional segment.
(D) There are a pair of prongs on the dor-
sal basal edge of the trochanter (Figs. 9-12).
They attach to the dorsal “tendon” of the
coxa. The prongs vary considerably in ex-
tent, and are reduced to two bumps in some
Nymphalidae (Fig. 12).
(E) The outside surface of the trochanter
is rounded in posterior aspect (Figs. 9-12),
and is indented anteriorly in lateral aspect
where the posterior coxa process articulates
with it (Figs. 5-8).
(F) The inner surface of the trochanter is
slightly concave in posterior aspect (Figs. 9-—
12), and is slightly indented ventrally where
it articulates with the femur process (Figs.
1-4).
(G) There is a slit/groove that extends
dorsally from the posterior edge of the in-
dentation for the femur process and that
forms the posterior edge of the concave area
on the inner surface of the trochanter (Figs.
1-3, 9-10). I presume that this slit/groove
allows the leg some lateral flexibility in
movement.
There are three clusters of small trichoid
sensilla (5 or more sensilla, less than 40 mi-
135
crons in length except in some larger but-
terflies, such as Papilionidae) on the tro-
chanter. (H) A cluster on the lateral
indentation just anterior and ventral to the
inside dorsal prong of the trochanter (Figs.
1-4, 9-12). (I) A cluster on the lateral in-
dentation just anterior and ventral to the
outside dorsal prong of the trochanter (Figs.
5-8, 9-12). (J) A third cluster just below the
articulation of the coxa process on the an-
terior face of the trochanter. It can be seen
from an inside lateral aspect (Figs. 1-4), but
is best seen in anterior aspect (Figs. 13-14).
I presume that these trichoid sensilla are
mechanoreceptors, at least in part, because
they occur where movements of the tro-
chanter would cause them to come into con-
tact with the coxa. There are also other
trichoid sensilla scattered over the foreleg,
but they occur singly or in a cluster of two,
and are often longer than 40 microns in
length.
Foreleg femur: A simple tubular structure
at its basal end, where it connects to the
trochanter.
(K) There is a basal process on the pos-
terior inner face of the femur (Figs. 1-4, 9-
11). This process may be rounded or some-
what tapered to a point.
Male distribution. — Hesperiidae: Poanes
Scudder, Megathymus Scudder, Autochton
Hiibner, Epargyreus Hubner.
Papilionidae: Papilio Linnaeus, Battus
Scopoli, Eurytides Hiibner, Parnassius La-
treille.
Pieridae: Eurema* Hiibner, Phoebis
Hiibner, Colotis Hiibner, Pieris* Schrank,
Euchloe Hiibner, Dismorphia Hiibner.
Lycaenidae (Theclinae, Polyommatinae,
Lycaeninae): Arawacus Kaye, Strymon
Hiibner, Calycopis Scudder, Evenus Hub-
ner, Allosmaitia Clench, Hypaurotis Scud-
der, Axiocerses Hiibner, Everes Hiibner, Ce-
lastrina* Tutt, Lycaena* Fabricius.
Female distribution. — Hesperiidae: Hes-
peria Fabricius, Poanes, Thorybes Scudder,
Erynnis Schrank.
Papilionidae: Papilio, Battus, Eurytides,
Parnasstus.
136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4. Foreleg coxa, trochanter, and basal femur in lateral inside aspect. Coxa horizontal on top. Letters
refer to structures in text. 1, Eurema male (Pieridae) (scale line 176 microns). 2, Celastrina female (Lycaenidae)
(scale line 176 microns). 3, Stvx female (Riodinidae) (scale line 176 microns). 4, Libythea female (Libytheidae)
(scale line 150 microns).
VOLUME 90, NUMBER 2 137
Figs. 5-8. Foreleg coxa, trochanter, and basal femur in lateral outside aspect. Coxa horizontal on top. Letters
refer to structures in text. 5, Hesperia female (Hesperiidae) (scale line 300 microns). 6, Arawacus male (Ly-
caenidae) (scale line 136 microns). 7, Stalachtis female (Riodinidae) (scale line 200 microns). 8, Prepona female
(Nymphalidae) (scale line 380 microns).
138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 9-12. Foreleg trochanter in posterior aspect. Outside of leg to left except in Danaus. Letters refer to
structures in text. 9, Eurema male (scale line 100 microns). 10, Styx female (scale line 150 microns). 11, Libythea
female (scale line 150 microns). 12, Danaus female (Nymphalidae) (scale line 150 microns).
VOLUME 90, NUMBER 2 139
Figs. 13-16. Foreleg trochanter in anterior and posterior aspects. Letter refers to structure in text. 13, Strymon
male (Lycaenidae) (scale line 136 microns), anterior aspect, outside of leg to right, coxa on top. 14, Marpesia
female (Nymphalidae) (scale line 67 microns), anterior aspect, outside of leg to left, coxa on top. 15, Poritia
male (Lycaenidae) (scale line 200 microns), posterior aspect, outside to right. 16, Curetis female (Lycaenidae)
(scale line 150 microns), posterior aspect, outside to right.
140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Pieridae: Eurema, Phoebis, Archonias
Hiibner, Pieris.
Lycaenidae (Theclinae, Polyommatinae,
Lycaeninae): Eumaeus Hibner, Calycopis,
Evenus, Axiocerses, Everes, Celastrina,* Ly-
caena.*
Riodinidae (Styginae, Hamearinae, Eu-
selasiinae, Riodininae): Styx Staudinger,
Laxita Butler, Hamearis Hiibner, Hades
Westwood, Stalachtis Hiibner, Ancyluris
Hiibner, Mesosemia Hiibner, Eurybia Ih-
ger.
Libytheidae: Libythea* Fabricius.
Nymphalidae: Dynamine* Hiibner, Pre-
pona* Boisduval, Doxocopa Hiibner, Dan-
aus Kluk, Marpesia* Hiibner, Chlosyne
Butler.
Type II
Morphology. — Foreleg coxa: This foreleg
type retains structures A-K (Figs. 15-24),
and its trochanter and femur do not differ
from the Type I foreleg. It differs only in
the shape of the coxa.
(L) The distal end of the coxa is arched
dorsally, but there is a lot of quantitative
variation within this character state. In some
genera (Allotinus Felder & Felder, Liphyra
Westwood, Pentila Westwood, Ornipholi-
dotos Bethune-Baker, Falcuna Stempffer &
Bennett), the dorsal coxa forms a “hump”
(Figs. 17, 21). In others (Feniseca Grote,
Poritia Moore), the hump points dorso-pos-
teriorly in a process that extends beyond (by
approximately 0.1 mm) the articulation with
the trochanter (Fig. 18). And in Curetis, the
process extends well beyond (by approxi-
mately 0.3 mm) the trochanter (Figs. 19, 20,
22). It may be possible to code this variation
in character states, but it would entail a more
detailed study of the genera that have the
Type II foreleg.
Male distribution. — Lycaenidae (Lipten-
inae, Poritiinae, Liphyrinae, Miletinae,
Curetinae): Pentila, Falcuna, Poritia, Allo-
tinus, Feniseca,* Curetis.*
Female distribution. — Lycaenidae (Lip-
teninae, Poritiinae, Liphyrinae, Miletinae,
Curetinae): Ornipholidotos, Falcuna, Pori-
tia, Liphyra, Allotinus, Feniseca,* Curetis.
Type III
Morphology.— Foreleg coxa and trochan-
ter: This foreleg type retains structures A-I
and K (Figs. 25-39), and its femur does not
differ from the Type II foreleg. It differs in
the structure of the coxa and trochanter, and
in that it is restricted to male forelegs.
(M) The dorsal, distal end of the foreleg
coxa extends beyond the lateral processes
of the coxa (structures A and B) and beyond
the articulation of the trochanter in a pro-
cess that is not arched dorsally (Figs. 25-
26, 29-30, 33-34, 37-38).
(N) The cluster of trichoid sensilla on the
inner anterior face of Type I and II foreleg
trochanters (structure J) is lacking (Figs. 25,
28-29, 32-33, 36-37). This group of sensilla
is lacking in all male butterflies that do not
use their forelegs for walking, including the
next two types. It is retained, however, in
female nymphalids (Fig. 14), which do not
use their forelegs for walking.
The Type III foreleg coxa shows two kinds
of quantitative variation. First, the dorsal
process of the coxa varies in length and
shape. At one extreme, the dorsal process
in Laxita and Libythea extends beyond the
trochanter in a blunt process (approximate-
ly 0.10-0.15 mm) (Figs. 33-34, 37-38). At
the other extreme in genera such as Anartia
(Fig. 29), the dorsal process is rounded and
barely extends beyond the trochanter
(< 0.05 mm). Second, in some genera, such
as Doxocopa, Prepona, and Marpesia, the
coxa has a flap on the distal outside lateral
side that “covers” the ventro-lateral process
(Figs. 26-27). This flap is less well devel-
oped in Dynamine and Memphis, poorly de-
veloped in Danaus, and is apparently lack-
ing in Anartia, Libythea, and Laxita. The
trochanter is somewhat twisted in species
with this flap so that the cluster of sensilla
on the outside of the trochanter is more
VOLUME 90, NUMBER 2 141
Figs. 17-20. Foreleg coxa and trochanter in inside lateral aspect. Coxa horizontal on top. Letter refers to
structure in text. All Lycaenidae. 17, A//otinus male (scale line 200 microns). 18, Feniseca female (scale line 176
microns). 19, Curetis male (scale line 380 microns). 20, Curetis female (scale line 380 microns).
142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 21-24. Foreleg coxa and trochanter in outside lateral aspect and trochanter in anterior aspect. All
Lycaenidae. 21, A//otinus male, coxa horizontal on top, (scale line 200 microns). 22, Curetis male, coxa horizontal
on top, (scale line 380 microns). 23, Feniseca male, outside of leg to right (scale line 136 microns). 24, Feniseca
female, outside of leg to left (scale line 136 microns).
VOLUME 90, NUMBER 2 143
Figs. 25-28. Foreleg coxae and trochanters. Letter refers to structure in text. All Nymphalidae. 25, Marpesia
male, inside lateral aspect with coxa horizontal on top (scale line 150 microns). 26, Marpesia male, outside
lateral aspect with coxa horizontal on top (scale line 136 microns). 27, Marpesia male, posterior aspect with
outside to left (scale line 150 microns). 28. Memphis male, anterior aspect with coxa on top and outside to left
(scale line 136 microns).
144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 29-32. Nymphalid foreleg coxae and trochanters. 29, Anartia male, inside lateral aspect with coxa
horizontal on top (scale line 150 microns). 30, Dynamine male, outside lateral aspect with coxa horizontal on
top (scale line 150 microns). 31, Dynamine male, posterior aspect with outside to right (scale line 86 microns).
32, Heliconius male, anterior aspect with coxa on top and outside to right (scale line 136 microns).
VOLUME 90, NUMBER 2 145
ia
Figs. 33-36. Foreleg coxa and trochanter of male Libythea. 33, Inside aspect of coxa and trochanter with
coxa horizontal on top (scale line 176 microns). 34, Outside aspect of coxa and trochanter with coxa horizontal
on top (scale line 176 microns). 35, Posterior aspect of trochanter (scale line 50 microns). 36, Anterior aspect
of trochanter (scale line 60 microns).
146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 37-40. Foreleg coxa and trochanter and trochanter in posterior aspect (Riodinidae). 37, Laxita male,
inside aspect with coxa horizontal on top (scale line 120 microns). 38, Laxita male, outside aspect with coxa
horizontal on top (scale line 120 microns). 39, Laxita male, posterior aspect with outside to left (scale line 43
microns). 40, Hamearis male, inside lateral aspect (scale line 120 microns).
VOLUME 90, NUMBER 2
ventral than the cluster on the inside (Figs.
27, 31). A more extensive survey of Type
III forecoxae might reveal phylogenetically
useful qualitative variation within the
Nymphalidae or between the Nymphalidae
and Libytheidae + Riodinidae (Hameari-
nae-Laxita).
The second source of quantitative vari-
ation is the development of the dorsal prongs
on the posterior trochanter (structure D).
They are reduced to bumps in most genera,
and in Libythea (Fig. 35), there 1s a third
small bump between the two reduced prongs.
Male distribution.—Riodinidae (Ha-
mearinae in part): Laxita.
Libytheidae: Libythea.*
Nymphalidae: Hypanartia Hibner, An-
artia Hiibner, Heliconius* Kluk, Pagyris*
Boisduval, Danaus, Marpesia, Dynamine*,
Callicore Hiibner, Taygetis Hiibner, Dox-
ocopa, Prepona, Anaea Hiibner, Memphis*
Hiibner.
Type IV
Morphology.—Foreleg trochanter: The
coxa and femur do not differ qualitatively
from the Type III foreleg (Figs. 40-48), but
the trochanter does.
(O) The cluster of trichoid sensilla on the
inside dorso-lateral posterior trochanter
(structure H) is absent (Figs. 40-43, 47) while
the Type III leg retains this cluster. Thus,
the Type IV male foreleg trochanter is miss-
ing both clusters of trichoid sensilla on the
inside, but retains the cluster on the outside
(Figs. 40, 44, 46, 47).
All the Type IV forelegs that I examined
under the SEM lacked the cluster of trichoid
sensilla except for male Ancyluris and Ha-
mearis, which had one sensillum (Figs. 40,
43). In the Ancyluris specimen, however, the
other leg had no sensilla. I do not know if
the presence of a single sensillum is a ves-
tigial condition or if the sensillum 1s differ-
ent from those in previous leg types clus-
tered on that part of the trochanter. In either
case, there is no cluster of trichoid sensilla.
147
The extension of the coxa beyond its ar-
ticulation with the trochanter is highly vari-
able in the Type IV foreleg. The amount
that the coxa extends beyond the trochanter
varies in the species that I examined from
0.23 mm in Hamearis and 0.28 mm in Sta-
lachtis to about 0.80 mm in Thisbe Hibner.
In Thisbe, the distal part of the coxa is long-
er than the basal part, but the opposite is
true in Stalachtis and Hamearis. The male
foreleg coxa of Curetis (Type II) extends be-
yond the trochanter more (approximately
0.30 mm) than in Stalachtis and Hamearis,
but it is arched upwards whereas it 1s bluntly
tapered in the riodinids.
The trochanter of the Type IV foreleg is
sometimes shaped like a cylinder (Figs. 41-
42), with the dorsal prongs completely re-
duced. In some genera, however, the tro-
chanter is shaped much like that in Type HI
forelegs.
Male distribution. — Riodinidae (Eusela-
siinae, Riodininae, Hamearinae in part):
Hades, Emesis* Fabricius, Thisbe, Stalach-
tis, Ancyluris, Mesosemia, Hamearis.
Type V
Morphology.—Foreleg trochanter: The
foreleg trochanter again differs in the ab-
sence of a cluster of trichoid sensilla. Oth-
erwise, the Type V foreleg retains the char-
acters of the Type IV foreleg (Figs. 49-52).
(P) The trochanter lacks the cluster of
trichoid sensilla on the outside dorso-lateral
posterior surface (structure I) (Fig. 50).
I have examined with the SEM two male
forelegs from one male specimen of Styx.
Both forelegs have one trichoid sensillum
on the trochanter in the general area where
the outside posterior cluster of the trochan-
ter occurs in other butterflies. It is unclear
whether this single sensillum is a remnant
of the cluster or a different kind of sensil-
lum. In either case, the lack of a cluster is
unique among the butterflies.
The inside dorsal cluster of the trochan-
ter, which is absent or reduced to one sen-
148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 41-44. Lateral aspect of foreleg coxae and trochanters (Riodinidae). 41, Stalachtis male, inside aspect
with coxa horizontal on top (scale line 200 microns). 42, Emesis male, inside aspect with coxa horizontal on
top (scale line 120 microns). 43, Ancy/uris male, inside aspect with coxa on top (scale line 176 microns). 44,
{ncyluris male, outside aspect with coxa on top (scale line 176 microns).
VOLUME 90, NUMBER 2
ff i \
—_ Vip? 5 SN
j ‘ i Py
i A PAN
“Se - IGPER
Figs. 45-48. Foreleg coxae and trochanters (Riodinidae). Letters refer to structures in text. 45, Emesis male,
outside lateral aspect with coxa horizontal on top (scale line 300 microns). 46, Sta/achtis male, posterior aspect
of dorsal trocanter showing cluster of trichoid sensilla on outside surface (scale line 30 microns). 47, Hades
male, posterior aspect of trochanter, outside to left (scale line 100 microns). 48, Ancy/uris male, anterior aspect
with coxa on top, outside to right (scale line 136 microns).
Hi,
150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lateral aspect with coxa horizontal on top (scale line 120 microns). 50, Outside lateral aspect with coxa horizontal
on top (scale line 120 microns). 51, Posterior aspect of trochanter, outside to left (scale line 120 microns). 52,
Enlargement of two setae on dorsal outside face of trochanter in Fig. 49 (scale line 15 microns).
VOLUME 90, NUMBER 2
sillum in the Type IV foreleg, is similarly
reduced in Styx. One foreleg has a single
trichoid sensillum while the other has two
(Figs. 49, 52), but the sensilla are aberrant.
On both legs, the sensillum socket is con-
siderably larger than the “‘stalk”’ of the sen-
sillum (Fig. 52). The space between the
socket walls appears to be solid, and it is
unclear whether the stalk goes through the
integument. It appears to be a different kind
of trichoid sensillum than those with foreleg
Types I, II, and III, or it is possible that it
is a vestigial structure.
The coxa has a dorsal posterior process
that barely extends beyond the articulation
of the trochanter (approximately 0.03 mm).
In this regard, it is more similar to the Type
III than the Type IV foreleg.
Male distribution. — Riodinidae (Stygin-
ae): Styx.*
CHARACTER MATRIX
I summarize the information above in a
character matrix (Table 1), and derive a most
parsimonious cladogram from it (Fig. 53).
I use the Hesperiidae as the outgroup for
the Papilionoidea (Kristensen 1976, Scott
1985), and put an asterisk (*) next to the
primitive character state for the papilio-
noids.
1. Foreleg coxa (A) in both sexes the dor-
sal surface is arched upwards at the distal
end, may extend beyond the articulation
with the trochanter, (B)* in both sexes the
coxa tapers distally, but is not arched up-
wards nor does it extend beyond its artic-
ulation with the trochanter, (C) females as
in the previous state, but in males the coxa
tapers distally in a blunt process that is not
arched upwards, but that extends beyond
the articulation with the trochanter.
2. In males, the trochanter (A)* has a clus-
ter (=5) of trichoid sensilla on the anterior,
inner lateral surface, (B) lacks this cluster of
trichoid sensilla.
3. In males, the trochanter (A)* has a clus-
ter of trichoid sensilla on the dorso-poste-
151
Table 1. Matrix of foreleg coxa and trochanter
characters. Foreleg types and character states are given
in the text. Lycaenidae #1: subfamilies Lycaeninae,
Theclinae, and Polyommatinae, sensu Eliot (1973). Ly-
caenidae #2: Lipteninae, Poritinae, Liphyrinae, Mile-
tinae, and Curetinae. Riodinidae #1: Hamearinae
(Laxita), sensu Harvey (1987). Riodinidae #2: Ha-
mearinae (Hamearis), Euselasiinae, and Riodininae.
Riodinidae #3: Styginae.
Character
Taxon 1 2 3 4
Type I male and female forelegs
Hesperiidae B A A A
Papilionidae B A A A
Pieridae B A A A
Lycaenidae #1 B A A A
Type II male and female forelegs
Lycaenidae #2 A A A A
Type I female and type III male forelegs
Libytheidae Gc B A A
Nymphalidae Cc B A A
Riodinidae #1 (6 B A A
Type I female and type IV male forelegs
Riodinidae #2 ( B B A
Type I female and type V male forelegs
Riodinidae #3 Cc B B B
rior inner lateral surface, (B) lacks this clus-
ter of trichoid sensilla.
4. In males, the trochanter (A)* has a clus-
ter of trichoid sensilla on the dorso-poste-
rior outer lateral surface, (B) lacks this clus-
ter of trichoid sensilla.
Of the three possible orders 1n which
the three states of character | could have
evolved, I chose transformation A-B-C. I
provisionally rejected transformation B-A-C
because it would require the female coxa to
evolve a dorsal arch and then lose it. I pro-
visionally rejected transformation B-C-A
because it would require the evolution of
sexual dimorphism in the coxa and then its
loss. Other characters that provide evidence
on the systematic placement of Lycaenidae
#2 may also provide a test of my transfor-
mation hypothesis.
152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
CLADOGRAM TO THE BUTTERFLIES
se
Fig. 53.
53.
Cladogram to the butterfly families based on distribution of character states of the foreleg coxa and
trochanter (Table 1). The numbers refer to characters and the letters to changes in character state.
DISCUSSION
Foreleg coxa and trochanter character
states are qualitatively invariant within pre-
viously recognized butterfly families except
for the Lycaenidae and Riodinidae, which
is significant in two respects. First, the Ly-
caenidae + Riodinidae are sometimes
lumped in a presumably homogeneous and
monophyletic taxon (e.g. Kristensen 1976),
perhaps because they are rich in species
whose morphology is poorly known. The
results in this paper and others (Robbins
1987, 1988) indicate that for leg characters,
at least, there is a great deal of morpholog-
ical variation among the Lycaenidae and
Riodinidae. Second, the lack of variation
within the Hesperiidae, Papilionidae, Pier-
idae, Libytheidae, and Nymphalidae in the
structure of foreleg coxae and trochanters
lends credence to their stability as evolu-
tionary characters (Kluge and Farris 1969).
The distribution of foreleg coxae and tro-
chanters— summarized in the character ma-
trix (Table 1)—provides evidence for four
presumably monophyletic taxa among the
butterflies (Fig. 53). The first taxon is the
riodinid subfamily Styginae, which has
uniquely evolved state B of character 4 (Fig.
53). I have not yet had the opportunity to
examine the legs of the monotypic riodinid
genera Petrocerus Callaghan and Corrachia,
but their forecoxae are apparently similar
to that of Styx (Callaghan 1979, Harvey
1987) in that they do not extend well beyond
the articulation with the trochanter.
The second monophyletic taxon is a com-
bination of the riodinid subfamilies Euse-
lasiinae, Riodininae, Styginae, and the ge-
nus Hamearis of the Hamearinae (Fig. 53).
It is characterized by the evolution of state
B of character 3. This result has not been
proposed previously, and suggests that the
New World Riodinidae plus Hamearis may
be a monophyletic group. The male foreleg
VOLUME 90, NUMBER 2
coxa and trochanter of the Old World rio-
dinid Laxita (Hamearinae) does not differ
qualitatively from those of Libytheidae or
Nymphalidae. Harvey (1987) considered the
Hamearinae to be monophyletic because
they share a posterior pointing beaked un-
cus in the male genitalia. My results conflict
with this classification, but clearly they are
preliminary since I have examined only two
genera in the Hamearinae.
The third monophyletic taxon is a com-
bination of the lycaenid subfamilies Lipten-
inae, Poritinae, Liphyrinae, Miletinae, and
Curetinae (Fig. 53). It is characterized by
the evolution of state A in males and fe-
males (character 1). Again, this combina-
tion of subfamilies has not been previously
recognized as monophyletic, and is incon-
sistent with Scott’s (1985) phylogeny of the
lycaenid subfamilies. If the transformation
of character | is B-A-C, however, then this
group could be paraphyletic. It consists of
hundreds of species restricted to the Old
World except for a single Nearctic species,
Feniseca tarquinius Fabricius.
The last monophyletic group is Riodini-
dae + Libytheidae + Nymphalidae, char-
acterized by the evolution of state C of char-
acter | and state B of character 2 (Fig. 53).
I have gotten the same phylogenetic result
using other leg characters (Robbins 1987).
Since all other published phylogenies (Ehr-
lich 1958, Kristensen 1976, Scott 1985)
consider the Lycaenidae + Riodinidae to
be a monophyletic group—in contradiction
to my results—either there has been a great
deal of convergence among leg characters or
the previous phylogenies have been based
on poorly analyzed characters whose dis-
tributions are also poorly known.
The results in this paper partly confirm
and partly contradict the morphological re-
sults of Ehrlich (1958b). They contradict
Ehrlich’s report that the male foreleg coxa
of Styx does not extend beyond the tro-
chanter, but its extension is smaller than in
most other riodinids, which 1s probably what
153
Ehrlich observed. Further, similar short ex-
tensions apparently occur in some other
riodinids, specifically Corrachia (Harvey
1987) and Petrocerus (Callaghan 1979),
Ehrlich’s finding that the foreleg coxa ex-
tends slightly below the articulation with the
trochanter in male Curetis (Lycaenidae) is
correct, but incomplete. He did not note
that the forecoxa also extends beyond the
articulation with the trochanter in male
Nymphalidae, Libytheidae, and Styx as well
as both sexes in some lycaenids with a Type
II forecoxa (Curetis, Feniseca, Poritia).
My results are inconsistent with Scott’s
(1985) phylogenetic hypothesis that Curetis
and Riodinidae (his Riodininae) form a
monophyletic group. He supported this hy-
pothesis in part by noting that the male fore-
coxa of these two groups extends beyond
the articulation with the trochanter. How-
ever, this “character state” occurs in many
other butterflies, as I have noted. Further,
the forecoxa of Curetis is qualitatively dis-
tinct from that in riodinids. It is arched dor-
sally, extends beyond the articulation with
the trochanter in both sexes, and its tro-
chanter retains a cluster of sensilla on its
inside anterior face. The forecoxa of riodin-
ids is not arched dorsally, extends beyond
the trochanter only in males, and its tro-
chanter does not retain the cluster of sensilla
on its inside anterior face. Thus, the simi-
larity in shape of the forecoxae of nodinids
and Curetis is superficial, and Scott’s hy-
pothesis would appear to be incorrect.
ACKNOWLEDGMENTS
I thank Don Harvey for numerous dis-
cussions and exchange of pertinent infor-
mation. I thank John Eliot, Don Harvey,
Gerardo Lamas, James Scott, and an anon-
ymous reviewer for critically reading and
commenting on the paper. I thank Brian
Kahn and Susann Braden for working with
me on the SEM. I thank Gerardo Lamas for
loaning me a male specimen of Styx infer-
nalis. | thank Phil Ackery and Dick Vane-
154
Wright for loaning me Ehrlich’s dissection
of a male S. infernalis.
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1958b. The comparative morphology, phy-
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PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 155-163
AEDES (STEGOMYIA) JOSIAHAE, A NEW SPECIES OF THE
SIMPSONI SUBGROUP (DIPTERA: CULICIDAE)!
YIAU-MIN HUANG
Systematics of dedes Mosquitoes Project, Department of Entomology, Smithsonian
Institution, Washington, D.C. 20560.
Abstract. —The adult male and female of Aedes (Stegomyia) josiahae n. sp. from Tan-
zania are described and illustrated. Diagnostic characters for distinguishing Ae. josiahae
from closely allied species are given. The distribution of Ae. josiahae is based on examined
specimens. 4edes josiahae is most closely related to de. kivwensis Edwards. These two
species together with de. bromeliae (Theobald), Ae. /ilii (Theobald), Ae. simpsoni (Theo-
bald), Ae. strelitziae Muspratt, Ae. subargenteus Edwards, and Ae. woodi Edwards form
the simpsonia subgroup within the aegypti group.
Key Words:
A new species of Aedes (Stegomyia), be-
longing to the simpsoni subgroup of the
aegypti group, was discovered among spec-
imens that were misidentified as Aedes
(Stegomyia) kivuensis Edwards from the
Division of Vector Borne Diseases (DVBD)
collection in Nairobi, Kenya. In view of the
medical importance of several species in the
simpsoni subgroup and the similarity of this
new species with Ae. Aivuensis Edwards, it
is desirable to describe the new species here
to make its name available and to avoid
future confusion between it and Ae. kivuen-
sis. As nothing is known about its biting
habits and potential as a vector of human
pathogens, it is hoped that this paper will
stimulate investigations on these subjects.
' This work was supported by Grant No. DAMD-
17-84-G-4033 from the U.S. Army Medical Research
and Development Command, Office of the Surgeon
General, Fort Detrick, Frederick, MD 21701, and by
the Walter Reed Biosystematic Unit, Museum Support
Center, Smithsonian Institution, Washington, D.C.
20560.
Culicidae, mosquitoes, Aedes, simpsoni subgroup
MATERIALS AND METHODS
This study is based on specimens that were
borrowed from the following institutions:
Musee royale de l’Afrique Centrale, Ter-
vuren, Belgium [CMT] and Division of
Vector Borne Diseases, Ministry of Health,
Nairobi, Kenya [DVBD]. Distributional
records are listed in the following order and
format: current country names are in capital
letters, administrative divisions, where
known, are in italics, and place names have
the first letter capitalized.
The terminology follows that of Harbach
and Knight (1980, 1981), with the exception
of “tarsal claws,” which is retained for “un-
gues.’ The venation terms follow those of
Belkin (1962).
Aedes (Stegomyia) josiahae Huang,
New SPECIES
Figs. 1-4
Female.— Head: Proboscis bearing dark
scales, lacking pale scales on ventral surface,
length as long as forefemur; maxillary pal-
pus 0.27 length of proboscis, dark, bearing
156
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Aedes (Stegomyia) josiahae n.sp. do
VOLUME 90, NUMBER 2
white scales on apical 0.50 of the total length;
pedicel covered with white scales except on
dorsal and ventral surfaces; clypeus bare;
occiput with few erect forked scales; a row
of broad white scales around eye margins;
vertex with a median stripe of broad white
scales, with broad dark scales on each side
interrupted by a lateral stripe of broad white
scales, followed ventrally by a patch of broad
white scales. Thorax: Scutum with narrow
dark scales and a distinct, median white spot
of narrow scales on anterior promontory,
followed by a submedian longitudinal stripe
of narrow yellowish scales on each side of
midline, reaching to prescutellar area and
connecting with prescutellar line of narrow
yellowish scales; fossal area with a large
patch of broader, crescent-shaped white
scales; posterior dorsocentral yellowish lines
present, reaching posterior 0.50 of scutum;
a patch of narrow white scales on lateral
margin just in front of wing root; acrostichal
setae absent; dorsocentral setae present;
scutellum with broad white scales on all
lobes and with a few broad dark scales at
apex of midlobe; antepronotum with broad
white scales; postpronotum with a patch of
broad white scales and a few narrow dark
scales dorsally; paratergite with broad white
scales; postspiracular area without scales;
hypostigmal area without scales; patches of
broad white scales on propleuron, subspi-
racular area, upper and lower portions of
mesokatepisternum, and on mesepimeron;
upper mesokatepisternal scale patch not
reaching to anterior corner of mesokatepi-
sternum; upper mesepimeral scale patch
connected to lower mesepimeral scale patch;
lower mesepimeron without setae; meta-
meron bare. Wing: With dark scales on all
veins and without a minute basal spot of
white scales on costa; cell R, 2.8 length of
LST
josiahae n. sp.
Fig. 2. Aedes (Stegomyia) josiahae n. sp. Anterior
surface of the allotype female legs.
—
Big!
Aedes (Stegomyia) josiahae n. sp., holotype male. A, Dorsal aspect of the thorax; B, Lateral aspect
of the thorax; C, Dorsal aspect of the wing; D, Anterior surface of the legs; E, Dorsal aspect of the abdomen;
F, Lateral aspect of the abdomen.
158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Postgenital \
plate ‘
t O1mm
0.2mm
$ 4 ea a C ~~ Se a
Y =
ff a ————
-” ————<$__—_— ’s
4 EL
spermathecae A Mobihal-
Aedes (Stegomyia) josiahae n. sp.
Fig. 3. Aedes (Stegomyia) josiahae n. sp. A, Sternal aspect of the female genitalia; B, Dorsal aspect of IX-
tergum; C, Dorsal aspect of VIII-sternum; D, Dorsal aspect of VIII-tergum.
VOLUME 90, NUMBER 2
— gonocoxite
claspette
Paraproct
aedeagus
IX tergum
PL -Hn LL LS
Aedes (Stegomyia) josiahae n. sp
Fig. 4.
R,..,. Halter: With dark and pale scales. Legs
(Fig. 2): Coxae with patches of white scales;
white knee-spot absent on forefemur, pres-
ent on mid- and hindfemora; forefemur an-
teriorly with a narrow, white longitudinal
stripe on ventral surface in basal 0.40; mid-
femur with a large white spot on anterior
surface about 0.67 from base; hindfemur
anteriorly with a broad white longitudinal
stripe in basal 0.66 that widens 0.25 from
base; foretibia anteriorly dark, with a basal
white band; mid- and hindtibiae all dark;
fore- and midtarsi with a basal white band
on tarsomeres 1, 2; foretarsomere | with
basal 0.20 white on dorsal surface: foretar-
somere 2 with basal 0.40-0.50 white on dor-
Aedes (Stegomyia) josiahae n. sp. Tergal aspect of the male genitalia.
sal surface; midtarsomere | with basal 0.30-
0.33 white on dorsal surface; midtarsomere
2 with basal 0.40 white on dorsal surface;
hindtarsus with a basal white band on tar-
someres I-3, the ratio of length of white
band on dorsal surface to the total length of
tarsomere is 0.25, 0.20-0.25, and 0.25;
hindtarsomere 4 all white, with a few dark
scales at apex on ventral surface; hindtar-
somere 5 all white, with a few dark scales
at apex on ventral surface; fore- and midlegs
with tarsal claws equal, all toothed; hindleg
with tarsal claws equal, both simple. Ab-
domen: Tergum I with white scales on la-
terotergite and with a median white spot;
terga II-VI each with a basal white band
160
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Aedes (Stegomyia) kivuensis Edwards °
VOLUME 90, NUMBER 2
and basolateral white spots not connecting
with basal white band; terga VII, VIII each
with basolateral white spots only; sterna III-—
VII each with a basal white band; segment
VIII largely retracted. Genitalia (Fig. 3):
Apical margin of sternum VIII with a me-
dian notch and with rounded lateral lobes;
insula longer than wide, with minute setae
and with 9 larger setae on apical 0.25; ter-
gum IX broader than long, apical margin of
tergum IX with well developed lateral lobes,
each with 4 setae; apical margin of postgen-
ital plate with a shallow median notch; cer-
cus short and broad; 3 spermathecae, one
larger than the other 2.
Male (Fig. 1).—Essentially as in the fe-
male, differing in the following sexual char-
acters: Head: Maxillary palpus 5-segment-
ed, as long as proboscis, predominantly dark,
with a white band at base of palpomeres 2-
5; those on palpomeres 4, 5 dorsally incom-
plete; palpomeres 4, 5 subequal, slender,
dorsally curved and with only a few short
setae; antenna plumose, shorter than pro-
boscis. Wing (Fig. 1C). Cell R, 3.1 length
of R,,,. Legs (Fig. 1D): Midfemur with a
large white spot on anterior surface 0.65—
0.67 from base; hindfemur anteriorly with
a broad white longitudinal stripe in basal
0.60-0.61 that widens 0.22 from base; fore-
tarsomere I with basal 0.20-0.25 white on
dorsal surface; foretarsomere 2 with basal
0.50 white on dorsal surface; midtarsomere
1 with basal 0.33 white on dorsal surface;
midtarsomere 2 with basal 0.50 white on
dorsal surface; hindtarsus with a basal white
band on tarsomeres 1-3, the ratio of length
of white band on dorsal surface to the total
length of tarsomere is 0.25—0.30, 0.25-0.33,
and 0.33; fore- and midlegs with tarsal claws
unequal, all simple. Abdomen (Fig. 1E, F):
Tergum II with basolateral white spots only;
sternum VIII with basolateral white spots
161
as long as wide (width measured 0.5 from
base), scales restricted to dorsolateral, lat-
eral and ventral surfaces, with setae on dor-
somesal surface, mesal surface membra-
nous; claspette large, broad, reaching to 0.54
of gonocoxite, distal expanded portion
square in shape in dorsal aspect, with nu-
merous simple setae on the expanded distal
portion and bearing 2(1—2) stronger, basally
widened spine-like setae on the apicomesal
angle; gonostylus simple, elongate, about
0.66 length of gonocoxite, with a short claw
process at apex and with a few setae on
apical 0.25; aedeagus strongly toothed;
paraproct with a sternal arm; cercal setae
absent; apical margin of tergum IX deeply
concave medially, with 5-7 setae on each
lateral lobe; sternum IX without setae.
Pupa and larva.— Unknown.
Type data.—Holotype male (MEP Acc.
808/#157/Tanganyika, 60-70 km. south of
Ifakara, Dr. H. Briegel) with genitalia on
slide (81/21), Ifakara, TANZANIA (Tan-
ganyika), no date (H. Briegel). Deposited in
Division of Vector Borne Diseases, Minis-
try of Health, Nairobi, Kenya. Allotype fe-
male (MEP Acc. 808/#157), same data as
holotype [DVBD]. Paratypes: | male (MEP
Acc. 808/#160) with genitalia on slide (81/
22) and | female (MEP Acc. 808/#154) with
genitalia on slide (81/23), same data as ho-
lotype [DVBD].
Other material examined. —TANZANIA
(Tanganyika). Morogoro Region: lfakara
(8°08'S, 36°41'E), Mselezi (8°46'S, 36°42’E),
H. Briegel, | F (#146) [DVBD].
Distribution.—This species is presently
known only from Tanzania (Tanganyika).
Etymology.—This species 1s named to
honor Mrs. Phoebe A. O. Josiah, Senior
Entomologist, Division of Vector Borne
Diseases, Ministry of Health, Nairobi, Ke-
—
Fig. 5.
Aedes (Stegomyia) kivuensis Edwards, holotype female. A, Dorsal aspect of the thorax; B, Lateral
aspect of the thorax; C, Dorsal aspect of the wing; D, Anterior surface of the legs; E, Dorsal aspect of the
abdomen; F, Lateral aspect of the abdomen.
162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nya, in recognition and appreciation of her
contributions to our knowledge of the mos-
quito fauna of Africa.
Taxonomic discussion.—Aedes (Stego-
myia) josiahae is a member of the simpsoni
subgroup of the aegypti group. The simp-
soni subgroup presently comprises at least
eight species (Ae. simpsoni (Theobald) 1905,
Ae. lilii (Theobald) 1910, Ae. bromeliae
(Theobald) 1911, 4e. woodi Edwards 1922,
Ae. subargenteus Edwards 1925, Ae. ki-
vuensis Edwards 1941, Ae. strelitziae Mus-
pratt 1950 and Ae. josiahae n. sp.) and is
characterized by the following combination
of characters: the scutum has a pair of sub-
median stripes, a white knee-spot is absent
on the forefemur but present on the mid-
and hindfemora; midfemur has a large, white
spot on the anterior surface and the hind-
tarsus has a basal white band on tarsomeres
1-3. Aedes josiaehae differs from all other
members of the s/mpsoni subgroup except
Ae. kivuensis, however, by the following
combination of characters: (1) scutum with
anterior median white spot of narrow scales;
(2) scutellum with broad white scales on all
lobes; (3) hindtibia anteriorly dark, without
a white stripe in basal area; (4) hindtarso-
meres 4 and 5 entirely white.
Aedes josiahae is most closely related and
similar to Ae. kivuensis, and I consider jo-
siahae to be a sister species of kivuensis.
Adults of Ae. josiahae are extremely similar
to those of Ae. kivuensis with which it has
been confused and misidentified. Aedes
joshiahae can be distinguished easily from
Ae. kivuensis, however, by the hindfemur,
which anteriorly has a broad white stripe in
basal 0.60-0.66 and by the presence of a
median white spot on tergum I. In Ae. ki-
vuensis, the hindfemur anteriorly has a broad
white stripe on the basal half, and has a
white spot about 0.62 from the base (the
white spot does not connect with the basal
white stripe) and the tergum I has white
scales only on the laterotergite (see Fig. 5).
The male genitalia of Ae. joshiahae are
easily differentiated from all other species
in the simpsoni subgroup by the claspette,
which has the distal expanded portion square
in the dorsal aspect, with numerous simple
setae on the expanded distal portion and
bearing 2(1-2) stronger, basally widened
spine-like setae on the apicomesal angle.
Gerberg and Van Someren (1970: 2) re-
ported that 4e. (Stg.) kivuensis was collect-
edin Tanzania by Dr. H. Briegel of the Swiss
Tropical Institute at Ifakara. However, the
specimens from Tanzania (Dr. H. Briegel)
in the DVBD collection are not Ae. kivuen-
sis, but are the new species joshiahae.
Aedes josiahae is apparently an East Af-
rican lowland species. Based on the present
collection data, Ae. josiahae occurs 1n hab-
itats with altitudes of 500 m (1500 ft) and
yearly rainfall of 88.90 cm (35 in.). Aedes
kivuensis is presently known only from Zaire
(Belgian Congo), where it occurs in habitats
with altitudes of 2166 m (6500 ft) and yearly
rainfall of 152.40 cm (60 in.).
Medical importance.—Unknown. How-
ever, the simpsoni subgroup is one of the
most important subgroups of Stegomyia
from the standpoint of transmission of
pathogens. Aedes bromeliae is an important
vector of yellow fever virus in East Africa.
Aedes simpsoni was incriminated in the
transmission of yellow fever virus during an
outbreak of yellow fever in Bwamba Coun-
ty, Uganda, in 1941 and yellow fever virus
has been isolated from wild caught mos-
quitoes (de. simpsoni) from Bwamba,
Uganda, (Mahaffy et al. 1942). The yellow
fever virus has also been isolated from wild
caught mosquitoes (4e. simpsoni) in Ugan-
da by Smithburn and Haddow (1946).
However, the species from which Mahaffy
et al. (1942) and Smithburn and Haddow
(1946) isolated yellow fever virus was Ae.
bromeliae, not Ae. simpsoni (see Huang
1986). Aedes simpsoni (probably Ae. bro-
meliae) from Nigeria has been shown to be
a laboratory transmitter of yellow fever
(Philip, 1929). Aedes strelitziae from South
Africa can transmit yellow fever virus from
one rhesus monkey to another under labo-
VOLUME 90, NUMBER 2
ratory conditions, as shown by Gillett and
Ross (1953).
ACKNOWLEDGMENTS
I express my sincere appreciation to
Wayne N. Mathis, Department of Ento-
mology, Smithsonian Institution, and Ron-
ald A. Ward and E. L. Peyton, Walter Reed
Biosystematics Unit, for critically reviewing
this manuscript and for their valuable com-
ments. I am most grateful to J. Decelle and
E. De Coninck, Department of Zoologie,
Section d’Entomologie, Musee Royale de
lAfrique Centrale, Tervuren, Belgium, for
the loan of the holotype of Aivwensis Ed-
wards used in this study; to J. M. D. Rob-
erts, Davy K. Koech (formerly of DVBD)
and Mrs. Phoebe A. O. Josiah, Division of
Vector Borne Diseases (DVBD), Ministry
of Health, Nairobi, Kenya, for the loan of
the specimens described above, and for their
cooperation and kind assistance during the
course of the museum and field studies on
African Stegomyia mosquitoes in Kenya. I
also express my gratitude to Young T. Sohn
and Vichai Malikul, formerly of the Medical
Entomology Project, for preparing the il-
lustrations.
LITERATURE CITED
Belkin, J. N. 1962. The mosquitoes of the South
Pacific (Diptera, Culicidae). Univ. Calif. Press,
Berkeley and Los Angeles. 2 vols., 608 and 412
pp.
Edwards, F. W. 1922. Mosquito notes.—III. Bull.
Entomol. Res. 13: 75-102.
163
1925. Mosquito notes.—V. Bull. Entomol.
Res. 15: 257-270.
1941. Mosquitoes of the Ethiopian region.
III. Culicine adults and pupae. Br. Mus. (Nat. Hist.),
Lond. 499 pp.
Gerberg, E. J. and E.C. C. Van Someren. 1970. Pic-
torial key to the mosquitoes dedes (Stegomyia) of
East Africa. WHO/VBC/70.236: 1-7.
Gillett, J. D. and R. W. Ross. 1953. The laboratory
transmission of yellow fever by the mosquito Aedes
(Stegomyia) strelitziae Muspratt. Ann. Trop. Med.
Parasit. 47: 367-370.
Harbach, R. E.and K. L. Knight. 1980. Taxonomists”
glossary of mosquito anatomy. Plexus Publishing,
Inc., Marlton, N.J. 415 pp.
1981(1982). Corrections and additions to
taxonomists’ glossary of mosquito anatomy. Mosq.
Syst. 13: 201-217.
Huang, Y.-M. 1986. Aedes (Stegomyia) bromeliae
(Diptera: Culicidae), the yellow fever virus vector
in East Africa. J. Med. Entomol. 23: 196-200.
Mahaffy, A. F., K. C. Smithburn, H. R. Jacobs, and J.
D. Gillett. 1942. Yellow fever in western Ugan-
da. Trans. R. Soc. Trop. Med. Hyg. 36: 9-20.
Muspratt, J. 1950. Notes on 4edes (Diptera, Culic-
idae) from Natal, with a description ofa new species
of the subgenus Stegomyia. J. Entomol. Soc. Afr.
13: 73-79.
Philip, C. B. 1929. Preliminary report of further tests
with yellow fever transmission by mosquitoes oth-
er than Ades aegypti. Am. J. Trop. Med. 9: 267-
269.
Smithburn, K. C. and A. J. Haddow. 1946. Isolation
of yellow fever virus from African mosquitoes.
Am. J. Trop. Med. 26: 261-271.
Theobald, F. V. 1905. A new Stegomyia from the
Transvaal. Entomol. 38: 224-225.
. 1910. A monograph of the Culicidae or mos-
quitoes. Vol. V. Br. Mus. (Nat. Hist.), Lond. 646
pp., 6 pl.
1911. Uganda Culicidae including thirteen
new species. Novae Culicidae. Part I, 35 pp.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 164-178
DESCRIPTION AND BIOLOGY OF ACROLOPHUS PHOLETER,
(LEPIDOPTERA: TINEIDAE), A NEW MOTH COMMENSAL
FROM GOPHER TORTOISE BURROWS IN FLORIDA
DONALD R. DAVIS AND ERIC G. MILSTREY
(DRD) Department of Entomology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560; (EGM) U.S. Army Environmental Hygiene Agency,
Field Support Activity, Fort Mead, Maryland 20755.
Abstract.—Vacuum sampling of gopher tortoise burrows in Putnam County, Florida,
has resulted in the discovery of a new species of tineid moth, Acrolophus pholeter Davis.
The larva feeds on both the fecal pellets of the gopher tortoise and upon decaying plant
debris within the burrow. Supplemented by numerous illustrations, the larval, pupal, and
adult stages are described, and the general biology 1s summarized.
Key Words:
toise
Recent vacuum sampling for invertebrate
commensals in the burrows of the gopher
tortoise, Gopherus polyphemus Daudin, by
the junior author has revealed the presence
of a few arthropods previously unreported.
Among these was a new species of Acrolo-
phus that was found in abundance feeding
on both tortoise fecal pellets and decaying
plant debris within the burrow.
This is the first record of an Acrolophus
commensal in an animal burrow. The sub-
terranean, tube-constructing habit of the ge-
nus, however, is well known. Other Tine-
idae (all Acrolophinae) have been reported
from rodent burrows (Hubbard 1901, Hub-
bell and Goff 1939, Davis et al. 1986), and
at least one other moth, /dia gopheri (Smith),
is known to inhabit the burrows of the go-
pher tortoise (Hubbard 1894, 1896, Smith
1899, Woodruff 1982). Hubbell and Goff
(1939) reported that some arthropod com-
mensals were true obligates and had not been
collected outside gopher tortoise burrows.
It is not known to what extent Acrolophus
pholeter n. sp. 1s restricted to burrows of
Lepidoptera, Tineidae, moth biology, Gopherus polyphemus, gopher tor-
this tortoise or if the moth also frequents
rodent burrows. The absence of previous
Acrolophus collecting records suggests that
its habitat may be rather restricted.
Since at least the Pleistocene, gopher tor-
toise burrows have provided a relatively
stable habitat for the establishment of a di-
verse community of organisms. The integ-
rity of individual burrows is normally main-
tained for five years or more. In terms of
numbers of both vertebrate and inverte-
brate species found using gopher tortoise
burrows, the diversity is one of the greatest
yet studied in North American animal bur-
rows (Milstrey 1986).
Sampling of organisms from gopher tor-
toise burrows can be a formidable task. De-
pending upon soil type and water table, bur-
rows may extend up to 40 feet long and 12
feet deep (Young and Goff 1939). Excava-
tion of such galleries can create a sizeable
trench (Hubbard 1894). In recent years the
use of vacuum suction devices (Butler et al.
1984) has greatly facilitated collecting from
burrows without decimating the landscape
VOLUME 90, NUMBER 2 165
Figs. 1-5. Acrolophus pholeter, habitat, adults, and cocoon. 1, Entrance to gopher tortoise burrow, Gopherus
polyphemus, Putnam Co., Florida. 2, Modified Echo R200 blower used for sampling invertebrate fauna from
burrows. 3, Adult male, length of forewing 6.7 mm. 4, Adult female, length of forewing 8.2 mm. 5, Cocoon
with pupal exuvium protruding, length of cocoon 13 mm.
166
(Fig. 2). One disadvantage of this technique
is that it does not allow direct observations
of the organisms’ biology.
All material used in this study was col-
lected by the junior author using a modified
Echo R200 (R) blower (Kioritz Corp., To-
kyo, Japan). Most adults (all type material)
were reared from larvae collected by vac-
uuming.
Deposition of specimens referred to in
this paper are: BMNH for British Museum
of Natural History, London, England; FSCA,
Florida State Collection of Arthropods,
Gainesville, Florida; and USNM, National
Museum of Natural History (formerly
United States National Museum), Smith-
sonian Institution, Washington, D.C.
Acrolophus pholeter Davis,
New SPECIES
Figs. 3-62
Adult (Figs. 3-4).—Length of forewing:
male, 5.5-6.7 mm; female, 7-9.5 mm. A
moderately small moth with uniformly
brownish gray wings, smooth head, and
short labial palpi.
Head: Vestiture smooth over vertex and
frons; scales uniformly brownish gray, rel-
atively slender with rounded apices, ap-
pearing to arise from lower frons and cury-
ing upwards over vertex until they reach
flattened, transversely oriented scale patches
across occiput. Eye small, interocular index
approximately 0.65, cornea relatively
smooth with only scattered microsetae (Figs.
10-11); eyelash absent. Antenna about 0.4—
0.6 the length of forewing, relatively longer
in male, 46-49 segmented; scape uniformly
brownish fuscous, smooth, without pecten;
flagellomeres of similar color, fully scaled,
simple in form with a few sensilla coelo-
conica along anterior margin (Figs. 12-13).
Pilifer reduced, minutely setose (Fig. 7).
Mandible absent. Maxillary palpus greatly
reduced, 2 segmented; approximately 8-10
elongate sensilla arising from apical pit.
Haustellum absent. Labial palpus_ short;
length approximately 2.5= eye diameter,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
uniformly brownish gray, relatively smooth
vestiture with slightly rough scales along
venter of second segment.
Thorax: Pronotum and both fore and
hindwings uniformly brownish gray. Venter
somewhat paler, more light brown. Legs with
smooth vestiture, light brownish gray dor-
sally, lighter buff ventrally; epiphysis greatly
reduced (Fig. 14), length only about twice
its width. Pretarsus of all legs unspecialized,
with symmetrical claws, pulvilli, well de-
veloped arolium, and unguitractor plate
bearing 5—6 transverse rows of scutes.
Abdomen: Uniformly light brownish gray.
Male genitalia: As shown in Figs. 18-21.
Uncus elongate, slender, and acute. Tegu-
men relatively broad and elongate. Vincu-
lum slender; anterior margin slightly con-
cave at middle. Gnathos a well developed
median lobe with a relatively broad, trun-
cate apex. Valva rather broad over basal
half to sharply defined saccular lobe,
abruptly narrowing beyond lobe to simple
apex. Aedoeagus relatively short, approxi-
mately two-thirds the length of valva, and
without cornuti; apex with a serrated cleft
extending over one-fourth down right side.
Female genitalia: As shown in Fig. 22.
Only a single pair of short, posterior apophy-
ses present. Caudal margin of lamella an-
tevaginalis smoothly curved. Ductus bursae
very short, slightly thickened and constrict-
ed just before corpus bursae; latter simple,
relatively small, membranous sac without
spicules.
Larva (Figs. 33-62).—Length of largest
larva 18 mm; diameter 2.1 mm. Body trans-
lucent, light yellowish brown with light
brown thoracic and anal plates.
Head: Uniformly light reddish brown,
darker around base of mandibles. Greatest
width 1.4 mm, length 0.9 mm. AF2 arising
well above (caudad) apex of frons. P2 more
distant from P1 than P1 is to ecdysial line.
Stemmata vestigial, probably non-function-
al; only three transparent vestiges remaining
(Fig. 53); one situated above S2, a very small
one well below S2, and an elongate hyaline
VOLUME 90, NUMBER 2 167
Figs. 6-11. Adult structure, Acrolophus pholeter. 6, Partial view of frons and mouthparts (88 um). 7, Maxillary
palpi and pilifers (37.5 um). 8, Sensilla at apex of maxillary palpus (2.5 um). 9, Sensilla at apex of labial palpus
(8.8 um). 10, Eye (88 um). 11, Detail of cornea showing scattered interfacetal microsetae (19 um). (Scale lengths
in parentheses.)
168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ail
Figs. 12-17. Adult structure, Acrolophus pholeter. 12, Antenna near middle of flagellum (37.5 wm). 13, Detail
of Fig. 12 showing sensilla coeloconica (16.5 um). 14, Reduced epiphysis on foretibia (25 wm). 15, Detail of
epiphysis (7.5 um). 16, Pretarsus of hindleg (19 um). 17, Detail of unguitractor plate (3.75 um). (Scale lengths
in parentheses.)
169
VOLUME 90, NUMBER 2
\
view (0.5 mm). 19, Lateral view. 20, Lateral
Figs. 18-24. Acrolophus pholeter. 18, Male genitalia, ventral
lia, ventral view (0.5 mm). 23, Pupa, ventral
view of valva. 21, Lateral view of aedoeagus. 22, Female genita’
view (2 mm). 24, Dorsal view. (Scale lengths in parentheses.)
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 25-30. Acrolophus pholeter, pupa. 25, Head, ventral view (0.3 mm). 26, Lateral view (0.22 mm). 27,
Caudal end (A7-10) of abdomen (0.22 mm). 28, Lateral view, dorsum up (0.19 mm). 29, Dorsal view (0.22
mm). 30, Detail of dorsal cremaster, A10 (88 um). (Scale lengths in parentheses.)
VOLUME 90, NUMBER 2 LA
Figs. 31-36. Acrolophus pholeter. 31, Pupa, dorsum of A4—5 (0.43 mm). 32, Detail of spine row, AS (30
um). 33, Larva, dorsal view of head (0.19 mm). 34, Dorsal view of labrum and mouthparts (68 um). 35, Lateral
view of head (0.17 mm). 36, Lateral view of stemmatal area (60 um). (Scale lengths in parentheses.)
172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 37-42. Acrolophus pholeter, larva. 37, Head, frontal view (0.15 mm). 38, Maxillae and labium (68
um). 39, Detail of maxilla (22 um). 40, Ventral view of maxillae and labium (88 wm). 41, Detail of spinneret
and labial palpi (37.5 um). 42, Detail of secondary labial setae in Fig. 41 (3.75 um). (Scale lengths in parentheses.)
VOLUME 90, NUMBER 2
Figs. 43-48. Acrolophus pholeter, larva. 43, Antenna (43 um). 44, Detail of antennal apex (12 um). 45,
Lateral view of prothorax (0.22 mm). 46, Ventral view of prothorax (0.22 mm). 47, Tarsal claw (11.5 um). 48,
Prolegs, AS (0.15 mm). (Scale lengths in parentheses.)
174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
RE PAR
:
Pee 77)
Figs. 49-54. Acrolophus pholeter, larva. 49, Crochets on proleg 5 (38.5 um). 50, Anal proleg, A10 (60 um).
51, Segments A9-10, dorsal view (0.19 mm). 52, Lateral view (0.19 mm). 53, Ventral view (0.19 mm). 54,
Caudal view (0.19 mm). (Scale lengths in parentheses.)
VOLUME 90, NUMBER 2
175
Figs. 55-62. Acrolophus pholeter, larva. 55, Chaetotaxy of body, segments T1—2, Al, 6, 8-9. 56, Head, dorsal
view (0.5 mm). 57, Ventral view. 58, Segments A8-10, dorsal view. 59, Head, lateral view. 60, Labrum, dorsal
view (0.2 mm). 61, Ventral view. 62, Mandible (0.2 mm). (Scale lengths in parentheses.)
area anterior to S2; the latter apparently the
remnant of three fused stemmata. Apical
segment of antenna relatively long; sensilla
as in Figs. 43-44. Labrum with M1 and 2
near anterior margin; M3 more remote.
Mandible somewhat tapered, with 4 small
cusps. Maxilla as in Figs. 38-39. Spinneret
elongate, slender; labial palpus 2-segment-
ed, basal segment elongate with a short api-
cal seta; apical segment greatly reduced,
176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
about 0.25 the length of basal segment, with
an elongate seta nearly equal in length to
both segments; apex of mentum with a pair
of minute secondary labial setae (Figs. 41—
42).
Thorax: Pronotal and mesonotal plates
light brown. Spiracular plate almost com-
pletely separated from pronotal plate; all 3
lateral setae together on spiracular plate.
Coxal plates separated slightly. Tarsal claw
as in Fig. 47.
Abdomen: A\-6 with 11 pairs of primary
setae, SV trisetose. Ventral crochets in a
uniserial ellipse of approximately 30 hooks;
sides of proleg densely covered with small,
scattered spines, anal proleg with approxi-
mately 21 hooks in a half ellipse open to
the rear. A8 with spiracle greatly enlarged,
equalling size of prothoracic spiracle; 10
pairs of primary setae, SV bisetose. A9 with
9 setal pairs, SV unisetose. Anal plate light
brown, bearing 4 pairs of setae.
Pupa (Figs. 23-32).—Length of largest
pupa: male, 7.2 mm; female, 11 mm. Light
reddish brown in color. Vertex smooth ex-
cept for a pair of minute setae. Antenna and
labial palpus of relatively equal length in
both sexes; antenna extending to caudal
margin of A3 and just short of wings which
extend to caudal margin of A4; labial palpus
short. Mesonotum with two pairs of minute
setae clustered together near midline. Dor-
sum of A3-8 with a transverse ridge like
row of minute spines near anterior margin.
A9 + 10 relatively smooth except for a clus-
ter of 3 pairs of minute spines ventrally and
a large, slightly bilobed ridge dorsally (Figs.
28-30).
Holotype.— Male, (with associated pupal
exuvium and cocoon) Roberts’ Ranch, ca.
6 km north of Hollister, Putnam Co., Flor-
ida; em. 12 May 1985, E. G. Milstrey,
(USNM).
Paratypes. FLORIDA: Same data as
holotype except: 2 6, 3 2, em. Sept. 1984; 5
6, 42,em. Nov. 1984; 3 2, em. 8 Dec. 1984;
3.6, 1 95 em: 19) Dec; 1984; 1d;,m:.5 Feb:
1985; 3 2,em. 5 May 1985; 1 2, 9 May 1985;
em. 11 May 1985; 2 6, 3 9, em. 12 May
1985; 4 6, 1 9, em. 17 May 1985; 1 2, 12
June 1985, em. 30 June 1985; 1 2, em. 20
July 1985; 1 9, 10 July 1985, em. 23 July
1985; 60 larvae, 26 July 1985; 68 larvae, 9
Sept. 1985. Paratypes deposited in BMNH,
FSCA, and USNM.
Host.—Larval substrate consists of both
decaying plant debris within burrow of go-
pher tortoise and fecal pellets of tortoise.
Flight period.— Difficult to assess; adults
were collected from burrows through much
of the year.
Distribution.— Known only from under-
ground burrows of the gopher tortoise in the
sandhill habitat of Putnam County, north-
eastern Florida.
Etymology.—The specific name is de-
rived from the Greek pholeter (one who lurks
in a hole), in reference to its subterranean
behavior.
Discussion.—Acrolophus pholeter does
not appear closely allied to any North
American Acrolophus. In addition to its dis-
tinctive male genitalia, this species is un-
usual in possessing a smooth head and
greatly reduced epiphysis. In color pattern,
it superficially resembles a nearly unicol-
orus, undescribed species from southern
Florida and Texas.
Even less can be summarized about larval
relationships because of the great inadequa-
cy of our knowledge. Compared with the
few Acrolophus larvae ever studied (e.g. Da-
vis 1987), the chaetotaxy of A. pholeter ap-
pears little differentiated. However, the
atypical stemmatal reduction in this species,
particularly the apparent fusion of the three
anterior stemmata, is probably character-
istic for the species.
Biological observations. — Biology of this
species has been determined from labora-
tory and field observations. Apparently, the
species is restricted to burrows of the gopher
tortoise, Gopherus polyphemus, in sandhill
habitats (Fig. 1). Tortoise burrows sampled
outside the sandhill biome were found not
to contain A. pholeter. The sandhills are rel-
VOLUME 90, NUMBER 2
ict dunes from the Pleistocene and earlier
epochs (Cooke 1945, Laessle 1958). The
vegetation of this habitat is characterized
by longleaf pines, Pinus palustris, several
oaks, Quercus laevis and margaretta, with
an understory of wiregrass, Aristida stricta,
and various herbs. The study site was in
Putnam County, 6 km north of Hollister
(29°41'40”N-8 1°48'10’"W).
Collections were made every six weeks by
vacuum extraction of the burrow using a
modified Echo leaf blower similar to that
described by Butler et al. (1984). Larvae,
pupae and adults were obtained but eggs
were not detected. Larval densities ranged
up to 30 per burrow. In repeatedly sampled
burrows, population estimates, based upon
removal sampling estimates (Carle and
Maughan 1980), indicated larval numbers
commonly were 3 to 16. A few burrows not
sampled routinely gave higher estimates, up
to as high as 200. First instar larvae were
found from May to November. All other
stages were present year round. In the later
part of the summer and early fall the pu-
pating larvae probably were of two gener-
ations; the previous year’s and offspring of
the spring emergence. Larval development
was estimated to require anywhere from 7
to 16 months and most likely around 11
months (+ one month) for most individu-
als. One lab reared larva, field collected in
its last larval stage, took 15 months to pu-
pate. The pupation period is relatively short,
normally requiring one to two weeks for
adult eclosion.
Pupae and adults were infrequently col-
lected in the burrows from May to Septem-
ber. Pupal cocoons (Fig. 5) were constructed
of loosely woven silk to which are attached
sand grains and larval frass. Apparently, pu-
pation was triggered by unknown narrow
microclimatic conditions present in indi-
vidual burrows, because when pupae were
found there were commonly more than one,
and in other adjacent burrows none were
found. Adults collected from burrows had
wings that were badly tattered with few scales
177
remaining on the body and wings. The poor
condition of the adults was not due to the
sampling method but probably due to abra-
sion occurring during normal adult activity
within the sandy burrows.
As larval size increased in the burrows,
density decreased. Larvae were found to be
cannibalistic in laboratory studies. Silk lined
larval galleries were usually constructed just
below the soil surface in the floor of the
burrow. Larval galleries could exceed 30 cm
in length and were often branched. Larvae
traveled forward and backward in these gal-
leries, always facing the same direction.
Feeding occurred at the entrance and fecal
pellets were deposited on the soil surface at
the other end.
Larvae were successfully reared on go-
pher tortoise fecal pellets. The fecal pellets
used were mostly partially disgested wire-
grass and some oak leaves. Although fecal
material was available in the burrows, its
abundance was always low and competition
for it was high. The amount of available
fecal material was generally too low to sup-
port the Acro/ophus population present.
Also, larvae were very common in burrows
that no longer had resident tortoises. In
active burrows, tortoises frequently re-ex-
cavated their burrows, and in those burrows
larval populations were significantly de-
creased if not exterminated. Larval density
was positively correlated with burrows that
accumulated leaves and other debris. Ap-
parently this rapidly decomposing organic
leaf litter was their primary food source.
Laboratory studies on limited numbers of
larvae found that they would survive on the
litter but that growth rates were lower.
The unique humidity situation 1n the bur-
row appears to be responsible for restricting
the species to this habitat. In the sandhill
gopher tortoise burrows studied, relative
humidities were in the mid to high 90’s but
the percent water in the soil was low (0-5%)
year round. In the laboratory inside an en-
vironmental chamber, larvae were found to
survive only within these limits. Mature lar-
178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vae left their silk galleries and moved to
drier conditions to pupate. Pupae in the lab-
oratory could be reared at room tempera-
ture and normal humidity levels.
The larvae of the antlion, Glenurus gratis
(Say), and larvae of an undescribed therivid
fly, Arenagena sp. were found to prey upon
the Acrolophus larvae. Lepidoptera larvae
were the only prey either would take under
laboratory conditions. Glenurus gratis was
probably the major predator. Its larval pop-
ulation numbers were more strongly cor-
related with Acrolophus larval numbers.
Both prefer the same burrows and both are
restricted to the drier, looser sand near the
entrances where the leaves collect; the ther-
ivid was more ubiquitous.
ACKNOWLEDGMENTS
The senior author (Davis) is responsible
for the systematic and morphological por-
tions of this paper and the biological ob-
servations are by the junior author (Mil-
strey). Thanks are given to Irwin Roberts
for allowing his land to be used for this re-
search, Judy Gilmore for adult preparation
and Dr. Dale Habeck for recognizing the
uniqueness of the species and encouraging
its investigation. We are indebted to Young
Sohn of the Department of Entomology,
Smithsonian Institution, for the line draw-
ings and to Susann Braden and Brian Kahn
of the Smithsonian SEM Lab and Victor
Kranz of the Smithsonian Photographic
Laboratory for photographic assistance. The
final draft was prepared by Silver West.
LITERATURE CITED
Butler, J. F., K. H. Holscher, O. Adeyeye, and E. P. J.
Gibbs. 1984. Sampling techniques for burrow
dwelling ticks in reference to potential African
swine fever virus vectors, p. 1065-1074. In D. A.
Griffiths and C. E. Bowman, eds., Acarology IV
Volume 2. Ellis Horwood Ltd. Chichester, En-
gland.
Carle, F. L. and O. E. Maughan. 1980. Accurate and
efficient estimation of benthic populations: A
comparison between removal estimation and con-
ventional sampling techniques. Hydrobiologia 71:
181-187.
Cooke, C. W. 1945. Geology of Florida. Florida Geo-
logical Survey Bulletin, 29: 1-339.
Davis, D. R., D. H. Clayton, D. H. Janzen, and A. P.
Brooke. 1986. Neotropical Tineidae, II: Biolog-
ical notes and descriptions of two new moths phor-
etic on spiny pocket mice in Costa Rica (Lepi-
doptera: Tineoidea). Proceedings of the
Entomological Society of Washington 88(1): 98—
109.
Davis, D. R. 1987. Tineidae, pp. 362-365. In F. W.
Stehr, ed., Immature Insects. Kendall/Hunt Pub-
lishing Co., Dubuque, Iowa.
Hubbard, H.G. 1894. The insect guests of the Florida
land tortoise. Insect Life 6(4): 302-315.
1896. Additional notes on the insect guests
of the Florida land tortoise. Proceedings of the
Entomological Society of Washington 3(5): 299-
302.
1901. Letters from the southwest: Insect fau-
na in the burrows of desert rodents. Proceedings
of the Entomological Society of Washington 4:
361-364.
Hubbell, T. H. and C. C. Goff. 1939. Florida pocket-
gopher burrows and their arthropod inhabitants.
Proceedings of the Florida Academy of Science,
4: 127-166.
Laessle, A. M. 1958. The origin and successional
relationship of sand-hill vegetation and sand pine
scrub. Ecological Monographs 28: 361-387.
Milstrey, E. G. 1986. Ticks and invertebrate com-
mensals in gopher tortoise burrows: Implications
and importance, Pp. 4-15. Jn D. R. Jackson and
R. J. Bryant, eds., The Gopher Tortoise and its
Community. Proceedings 5th Annual Meeting of
the Gopher Tortoise Council. Florida State Mu-
seum, University of Florida, Gainesville, Florida.
Smith, J. B. 1899. Description of the gopher moth.
Canadian Entomologist 31: 94-95.
Woodruff, R.E. 1982. Arthropods of gopher burrows,
pp. 25-49. Jn R. Franz and R. J. Bryant, eds., The
Gopher Tortoise and its Sandhill Habitat. Pro-
ceedings of the 3rd Annual Meeting of the Gopher
Tortoise Council. Tall Timbers Research Station,
Tallahassee, Florida.
Young, F. N. and C. C. Goff. 1939. An annotated list
of the arthropods found in the burrows of the Flor-
ida gopher iortoise, Gopherus polyphemus (Dau-
din). The Florida Entomologist 22(4): 53-62.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 179-188
NEW SPECIES OF CERAMBYCIDAE FROM TWIN CAYS,
BELIZE (COLEOPTERA)
JOHN A. CHEMSAK AND CANDY FELLER
(JAC) University of California, Berkeley, California 94720; (CF) McAAP, McAlester,
Oklahoma 74501.
Abstract.—New species of Cerambycidae (Coleoptera) from Twin Cays, Belize, asso-
ciated with mangrove, are described along with a description of the male of Derancistrus
fellerae Chemsak. New taxa are: Methia rhizophorae, Ataxia cayensis, Leptostylopsis latus,
Stvloleptus rhizophorae, and Urgleptes ozophagus.
Key Words:
The recent interest in and emphasis on
Neotropical natural history has resulted in
a great increase in biological investigations
on insects. The Cerambycidae, because of
their larval habits, are directly affected by
the well documented loss of tropical forests
(Janzen 1986). For this reason, studies deal-
ing with habits and behavior of immatures
and adults will attain increasing impor-
tance.
One such study, presently being con-
ducted by C. Feller at Twin Cays, Belize,
has produced a number of new species. Twin
Cays (16°50'N, 18°06'W), locally called
Water Range, 1s a swampy mangrove island
within the Belize Barrier Reef. The island
is approximately | km in diameter and is
22 km SE of Dangriga (Stann Creek), Dan-
griga District, Belize. The Cerambycidae
present appear to be associated with red
mangrove, Rhizophora mangle L., black
mangrove, Avicennia germinans (L.) Stearn,
white mangrove, Laguncularia racemosa
(L.) Gaertn. f., and buttonwood, Conocar-
pus erectus (L.). The beetle genera repre-
sented are typical West Indian—Central
American groups containing numerous
species (Chemsak and Linsley 1982).
In 1983, Chemsak described Derancistrus
Insecta, Coleoptera, Cerambycidae, Belize
fellerae from Twin Cays from two female
specimens. Subsequent collections and rear-
ings have produced additional specimens,
including the male, which is described be-
low along with five new species. The new
taxa are presented to make their names
available for current ecological studies.
Types are deposited in the United States
Museum of Natural History, Washington,
D.C. and paratypes in the Essig Museum of
Entomology, University of California,
Berkeley.
Derancistrus fellerae Chemsak
(Prioninae)
Fig. |
Derancistrus fellerae Chemsak 1983, Proc.
Entomol. Soc. Wash., 85: 714.
Male.—Form moderate-sized, rather
slender, tapering posteriorly; integument
black, elytra black basally, dark reddish to-
ward apex, femora reddish, narrowly black
at apices and bases, tibiae reddish toward
apical one-half. Head narrow, front short,
deeply impressed, impression extending
onto vertex; punctures coarse, subconfluent,
becoming sparse on neck; pubescence sparse,
recurved; antennae serrate, extending to
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
180
male.
>
Derancistrus fellerae Chemsak
Fig. 1.
VOLUME 90, NUMBER 2
about third abdominal segment, segments
broad, flattened, basal segments sparsely,
shallowly punctate, poriferous area small on
third segment, areas increasing to sixth seg-
ment, segments from seventh entirely
opaque, third segment longer than first,
fourth subequal to first, eleventh longer than
tenth. Pronotum broader than long, sides
gradually expanding back from apex to be-
hind middle then narrowing to base, mar-
gins crenulate, angles dentate; disk shallow-
ly impressed at middle behind apical margin;
punctures irregular, middle with irregular
glabrous areas; sides behind dentate angles
with a small patch of whitish recumbent
pubescence; prosternum sparsely pubes-
cent; meso- and metasternum with patches
of white appressed pubescence at sides. Ely-
tra about twice as long as broad; punctures
at base coarse, confluent, becoming finer to-
ward apex; apices serrated. Legs slender,
sparsely punctate and pubescent. Abdomen
with sternites narrowly glabrous on poste-
rior margins; last sternite emarginate at apex.
Length, 18 mm.
The male is much smaller and more slen-
der than the female. The antennae are long-
er with opaque outer segments, and pubes-
cent patches are lacking along the anterior
margins and base of the pronotum.
Larvae of this species have been collected
from the wood of red mangrove, black man-
grove, white mangrove, and buttonwood.
One specimen is deposited in the USNM
and the other in the Essig Museum.
Methia rhizophorae Chemsak & Feller,
New Species (Methiini)
Female.— Form slender, rather elongate,
elytra dehiscent toward apex; color fuscus,
head, pronotum and appendages at least
partially orange-brown, elytra often paler
along suture at basal one-half. Head slightly
wider than pronotum; eyes separated on
vertex by less than diameter of third anten-
nal segment, separated beneath by more than
diameter of scape, lobes of eyes connected
by a single row of facets; front finely asper-
181
ate; antennae extending about four seg-
ments beyond body, scape lacking an apical
tooth, basal segments dark apically, pubes-
cence rather short, dense, suberect; neck
deeply, narrowly impressed medially be-
hind eyes. Pronotum broader than long,
sides broadly rounded; apex and base trans-
versely impressed; disk convex, medially
impressed before basal margin; punctures
finely scabrous; pubescence pale, dense,
subdepressed, long; stridulatory plate of
mesonotum not grooved; prosternum nar-
rowly impressed at apex, finely plicate. Ely-
tra extending to second abdominal segment;
each elytron vaguely bicostate; punctures
moderately coarse, shallow, subconfluent;
pubescence pale, moderately dense, sube-
rect. Legs moderate; femora finely, trans-
versely plicate; pubescence moderately
dense. Abdomen with last sternite deeply
v-shaped at apex, margins with a row of
setae. Length, 10-11 mm.
Male.— Form small, slender; color brown-
ish, antennae pale orange-brown on basal
segments, scape infuscated over apical two-
thirds, elytra pale at middle and at apex.
Head with front asperate; eyes separated
above by less than diameter of third anten-
nal segment, beneath by more than diam-
eter of scape, lobes connected by a single
row of facets; antennae extending almost
five segments beyond body, segments
densely clothed with short erect and sub-
erect pubescence. Elytra brownish basally
with vague brownish vittae extending down
middle of disk to preapical depressions to
form transverse dark spots; punctures fine,
dense, confluent basally and contiguous to-
ward apex; pubescence pale, short, subde-
pressed. Abdomen not modified. Length, 6—-
7 mm.
Holotype 2, allotype, from Twin Cays,
Belize, 4 June 1985, 21 May 1986, in red
mangrove (C. Feller); paratypes, | 6, 21 May
1986; 2 2, 2 June 1985, 21 May 1986, from
red mangrove.
This species resembles WV. constricticollis
Schaeffer from Texas and Mexico by the
182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
coloration of the antennae. The two differ
by the coarsely punctate head, the eye lobes
connected by 2-3 facets and by the feebly
rugose elytra in M. constricticollis.
The female paratypes are slightly paler in
color with a pale vitta along the basal half
of the suture of the elytra.
Ataxia cayensis Chemsak & Feller,
NEw SPECIES (Ataxiini)
Fig. 2
Male.— Form moderate-sized, slightly ta-
pering posteriorly; integument piceous, ap-
pendages slightly reddish-piceous; pubes-
cence moderately dense, grayish, fine,
appressed. Head with front rather coarsely,
deeply punctate, punctures well separated,
pubescence appressed, interrupted by punc-
tures, recurved setae arising from each
puncture with longer erect hairs around eyes
and mouthparts; vertex with a median line,
sparsely punctate; antennae about as long
as body, segments from fourth paler at bas-
es, segments finely pubescent, long, erect
hairs beneath numerous, scape with a fine,
apical cicatrix, third segment subequal to
first, fourth longer than third. Pronotum
slightly broader than long, lateral tubercles
small; disk very coarsely, irregularly punc-
tate, medially shallowly impressed behind
middle; pubescence mottled, appressed, with
recurved setae arising from punctures; pro-
sternum coarsely, sparsely punctate, rather
sparsely pubescent; mesosternum with in-
tercoxal process arcuately declivous; meta-
sternum with pubescence interrupted by
small glabrous spots. Elytra about twice as
long as broad; disk shallowly costate with
coarse punctures arranged linearly down in-
tervals, punctures becoming finer toward
apex; pubescence fine, mottled, with whitish
flecks of denser pubescence interspersed,
suberect hairs short; apices subtruncate. Legs
moderately densely pubescent, pubescence
interrupted by small spots. Abdomen mod-
erately densely pubescent, small glabrous
spots numerous; last sternite very shallowly
emarginate at apex. Length, 11 mm.
Female.—Form similar to male. Anten-
nae slightly shorter than body. Abdomen
with last sternite shallowly impressed, trun-
cate at apex. Length, 10.5-12 mm.
Holotype 4 and one 2 paratype from Twin
Cays, Dangriga Dist., Belize, 4-5 June 1985.
‘““Fogging Proj. Black Mang.” (T. L. Erwin,
L. L. Sims, W. N. Mathis); one female para-
type, Twin Cays, emerged 29 May 1986,
from larva from red mangrove twig termi-
nal (C. Feller); one 2 paratype, Twin Cays,
mudflat nr. Lairchan, 15-19 January 1987
(W. N. Mathis, C. Feller).
This species 1s distinctive by its small size,
dark integument with paler appendages, and
fine, dense pubescence of the elytra with
small whitish flecks interspersed.
The antennae and legs of the first paratype
are paler reddish than those of the type.
Leptostylopsis latus Chemsak & Feller,
New Species (Acanthocini)
Fig. 3
Female.—Form moderate-sized, broad;
integument reddish brown, underside par-
tually infuscated; pubescence dense, short,
fine, appressed, pale and dark brownish.
Head with front about as broad as long,
micropunctate with larger punctures sparse-
ly interspersed, pubescence fine, pale
brownish, interrupted by glabrous spots;
genae as long as lower eye lobes; upper eye
lobes small, separated by diameter of an-
tennal scape; antennae a little longer than
body, segments dark annulate at bases and
apices, segments to fifth with small dark
spots, third segment slightly longer than first,
fourth shorter than first. Pronotum much
broader than long, sides tumid, vaguely tu-
berculate slightly behind middle; disk with
five shallow calluses, three median more
prominent; punctures around median callus
sublinearly arranged between transverse ru-
gosities, punctures at sides finer, scattered;
pubescence fine, interrupted by punctures,
broadly dark medially, small dark spots in-
terspersed over remaining surface; proster-
num finely pubescent, intercoxal process
VOLUME 90, NUMBER 2
Fig. 2. Ataxia cayensis Chemsak & Feller, male.
more than one-half as broad as coxal cavity;
meso- and metasternum with darker spots,
mesosternal process broader than coxal cav-
ity. Elytra about 1!” times as long as broad,
tapering apically; basal gibbosities shallow,
impression behind deep; disk with a few
tufted tubercles down costae, basal ones
elongate; pubescence dense, appressed,
brownish, each side with a dark lateral vitta
183
5
extending from humerus to a little behind
middle, basal impressions dark, apical one-
half with dark linear markings on suture and
on disk before apex, tubercles pale pubes-
cent; apices narrowly, shallowly emarginate
truncate. Legs robust, femora with small
spots; tibiae dark biannulate; tarsi dark, first
segments basally and claws pale. Abdomen
thinly pubescent, sternites dark along apical
184
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Leptostylopsis latus Chemsak & Feller, female.
VOLUME 90, NUMBER 2
margins; last sternite slightly elongate, apex
narrowly truncate. Length, 13 mm.
Holotype 2? from Twin Cays, Belize, 5 May
1985 (26-85), in red mangrove (C. Feller
and Canupp).
The broad form and distinctive puncta-
tion and rugosities of the pronotum make
this species distinctive. The type is slightly
teneral, probably making the integumental
coloration paler than in fully mature spec-
imens.
Styloleptus rhizophorae Chemsak & Feller,
New Species (Acanthocini)
Fig. 4
Male.— Form small, subdepressed; integ-
ument pale reddish-brown, head infuscated,
antennae with scape dark dorsally, other
segments dark annulate at apices, legs with
tibiae broadly dark biannulate, femora with
clavate portion dark on outside and inside,
elytra with dark fasciae and an irregularly
margined, oblique whitish fascia on each
side at middle; pubescence very short, de-
pressed, grayish and black. Head with front
convex, micropunctate, pubescence short,
depressed, mouthparts with several erect se-
tae; antennal tubercles moderate, widely di-
vergent; vertex impressed between eyes,
convex behind; eyes coarsely faceted, upper
lobes separated by more than width of lobes,
lower lobes almost twice as long as genae;
antennae rather short, extending about four
segments beyond elytra, scape barely at-
taining middle of pronotum, third segment
longer than first, fourth equal to first, seg-
ments dark mottled, dark annulate at api-
ces, pubescence very fine, depressed. Prono-
tum almost twice as broad as long, sides
with broadly rounded calluses behind mid-
dle; disk with three small calluses, one me-
dian and one on each side near apical mar-
gin; punctures fine, scattered, linear on basal
impression; basal transverse impression
broad, extending onto sides; pubescence fine,
appressed, variegated gray and brownish,
sides at base with two long erect setae; pro-
sternum with intercoxal process plane,
185
slightly narrower than coxal cavities; meso-
sternum with intercoxal process almost
plane, as broad as coxal cavities, broader
than prosternal process; metasternum finely
gray pubescent, pubescence interrupted by
small spots. Elytra about 1.75 times longer
than basal width, sides slightly expanding
behind middle then tapering at about apical
one-fourth; basal calluses low, not tufted,
basal impression rather small, extending
down outside and below calluses; costae
vague; punctures moderately coarse basally,
becoming finer and sparser toward apex; pu-
bescence fine, depressed, middle with a
broad, irregularly margined, oblique, whit-
ish vitta which extends back from suture to
lateral margins, sides behind humeri with
black vittae, an irregular dark vitta present
behind oblique whitish vitta, surface with a
few small, black spots particularly along su-
ture; apices narrowly, obliquely truncate.
Legs short; femora clavate: pubescence fine,
appressed, mottled grayish and brownish;
metatibiae with an external sinus. Abdo-
men finely punctate and pubescent: fifth
sternite slightly longer than fourth, subtrun-
cate at apex. Length, 5.5 mm.
Female.— Form similar. Antennae slight-
ly shorter. Abdomen with fifth sternite much
longer than fourth, narrowly, shallowly
emarginate at apex. Length, 6 mm.
Holotype ¢ from Twin Cays, emerged
from red mangrove 14 July 1986 (C. Feller)
(twigs collected 26 May 1986). One 2 para-
type, same data, emerged | July 1986.
The whitish vitta of the elytra is more
pronounced in the male. In the female this
band does not extend to the margins and a
few narrow pale vittae extend forward and
back on the disk. The suture is also narrowly
pale.
The previously described species of Sty-
loleptus are West Indian with two extending
into the United States. This species is the
first of this genus from the Central Ameri-
can mainland area. The possibility does ex-
ist, of course, that the placement of this
species into Sty/oleptus is incorrect. Such
186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Styloleptus rhizophorae Chemsak & Feller, male.
VOLUME 90, NUMBER 2 187
——a
Fig. 5. Urgleptes ozophagus Chemsak & Feller, female.
188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
problems must be resolved by a generic re-
vision of the very large and taxonomically
difficult tribe Acanthocini.
Urgleptes ozophagus Chemsak & Feller,
New Species (Acanthocini)
Fig. 5
Female.—Form small, depressed; integ-
ument pale reddish brown, antennae yel-
lowish brown and dark, legs yellow brown,
tarsi and most of tibiae dark, elytra dark
vittate basally at sides and with narrow,
connected, dark vittae on apical one-half;
pubescence very fine, appressed, grayish and
dark brown. Head with front slightly con-
vex, broad, micropunctate, pubescence
short, pale, appressed, margin of frons with
several long, suberect setae near sides; ver-
tex convex, medially impressed before inner
eye margins; eyes with lower lobes rounded;
genae slightly shorter than lower eye lobes;
antennae extending about five segments be-
yond elytra, scape shorter than third seg-
ment, fourth longer than third, scape broad-
ly dark annulate, segments very finely
pubescent, third segment with several short,
subdepressed, dark setae beneath. Prono-
tum broader than long; disk lightly convex,
densely micropunctate, basal impression
with a row of fine, deep punctures; pubes-
cence pale, fine, appressed; prosternum with
intercoxal process narrow; mesosternum
with intercoxal process narrow; metaster-
num densely clothed with pale, appressed
pubescence. Elytra about twice as long as
broad, sides tapering behind middle; disk
feebly impressed at basal one-fourth; punc-
tures moderately coarse, dense, becoming
obsolete near apex; pubescence short, fine,
grayish, appressed, sides with broad dark
vittae extending from base to about middle,
apical one-half with dark, narrow reticulate-
like vittae; apices narrowly obliquely trun-
cate. Legs finely pubescent; femora pale with
infuscated patches near apices; tibiae pale
basally, dark over apical two-thirds; tarsi
dark. Abdomen pale, finely densely pubes-
cent; fifth sternite twice as long as fourth,
apex narrowly, shallowly emarginate.
Length, 5.5 mm.
Holotype 2 from Twin Cays, emerged 15
July 1986, from twig terminal of red man-
grove (C. Feller). Two 2 paratypes, Twin
Cays, N. shore of W. Island, 20 January
1987, on R. mangle (W. Mathis) and
Weather Station, 21 January 1987, ex R.
mangle.
The small size and reticulate-like vittae
on the apical half of the elytra make this
species distinctive.
ACKNOWLEDGMENTS
This is contribution no. 196, Caribbean
Coral Reef Ecosystems Program (CCRE),
Smithsonian Institution, partially funded by
a grant from Exxon. Candy Feller prepared
the illustrations.
LITERATURE CITED
Chemsak, J. A. 1983. A new Central American species
of Derancistrus (Coleoptera: Cerambycidae). Proc.
Entomol. Soc. Wash. 85: 714-716.
Chemsak, J. A. and E. G. Linsley. 1982. Checklist of
the Cerambycidae and Disteniidae of North
America, Central America, and the West Indies
(Coleoptera). Plexus. Medford, NJ. 138 pp.
Janzen, D. H. 1986. The Eternal external threat. /n
M. E. Soule, ed., Conservation Biology. Sinauer
Publ., Mass. 584 p.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 189-192
CEROPTERA LONGICAUDA, A SECOND NORTH AMERICAN
SPECIES IN THE KLEPTOPARASITIC GENUS CEROPTERA
MACQUART (DIPTERA: SPHAEROCERIDAE)
S. A. MARSHALL AND D. J. S. MONTAGNES
Department of Environmental Biology, University of Guelph, Guelph, Ontario NIG
2W 1, Canada.
Abstract. — Ceroptera longicauda, new species, is described from specimens collected in
Levy and Leon Counties, Florida. It is compared to C. sivinskii Marshall, and new dis-
tributional data for the latter species is presented.
Key Words:
Ceroptera is widespread in the warmer
parts of the Old World, with 16 African
species, 8 Palaearctic species, and a single
species described from Ceylon. This genus
was first recorded from the Nearctic region
with the description of Ceroptera sivinskii
Marshall 1983. All species of Ceroptera for
which biological information is available are
associated with dung-rolling scarab beetles.
Adult Ceroptera have been observed fre-
quently clinging to dung beetles and were
first observed to oviposit in scarab dung
balls by Roubaud (1916). Sivinski (1983)
demonstrated that Ceroptera sivinskii is
phoretic and kleptoparasitizes dung caches
of scarab beetles in north Florida. Ceroptera
sivinskii has been observed in association
with Phanaeus vindex MacLeay, Geotrupes
egeriei Germar, Copris minutus (Drury), and
Canthon pilularius (L.). It has also been
reared from the dung cache of Copris sp.
(Sivinski 1983). Ceroptera longicauda, new
species, was observed to cling to the elytra
of the scarab Mycotrupes gaigei Olson and
Hubbell. Ceroptera longicauda is similar to
other Ceroptera in possessing strikingly
small eyes, a long narrow interfrontal area,
2 long rows of orbital setulae, a small apical
spur on the hind tibia, a retractile female
abdomen with tubercle-based setae in the
Sphaeroceridae, Diptera, taxonomy, Kleptoparasite
membrane, and a deeply bilobed surstylus.
Similarities shared between the two Nearc-
tic species include the apparently synapo-
morphic, strongly modified male sternite 5
and several probably plesiomorphic char-
acter states such as the incompletely cleft
surstylus and relatively well sclerotized ab-
domen. Furthermore, the legs of C. /ongi-
cauda are not as strongly modified (for
grasping the host) as those of the Old World
species. Ceroptera longicauda can be differ-
entiated from C. sivinskii by the following
key:
1. Lunule broadly rounded anteriorly. Wing mem-
brane brown tinted. Two pairs of dorsocentral
bristles. Epandrium posterodorsally elongated
and tapered, covered with microtubercles (Fig.
3). Anterior surstylar lobe with a large outer lobe
(Figs. 1, 2). Female cerci less than twice as long
as wide (Fig. 9) C. longicauda
— Lunule angulate anteriorly. Wing membrane
clear. One pair of dorsocentral bristles. Epan-
drium simple. Anterior surstylar lobe with a
straight outer surface. Female cerci more than
twice as long as wide C. sivinsku
Ceroptera longicauda Marshall
Figs. 1-11
Description.— Body length 2.1-2.7 mm.
Color dark brown; tarsi, apices of tibiae,
gena, frons and face paler. Interfrontal area
190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-8.
minalia, left lateral. 4, Sternite 5. 5, Terminalia, posterior. 6, Aedeagus, aedeagal apodeme and paramere, left
lateral. 7, Left tibia, dorsal. 8, Left wing. Scales for Figs. 1-2 as on Fig. 2; for Figs. 3-6 as on Fig. 6. Scale bar
for Fig. 8 = 0.5 mm; other scale bars = 0.1 mm.
Ceroptera longicauda, male. 1, Left surstylus, posteroventral. 2, Left surstylus, lateral. 3, Ter-
VOLUME 90, NUMBER 2
191
Figs. 9-11.
Scale bars = 0.1 mm.
long and narrow, 0.2 as wide as frons,
0.4 x as wide as high; bordered by 5S—6 equal
interfrontal bristles. Orbital setulae forming
2 rows; only outer, exclinate row well de-
veloped and extending below eye. Face
broadly carinate, lunule triangular with a
broadly rounded apex. Flagellomere | small,
1.5 times as broad as long; arista arising
dorsobasally, 2.1 x length of rest of anten-
nae. Eye small, 0.8 x genal height. Thorax
with 2 pairs of dorsocentral bristles, anterior
pair slightly longer than acrostichal setulae,
posterior pair subequal in length to broad
scutellum. Acrostichal setulae in 7-8 rows
between dorsocentral areas; prescutellar pair
not enlarged. Katepisternum with a large
posterodorsal bristle reaching 0.8 dis-
Ceroptera longicauda, female. 9, Terminalia, dorsal. 10, Terminalia, ventral. 11, Spermathecae.
tance to wing base, | or 2 small anterodorsal
setulae, and several ventral setulae. Legs
long, all tarsi elongate and curved, distinctly
longer than tibiae, with enlarged pulvulli
and claws. Mid tibia with 2 anterodorsal
and | posterodorsal bristle on proximal half;
1 anterodorsal, | dorsal and | posterodorsal
bristle just below middle; | very long dorsal
bristle in apical quarter; apical part of mid
tibia with large anterior and smaller pos-
terior preapical bristles and with a long api-
coventral bristle; ventral surface of mid tib-
ia with | long bristle near middle. Hind tibia
with a weak apicoventral spur and 2 weak
distal dorsal bristles, each shorter than tibial
width. Hind tarsomeres with short, stout
apicoventral bristles. Wing membrane dis-
192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tinctly brown tinted, second costal sector
1.1 x as long as third in male, 1.2 in fe-
male (Fig. 8).
Male terminalia: Sternite 5 elongate,
posteromedial margin bilobed, setulose,
projecting over genital pouch (Fig. 4). Epan-
drium narrowed and elongated posterodor-
sally; posterodorsal surface covered with
microtubercles (Figs. 3, 5). Cerci bulbous,
distinctly differentiated from epandrium.
Hypandrium small short, simple (Fig. 3).
Surstyli deeply cleft; anterior lobe with a
large lateral process; posterior lobe more
slender, with a stout apical spine (Figs. 1,
2). Distiphallus elongate, well sclerotized,
with a prominent distal ventral medial
sclerite. Basiphallus short, simple; ejac-
ulatory apodeme apparently absent. Para-
mere almost parallel sided, weakly bent an-
teriorly near apex (Fig. 6).
Female terminalia: Abdomen strongly
telescoping; membrane with short, scat-
tered, tubercle-based setae. Tergite 8 with
tripartite pigmentation, median part bare
(Fig. 9). Epiproct pale medially and poste-
riorly, with 2 small setae. Hypoproct broad,
setulose posteriorly (Fig. 10). Cerci short,
less than twice as long as wide, with long
apical bristles. Spermathecae ovate, bent,
swollen apically, with a small invagination
at apex; ducts short, inserted eccentrically
(Fig. 11).
Types.— Holotype ¢, Florida, Levy Co.,
west of Archer, 15.11.1987, ex. Mycotrupes
gaigei, Paul Skelley (BRI). Paratypes: Flor-
ida, 26 4, 43 2, same data as holotype; 2 4,
5 2same data except 1.111.1987, (GUE, FSC,
BRI); Levy Co., 17.11.1976, pitfall trap, L.
R. Davis (3 2, FSC); Leon Co., Tall Timbers
Research Station, 8-15.x.1969, pitfall, D.
L. Harris (2 8, 2 2, FSC).
Ceroptera sivinskii Marshall
Ceroptera sivinskii Marshall 1983: 139.
Material made available since 1983 has
extended the known distribution and al-
lowed two amendments to the original de-
scription. The size range is modified to 1.4—
2.5 mm, with the majority of specimens less
than 2.0 mm (outside the range of the larger
C. longicauda). The dorsal part of the epi-
proct, transparent and apparently membra-
nous on the type material, is lightly scler-
otized and weakly pigmented at least
posteriorly on most other specimens, as in
C. longicauda.
New records (75 specimens): Florida, Lib-
erty Co., 10 mi. SW. Juniper, Rt. 12,
26.11.1983, pig dung among Turkey Oaks,
Woodruff and Thomas (FSC); Marion Co.,
Ocala National Forest, Rd. 65, 1.5 mi. W.
State Rd. 19, 15-16.i1.1984, dung trap, R.
Woodruff (FSC); Alachua Co., Gainesville,
Hogtown Creek, 12.x.1976, P. M. Choat &
R. E. Woodruff (FSC); Okaloosa Co., | mi.
N. Holt, Blackwater River Nat. For.,
23.x.1978, L. Stange, human dung trap
among Turkey Oaks (FSC). Alabama, Cov-
ington Co., 1.7 mi. E. jet. Rt. 84 & Rt. 55,
pig dung trap, 4—-10.111.1977, Woodruff and
Wiley (FSC). Massachusetts, Nonamesset
Id., vi.24.23, sheep dung (1 4, A. H. Stur-
tevant Collection, USNM).
ACKNOWLEDGMENTS
We thank NSERC for financial support
and T. Wheeler for helpful comments. P.
Skelley and R. E. Woodruff of the Florida
State Collection of Arthropods, Gainesville
(FSC) made much of this material available
to us. Specimens not deposited at FSC are
at the University of Guelph (GUE), the Bio-
systematics Research Centre in Ottawa, On-
tario (BRI) or the United States National
Museum, Washington (USNM).
LITERATURE CITED
Marshall, S. A. 1983. Ceroptera sivinskii, anew species
of Sphaeroceridae (Diptera) in a genus new to North
America, associated with scarab beetles in south-
eastern United States. Proc. Entomol. Soc. Wash.
85: 139-143.
Roubaud, E. 1916. Nouvelles observations de pho-
resie chez les Dipteres du groupe des Borboridae.
Bull. Soc. Zool. Paris 41: 43-45.
Sivinski, J. 1983. The natural history of a phoretic
sphaerocerid Diptera fauna. Ecol. Entomol. 8: 419-
426.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 193-195
A REVIEW OF THE SPECIES OF ACRITISPA UHMANN
(COLEOPTERA: CHRYSOMELIDAE, HISPINAE)
C. L. STAINES, JR.
3302 Decker Place, Edgewater, Maryland 21037.
Abstract. —The literature on Acritispa is reviewed. Octotoma germaini Pic is transferred
to Acritispa. The two included species are redescribed and a key is presented; these species
occur in Central and South America.
Key Words:
The species of the genus Acritispa were
examined in connection with ongoing re-
visionary studies on the subfamily Hispinae
(Staines 1986a, b, 1987). The name dAc-
ritispa was first published by Uhmann
(1937); in this genus he placed the species
Physocoryna dilatata Uhmann. Under Ar-
ticle 13b (Internat. Code Zool. Nomen.
1985) the generic name was a nomem nu-
dem because no type species was designat-
ed. Uhmann (1940) designated Acritispa di-
latata (Uhmann) as the type species of the
genus validating the name. Papp (1953) list-
ed five species in Acritispa: dilatata, nigri-
tarsis (Weise), triquetra (Uhmann), viridi-
ceps (Pic), and viridinotata (Pic). Uhmann
(1957) placed all but di/atata in the genus
Probaenia Weise.
Acritispa is in the tribe Uroplatini (Weise
1911) which is characterized by having the
last antennal segments very closely united
and the antennae appearing as three to eight
segmented. The tribe is represented by thir-
ty-two genera. Acritispa is distinguished by
the following combination of characters:
antennae short, not exceeding the base of
the pronotum; 8-segmented, segments I and
II subequal, VII as wide as VIII and difficult
to distinguish as a separate segment; VIII
not longer than the four preceeding seg-
ments combined. Pronotum with two
Coleoptera, Chrysomelidae, Hispinae, Acritispa
toothlike projections on lateral margins.
Elytra expanded to apex; plications over en-
tire surface.
Acritispa Uhmann
Acritispa Uhmann 1940: 143 (type species:
A. dilatata (Uhmann)). Uhmann 1937:
336, Blackwelder 1946: 729, Uhmann
1950: 267, Papp 1953: 93, Uhmann 1957:
115, 1964: 11, Gaedike & Dobler 1971:
347, Seeno & Wilcox 1982: 161.
General habitus: Size small (4.0—5.8 mm),
with elytra greatly expanded at apex. Head:
Vertex micropunctate; median sulcus pres-
ent; antennae 8-segmented, I-II subequal,
III widened apically, VII as wide as VIII
and difficult to distinguish as a separate seg-
ment, VIII clavate, hirsute. Pronotum:
Wider than long; tooth present on anterior
margin on each side of head; two toothlike
projections on lateral margins; covered with
coarse punctures. Elytra: Greatly expanded
at apex; translucent at exterior apical angles;
plications over entire surface; lateral mar-
gins flattened, dentate; apical margins flat-
tened, dentate. Profemur expanded apical-
ly.
Measurements were taken with an ocular
micrometer. Pronotal length and width were
taken along the midlines. Elytral width was
measured at the humeri. Elytral length was
194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
measured from the base to apex. Total length
was measured from the base of the frontal
sulcus to the apex of the elytra.
Key TO THE DESCRIBED SPECIES OF
AcRITISPA UHMANN
1. Vertex of head with median sulcus faint; body
color black ae dilatata (Uhmann)
— Vertex of head with median sulcus deep and
wide; body color reddish-brown .. germaini (Pic)
Acritispa dilatata (Uhmann)
Physocoryna dilatata Uhmann 1932: 266
(type not seen; type locality Brazil, Bahia;
holotype deposited in Deutschen Ento-
mologischen Institutes (Gaedike & Dob-
ler 1971)).
Acritispa dilatata (Uhmann): Uhmann 1937:
336, 1940: 143, 1950: 267, Papp 1953:
93, Uhmann 1957: 115, 1964: 11, Gae-
dike & Dobler 1971: 347.
Body color black. Head: Median sulcus
present, faint; three punctures on inner mar-
gin of each eye; vertex rugose, micropunc-
tate; antennae reddish; segments I-II trans-
verse, I largest; II-VI compressed, smaller
than I or II. Pronotum: Two tubercles pres-
ent on disc behind midline; covered with
coarse punctures; basal margin bisinuate;
surface micropunctate; length 0.7-0.8 mm
(avg. 0.75, n = 4); width 1.1-1.3 (avg. 1.2).
Elytra: Covered with large plicae; lateral
margins dentate, more so at base; rows of
punctures visible between plicae; length 3.1-
3.6 (avg. 3.3); width 1.7-1.9 (avg. 1.75).
Legs: Reddish. Venter: Pro- and metasterna
alutaceous; mesosternum punctate in mid-
dle; abdomen red. Total length: 4.0-4.4 (avg.
4.2).
Discussion: Life history unknown. Im-
mature stages undescribed. This species can
be distinguished from A. germaini by the
less pronounced basal elytral plications, the
faint median sulcus on the vertex of the
head, and the black body color.
Larval host plant: Unknown.
Distribution: Brazil and Paraguay.
Specimens examined: BRAZIL: Lam-
bary, XI/1926 (USNM); Parahyba, [X/1884
(USNM); Sao Paulo (USNM). PARA-
GUAY: central, 1885 (USNM). Total: 4.
Acritispa germaini (Pic),
NEw COMBINATION
Octotoma germaini Pic 1925: | (Holotype:
Cochabamba (Bolivie), Germain/Octo-
toma n. sp./Museum Paris, coll. M. Pic/
Type/germaini n. sp. (MNHN)). Uhmann
1927: 136, Blackwelder 1946: 729, Papp
1953: 93, Uhmann 1957: 116, Descar-
pentries & Villiers 1959: 149.
Body color reddish-brown. Head: Medi-
an sulcus deep, wide; ledge over base of an-
tennae; antennae reddish-brown, segment
I-II punctate; HI widened apically, longer
than I or II, punctate, IV-VI transverse,
punctate, VII wider, fringe of setae on apical
margin, VIII hirsute, pointed apically, three
whorls of setae. Pronotum: Two tubercles
present on disc near midline; areas between
punctures micropunctate; callous present
near right tubercle; length 0.9-1.4 mm. (avg.
1.1; n = 3); width 1.3-1.5 (avg. 1.4). Scu-
tellum: Light reddish-brown; rounded at
apex. Elytra: Apical margin less dentate than
lateral margins; puncture rows visible be-
tween plications; base explanate, expanded
over base of pronotum; plications very large,
especially on basal half, less raised on apex;
length 3.6—-4.4 (avg. 3.9); width 1.9-2.5 (avg.
2.1). Legs: Reddish-brown, except femur
which is black; femur with large punctures
at apex. Total length: 4.3-5.8 (avg. 4.8).
Discussion: Life history unknown. Im-
mature stages undescribed. This species can
be distinguished from 4. dilatata by the
more pronounced basal elytral plications,
the deep median sulcus on the vertex of the
head, and the reddish-brown body color.
Larval host plant: Unknown.
Distribution: Bolivia and Panama.
Specimens examined: PANAMA: Porto
Bello, 11/VIII, 19/III/1911(USNM); Canal
Zone, Fort Kobbe, 20/VI/1976 (EGRC).
BOLIVIA: Cochabamba (MNHN). To-
tal: 4.
VOLUME 90, NUMBER 2
ACKNOWLEDGMENTS
I thank N. Berti, Museum National
d’Histoire Naturelle, Paris (MNHN), for the
loan of the Pic type; R. E. White, Systematic
Entomology Lab, USDA (USNM) for ac-
cess to that collection and E. G. Riley
(EGRC) for the loan of the material from
his collection. R. E. White and L. LeSage,
Biosystematics Research Institute, Agricul-
ture Canada, commented on an earlier draft
of this manuscript.
LITERATURE CITED
Blackwelder, R. E. 1946. Checklist of the Coleopter-
ous insects of Mexico, Central America, the West
Indies, and South America. U.S. Nat. Mus. Bull.
185(4): 551-763.
Descarpentries, A. and A. Villiers. 1959. Les types
de la collection M. Pic. I. Chrysomelinae et His-
pinae du Nouveau-Monde. Bull. Mus. Hist. Nat.
Paris (2)31: 137-154.
Gaedike, R.and H. Dobler. 1971. Katalog der in dem
Sammlungen des ehemaligen Deutschen Ento-
mologischen Institute aufbewahrten Typen-VII
(Coleoptera: Hispinae). Beitr. Entomol. 21: 341-
395.
Papp, C.S. 1953. The Hispinae of America. The 3rd
contribution for promoting the scientific results of
the International Hylean Amazon Institute in
Manaos, Brazil. Port. Acta Biol. (B)4: 1-147.
Pic, M. 1925. Nouveautes diverses. Mel. Ext. Ento-
mol. 44: 1-36.
Seeno, T. N. and J. A. Wilcox. 1982. Leaf beetle
genera (Coleoptera: Chrysomelidae). Entomogra-
phy 1: 1-221.
Staines, C. L. 1986a. New combination and new syn-
195
onymy in North American Stenopodius (Coleop-
tera: Chrysomelidae: Hispinae) with a taxonomic
note on Uroplatini. Proc. Entomol. Soc. Wash. 88:
192"
—. 1986b. A revision of the genus Brachycoryna
(Coleoptera: Chrysomelidae: Hispinae). Insecta
Mundi |: 231-241.
. 1987. The correct generic placement of Sum-
itrosis gracilis (Horn) (Coleoptera: Chrysomelidae:
Hispinae). Coleopts. Bull. 41: 319-321.
Uhmann, E. 1927. Hispinen des Deutch. Ent. Insti-
tutes (Col.). 4. Beitrag zur Kenntnis der Hispinen.
Entomol. Mittlungen 16: 134-136.
1932. Sudamerikanische Hispinen aus der
Sammlung des Stettiner Museums und der mei-
nigen. 38. Beitrag zur Kenntnis der Hispinen (Col.
Chrys.). Stett. Entomol. Zeit. 93: 260-266.
1937. Ubersicht uber die ersten Gattungen
der Uroplatini (Col. Chrys.). 67. Beitrag zur
Kenntnis der Hispinen. Entomol. Blatt. 33: 336-
337.
1940. Die Genotypen der von mir aufges-
tellten Hispinen-Gattungen. 88. Beitrag zur
Kenntnis der Hispinen (Col. Chrys.). Entomol.
Tidskrift 61: 143-144.
1950. Die Deckenskulptur von Octotoma
Suffr. und verwandten Gattungen. 118. Beitrag zur
Kenntnis der Hispinae (Coleopt. Chrysom.). Rev.
Entomol. 21: 259-274.
1957. Coleopterum Catalogus. Chrysomeli-
dae: Hispinae, Hispinae Americanae. W. Junk,
Gravenhage. Pars 35(1): 1-153.
1964. Hispinae aus dem Staate Sao Paulo,
Brasilien. 209. Beitrag zur Kenntnis der Hispinae
(Coleoptera, Chrysomelidae). Iheringia (Zool.) 32:
1-28.
Weise, J. 1911. Coleopterum Catalogus, Chrysomel-
idae: Hispinae. Pars 35: 1-94. W. Junk. Berlin.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 196-200
DIGONOGASTRA: THE CORRECT NAME FOR NEARCTIC
IPHIAULAX OF AUTHORS (HYMENOPTERA, BRACONIDAE)
DONALD L. J. QUICKE
Department of Zoology, University of Shefheld, Shefheld S10 2TN, United Kingdom.
Abstract.—The braconine genus /phiaulax Foerster although frequently taken to include
Nearctic and Neotropical species, is in fact restricted to the Old World. The correct generic
name for most New World /phiaulax of authors 1s Digonogastra Viereck. Digonogastra
and its type species (D. epicus (Cresson)) are redescribed and illustrated. Diagnostic fea-
tures of Digonogastra and Iphiaulax are provided. Monogonogastra Viereck 1s a junior
synonym of Digonogastra Viereck.
Key Words:
Braconinae
The Nearctic fauna of braconine wasps
contains a number of described and many
undescribed species that have traditionally
been treated as species of [phiaulax Foers-
ter. [phiaulax (type-species: Ichneumon im-
postor Scopoli) was originally described from
Europe and 1s distributed throughout the
Palearctic, Afrotropical and Indo-Austra-
lian regions. However, during the course of
a revision of the World genera of Braconi-
nae, it has become apparent that most, if
not all, of the New World ‘/phiaulax’ species
are not congeneric with those from the Old
World despite some superficial resem-
blance. Viereck (1912) erected two new gen-
era, Digonogastra and Monogonogastra, to
receive a number of species of Nearctic
Iphiaulax of authors on the basis of small
differences in metasomal sculpture, but he
still considered that [phiaulax occurred in
North America, and subsequently, Muese-
beck & Walkley (1951) synonymized both
of Viereck’s genera with Iphiaulax. Digon-
ogastra which is a senior synonym of Mon-
ogonogastra, appears to be the oldest avail-
able name for the Nearctic /phiaulax group.
In order to clear up these misunderstand-
Digonogastra, Iphiaulax, Monogonogastra, Nearctic fauna, Braconidae,
ings Digonogastra is redescribed below and
features are given which enable its separa-
tion from /phiaulax Foerster, and from the
other New World genera of Braconinae.
Many species will be reclassified elsewhere
(Quicke, in press a).
Terminology follows that of van Achter-
berg (1979). The type material is located in
the United States National Museum, Wash-
ington (USNM).
Digonogastra Viereck
Figs. 1-9
Digonogastra Viereck, 1912. Type-species:
Bracon epicus Cresson, 1872; monobasic
and original designation.
Monogonogastra Viereck, 1912. Type-
species: Bracon atripectus Ashmead,
1889; monobasic and original designa-
tion.
Females.—Antennae approximately as
long as the forewing. Median flagellomeres
wider than long. Scapus sub-cylindrical,
longer ventrally than dorsally, apicolater-
ally and (weakly) apicomedially emarginate
(Fig. 2). Labiomaxillary complex not elon-
VOLUME 90, NUMBER 2
Figs. 1-9.
scapus, pedicellus, and basal flagellomere, lateral aspect. 3, Head, dorsal aspect. 4, Junction of veins 1-SR-M
and I-SR of right forewing. 5, Distal part of ovipositor. 6-9. Digonogastra selected species. 6, Metasomal tergites
2-4, dorsal aspect. 7, Junction of veins 1-SR+M and 1-SR of right forewing. 8, Ist metasomal tergite dorsal
aspect. 9, Ovipositor. Scale line: Figs. 1-3, 6-7, 1.0 mm; Fig. 4, 0.67 mm; Figs. 5, 9, 1.3 mm; Fig. 8, 0.8 mm.
gate. Lower part of clypeus more or less
strongly reflexed into the hypoclypeal
depression, separated from the upper part
by a carina. Clypeus usually separated from
the face by a weak carina though sometimes
the two are more or less contiguous. Face
usually densely long setose. Eyes virtually
glabrous. Frons usually generally, evenly
impressed with a mid-longitudinal sulcus;
usually extensively densely setose (Fig. 3),
the setae often being more or less prostrate
Features of Digonogastra spp. 1-5. D. epicus 1, Metasomal tergites | & 2, dorsal aspect. 2, Right
and silvery in appearance. Prosoma mod-
erately contracted behind the eyes.
Mesosoma usually extensively setose, es-
pecially the scutellum, mesosternum and
propodeum, smooth and shiny between the
setae. Notauli usually distinctly impressed
along most of the length of the mesoscutum.
Scutellar sulcus usually narrow and distinct-
ly crenulate. Precoxal suture absent. Pleural
suture smooth and almost obliterated. Pro-
podeal spiracle situated at about the middle
198
of the propodeum, approximately 2 x taller
than long.
Forewing: Marginal and 2nd submarginal
cells long. Vein 1-SR+M usually distinctly
angled posteriorly shortly after arising from
1-SR (Fig. 4), but more or less straight in
some species (Fig. 7). Vein cu-a usually in-
terstitial, sometimes marginally postfurcal.
Veins 1-SR and C+SC+R forming an angle
of more than 55° (often 80°). Vein 1-M
straight. Vein 3-CU1 not or hardly expand-
ed posteriorly.
Hindwing: Vein l|r-m at least slightly
shorter than SC+RI. Apex of vein
C+SC+R with more than | especially
thickened bristle (hamule), unless the length
of the forewing is less than 5 mm. At least
with a small glabrous area postero-distal to
vein cu-a.
Claws with small, rounded basal lobes.
Anterolateral aspect of fore tibia densely se-
tose, without an apical transverse row of
thick, peg-like bristles. Hind tibial spurs
densely setose.
Metasoma generally depressed; general
sculpture variable from largely smooth to
rugose or foveate. Ist metasomal tergite with
very well-developed dorso-lateral carinae
and with a raised medial area which is well-
separated from the dorso-lateral carinae.
Median area of Ist metasomal tergite usu-
ally with at least a trace of a mid-longitu-
dinal carina for a short distance on its pos-
terior third, though sometimes this is only
indicated by a pair of sub-medial pits (Fig.
8). 2nd metasomal tergite always with a dis-
tinct mid-basal area which is usually pro-
duced to form a mid-longitudinal carina
(Figs. 1, 6); with a pair of posteriorly di-
verging furrows running from the anterior
corners of the mid-basal area. Posterior
margin of 2nd metasomal tergite moderate-
ly sinuate. 2nd suture variable, smooth or
crenulate. 3rd metasomal tergite with large
antero-lateral areas defined by a pair of pos-
teriorly diverging furrows; often with a dis-
tinct, sometimes large, mid-basal triangular
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
area (Fig. 1). 4th-6th metasomal tergites only
rarely with a distinct transverse, subposte-
rior groove. 4th—6th tergites with a trans-
verse peri-basal groove but this groove not
divided laterally to demark anterolateral
areas. Ovipositor between 0.25 & 3.0 x
length of metasoma; highly variable (Figs.
5, 9), sometimes slender with a pre-apical
dorsal nodus and apico-ventral serrations,
but sometimes thickened without a pre-api-
cal dorsal nodus and with very reduced ap-
ico-ventral serrations.
Males.—Similar to females. Digitus of
genitalia with 4 (or sometimes 3) well-de-
veloped and widely separated, tooth-like
processes dorso-laterally. Basal ring ante-
riorly pointed but not produced into a spine.
Notes on Digonogastra. Digonogastra
species display a remarkable superficial re-
semblance to Jphiaulax species, many also
having evolved a thickened ovipositor
without a dorsal nodus. Digonogastra
species may however, be distinguished from
those of Jphiaulax by a number of features.
Perhaps surprisingly, the most consistent
feature appears to be the presence ofa clear-
ly-defined and often large raised mid-basal
triangular area on the 2nd metasomal tergite
of Digonogastra; such an area is never pres-
ent in /phiaulax. In addition, most Digon-
ogastra species have an extensively, densely
setose frons, the setae often being relatively
long and lying rather flat, and the 3rd to Sth
metasomal tergites lack a transverse sub-
posterior groove (though this is present in
Digonogastra ornatus (Provancher)). In
Iphiaulax the frons is always completely
glabrous except adjacent to the eye, and the
3rd—5th metasomal tergites nearly always
have a transverse, subposterior groove.
However, perhaps the most important fea-
ture separating Digonogastra from Iphiau-
lax is the presence of four (occasionally
three) well developed tooth-like processes
on the digitus of the male genitalia in Di-
gonogastra whereas in Iphiaulax there 1s
only one. The presence of four digital teeth
VOLUME 90, NUMBER 2
has recently been shown to characterize a
group of apparently closely related genera
from both the Old and New worlds, (Quicke,
in press b) typified by Afrotropical genera
Archibracon Saussure and Sororarchibra-
con Quicke. Of the Neotropical genera, four
digital teeth are also present in Megabracon
Szepligeti and Lasiophorus Haliday, and this
may indicate that Digonogastra 1s derived
from this Neotropical assemblage of bra-
conine genera.
For practical purposes, Digonogastra can
be separated from /phiaulax by the presence
in the former of a medium sized to large
mid-—basal triangular area on the 2nd meta-
somal tergite. In the recent key to the Old
World genera of Braconinae provided by
Quicke (1987), Digonogastra spp. with a pre-
apically smooth ovipositor will key to cou-
plet 95 and some will run out to Braco-
morpha Papp at couplet 96. Digonogastra
spp. with a nodus on the ovipositor will run
to couplet 125 and most will run (with some
difficulty) to Poecilobracon Cameron.
Digonogastra epicus (Cresson)
Figs. 1-5
Bracon epicus Cresson, 1872.
Material examined. — Female holotype in
USNMW: “Texas Belfrage” & “Type No.
1611 U.S.N.M.” One female in the author’s
collection: **Merivale. Ont. 5.viil. 1980 J. J.
de Gryse”’; one female in USNM: “Dawson
Camp, Salt River Ariz” & ““CHT Townsend
coll. sep 4” both compared with the holo-
type.
Females. — Length of body 10-12 mm, of
forewing 11-13mm and of ovipositor
(exserted part) 7.0-7.5 mm.
Antennae with 51 flagellomeres. Penul-
timate flagellomere 1.3 = longer than wide.
Ist flagellomere 1.6 and 1.8 x longer than
the 2nd and 3rd respectively, the latter being
1.1 longer than wide. Scapus almost cy-
lindrical, approximately 2.2 = longer than
maximally deep (Fig. 2). Hypoclypeal hair-
199
brushes well developed. Upper part of clyp-
eus finely punctate. Clypeus clearly de-
marked from face by elevation and by a
finely crenulate groove. Height of clypeus:
intertentorial distance: tentorio-ocular dis-
tance = 10:29:18. Face densely silvery se-
tose and punctate, smooth and shiny be-
tween the punctures. Width of face : width
of head : height of eye = 29:59:27. Lateral
half of frons on either side moderately
densely covered with rather prostrate sil-
very setae. Distance between posterior ocel-
li: diameter of posterior ocellus : shortest
distance between posterior ocellus and eye =
10:9:23. Head rather strongly contracted
behind the eyes. Occiput sparsely setose.
Mesosoma 1.6 x longer than high. Prono-
tum largely gabrous laterally; lateral pro-
notal groove only (weakly) crenulate at the
front of the pronotum. Propleuron moder-
ately densely long setose. Middle lobe of
mesoscutum rather strongly protruding in
front of the lateral lobes. Mesoscutum large-
ly glabrous except along line of notaull. Scu-
tellar sulcus crenulate. Scutellum rather
sparsely setose. Mesopleuron sparsely se-
tose posteriorly. Mesosternum moderately
densely setose. Median area of metanotum
glabrous. Propodeum and metapeuron ex-
tensively, densely long setose.
Forewing.—Lengths of SR1:3-SR:r =
80:51:10. Vein 1-SR+M distinctly angled
posteriorly shortly after arising from 1-SR.
Veins C+SC+R and 1-SR forming an angle
of approximately 60°.
Hindwing.—Lengths of veins Ir-m:
SC+RI1 = 19:27. Apex of vein C+SC+R
with 2 to 3 thickened bristles (hamules).
Postero-basal part of wing with a moder-
ately large glabrous area.
Length of fore femur : tibia: tarsus = 47:
57:76. Fore basitarsus approximately 5 x
longer than maximally deep.
Metasoma largely smooth and shiny. El-
evated median area of Ist metasomal tergite
largely smooth, the mid-longitudinal carina
at most only indicated by a pair of small
200
weak submedial depressions, often absent;
bordered antero-laterally by a few rugae. 2nd
metasomal tergite approximately 1.9 x wid-
er than maximally long; with a clearly-de-
fined though rather small, raised mid-basal
area which is produced into a mid-longi-
tudinal carina posteriorly. 2nd metasomal
suture and transverse peribasal grooves of
the 4th to 6th tergites smooth. Tergites 3 to
6 sparsely setose. Ovipositor (part extend-
ing beyond the apex of the metasoma) ap-
proximately 0.7 length of forewing; with
a pre-apical dorsal nodus and apico-ventral
serrations.
Antennae, head, mesosoma, legs and ovi-
positor sheaths black; metasoma bright red;
wings pale brown with dark brown vena-
tion.
ACKNOWLEDGMENTS
I am indebted to Paul M. Marsh (USNM)
for loan of the type specimen of Digono-
gastra epicus and of other specimens rele-
vant to this study.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LITERATURE CITED
Cresson, E. T. 1872. Hymenoptera Texana. Trans.
Am. Entomol. Soc. 4: 153—292.
Muesebeck, C. F. W. and Walkley, L. M. 1951. Bra-
conidae, pp. 90-184 (vol. 2). Jn C. F. W. Muese-
beck, K. V. Krombein and H. K. Townes, eds.,
Hymenoptera of America North of Mexico. Syn-
optic Catalogue U.S. Dept. Agric., Agric. Mono. 14—
20 pp.
Quicke, D. L. J. 1987. The Old World genera of
Braconine wasps (Hymenoptera: Braconidae). J.
Nat. Hist. 21: 43-157.
In press a. Reclassification of some Neo-
tropical species of Braconinae (Hym., Braconi-
dae). Entomol. Mon. Mag.
. Inpress b. Inter-generic variation in the male
genitalia of the Braconinae (Insecta, Hymenop-
tera, Braconidae). Zool. Scripta.
van Acterberg, C. 1979. A revision of the subfamily
Zelinae auct. (Hymenoptera, Braconidae). Tijdschr.
Entomol. 122: 241-479.
Viereck, H. L. 1912. Contributions to our knowledge
of bees and ichneumon-flies, including descrip-
tions of twenty-one new genera and fifty-seven
new species of ichneumon-flies. Proc. U.S. Nat.
Mus. 42: 613-648.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 201-203
EVIDENCE FOR MULTIVOLTINISM IN PRODIPLOSIS PLATANI
GAGNE (DIPTERA: CECIDOMYIIDAE), A LEAF CURL
MIDGE OF AMERICAN SYCAMORE
JOHN W. NEAL, JR. AND K. M. Gott
Florist and Nursery Crops Laboratory, Building 470, Horticultural Science Institute,
Agriculture Research Service, United States Department of Agriculture, Beltsville, Mary-
land 20705.
Abstract. —Seasonally observed pulses of leaf curls caused by larval feeding suggest a
sycamore leaf curl midge, Prodiplosis platani Gagné is multivoltine with five to six gen-
erations per year. Larvae feed on the abaxial lateral surface of new or expanding leaves
and cause the margins to curl tightly, disfiguring the leaf. Frequency of larval activity was
measured by periodically counting new leaf curls. In 1983 and 1984, from early June to
early October in Maryland, five to six peaks occurred at 16-17 day intervals. Larvae are
actively sought as food by adults of many species of coccinellids and the curls serve as
retreats for spiders. Pteromalid and eulophid parasites were recovered from the pupae.
Key Words: | Sycamore, Prodipolis platani, generations, parasites
The recently described leaf curl midge
Prodiplosis platani Gagné, (Gagné 1986) is
a potential pest of American sycamore sap-
lings (Platanus occidentalis L.) in eastern
North American nurseries. Larvae feed on
the surface of abaxial lateral margins of
young leaves. These become disfigured as
feeding causes the leaf margins to curl tight-
ly during growth. The literature of the nine
known species of Prodiplosis in N. America
includes reports on the biology of P. citrulli
(Felt) (Wehrle 1946), P. morrisi Gagné
(Morris 1981), P. vaccinii (Felt) (Driggers
1926) and P. violicola (Coquillett) (Garman
1922). Each of these reported species has
been described as a pest causing economic
damage. There are no previous reports on
the biology of P. platani. Gagné (1986) not-
ed that P. platani was first collected in New
York [= Cecidomyia sp., Felt (1940)] and
later from New Jersey, Pennsylvania, and
Maryland. Larval feeding damage by P. pla-
tani was first observed on young American
sycamores in research plots at the United
States Department of Agriculture, Agricul-
tural Research Center, Beltsville, Maryland
throughout the summer of 1982 and has
since been observed on sycamores in Prince
George’s and Howard counties, Maryland.
The observed chronic appearance of leaf
curls during the growing season prompted
our study to determine the frequency of oc-
currence that would provide data on sea-
sonal activity that could be quantified. Lim-
ited observations on the cocoon and
predators and parasites are provided.
MATERIALS AND METHODS
In this study six sycamores at the Agri-
cultural Research Center that had been
propagated four years earlier by rooted cut-
tings from a single sycamore were used. The
trees had been spring-pruned in the early
growth stage to promote sprouting and each
plant consisted of several long shoots not
exceeding 10 feet in height. Sycamores have
of Curled Leaves per Tree
Mean No.
om 270 1
June
Fig. 1.
under peaks represent larval pulses.
indeterminant growth (oaks, Quercus have
determinate growth and produce two flush-
es of annual growth) during the growing sea-
son and will continue to produce new ter-
minal growth and new leaves as a result of
available water. During the first year of study
(1983) the sycamores were watered on July
7 and August 22. A repeat study was con-
ducted in 1984 because the first year data
suggested several generations. During 1984
Mean number of new leaf curls per America
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
—e 1983
e---- 1984
>
wr
20 1
n sycamore tree in 1983 and 1984. Coincident numbers
trees were watered frequently to promote
terminal growth and thus increase the num-
ber of new leaves and the potential rate of
infestation. The relatively large number of
new growth leaves would amplify the larval
activity pulses that would occur. The six
plants were observed three times a week in
1983 and twice a week in 1984. All leaves
on each plant were examined independently
by two observers. Each leaf with surface
VOLUME 90, NUMBER 2
wrinkling or an incipient curl was marked
on the adaxial surface to indicate that 1t had
been observed and recorded.
RESULTS AND DISCUSSION
Continuous new leaf production by the
sycamores was obtained during the 1984
growing season as a result of frequent water-
ing. The higher number of new leaves re-
sulted in a corresponding higher incidence
of infected leaves than in 1983. The mean
number of observed new curled leaves by
tree per date is presented in Fig. 1. In 1983
a total of 737 curled leaves (X = 122.8/tree)
was counted and in 1984 2216 (x = 369.3/
tree) were counted. In 1983 there were five
pulses of larval activity. Pulses one and two
at 16 and 17 days apart were recorded on
the relatively young growth during June and
July, three subsequent minor increases of
leaf curls with similar temporal spacing fol-
lowed. In 1984 six pulses of larval activity
were observed with peaks |, 2 and 3 coin-
cident with the first three peaks of 1983.
The first pulse, a result of spring-emerged
adults, was minor. Activity pulses 2, 3 and
4 were each followed by minor emergences
and may result from the sampling intervals.
Pulses 5 and 6 in September were minor
which may have resulted due to the normal
seasonal decline in tree growth. Larval ac-
tivity for both years commenced in early to
mid June and ended 1n early October. These
findings are similar to those seasonal studies
of P. morrisi, found from June to August
on poplar (Populus deltoides Bartram) and
hybrid poplars (Morris 1981); Morris (1981)
reported five generations with an average
development period per generation of 16
days based on adult emergences. Larval
pulses by P. platani suggest a corresponding
development time. Our observations sug-
gest that the unsightly feeding damage is
accumulative and results in a general am-
biguity of the existing 5 to 6 generations
203
without frequent sampling. Because several
generations occur on a season’s foliage and
the damaged leaves do not drop, the cu-
mulative damage can be aesthetically det-
rimental.
Midge larvae construct white, double
convex silken cocoons in the leaf curl. These
are formed near the leaf surface or among
clots of stellate hairs that form as a result
of larval feeding and movement on the sur-
face. Many species of adult coccinellids were
observed searching for larvae and pupae in
the curl. The curls also provide retreat for
hunting spiders such as Salticids. Parasites
included Pteromalidae (Zatropis sp.) and
Eulophidae (Tetrastichus sp.) that emerged
in the laboratory from pupae collected June
15, 1983. Data from this study indicate that
P. platani is multivoltine and have estab-
lished it as a potential pest on rapid growing
sycamores.
ACKNOWLEDGMENTS
We thank E. E. Grissell Agriculture Re-
search Service, SEL for identification of pu-
pal parasites and Laurie J. Davis, Efhe Haw-
ley, Donald C. Johnson and Carol Jo Osiecki
of this laboratory for their attentiveness in
making leaf counts.
LITERATURE CITED
Driggers, B. F. 1926. The blueberry tp worm (Con-
tarinia vaccinii Felt), a new species of midge at-
tacking cultivated blueberries. J. N.Y. Entomol.
Soc. 34: 82-85.
Felt, E. P. 1940. Plant galls and gall makers. Com-
stock, Ithaca, N.Y.
Gagné, R. J. 1986. Revision of Prodiplosis (Diptera:
Cecidomyiidae) with descriptions of three new
species. Ann. Entomol. Soc. Am. 79: 235-245.
Garman, P. 1922. The violet gall midge. Phytophaga
violicola (Coquillett). Conn. Agric. Exp. Stn. New
Haven Bull. 234: 152-156.
Morris, R.C. 1981. Prodiplosis morrist: Gagné a new
pest of poplars (Diptera-Cecidomytidae). Ont. Min.
Nat. Res. Pest Control Rep. No. 13: I-7.
Wehrle, L. P. 1946. The cucurbit midge, /tonida ci-
trulli J. Econ. Entomol. 39: 415-416.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 204-206
NOTES ON THE BIOLOGY OF CAENOCEPHUS ALDRICHI BRADLEY
(HYMENOPTERA: CEPHIDAE)
PAUL E. HANSON AND JEFFREY C. MILLER
Department of Entomology, Oregon State University, Corvallis, Oregon 97331-2907
(PEH present address: Escuela de Biologia, Universitad de Costa Rica, Ciudad Univer-
sitaria, Costa Rica).
Abstract.—The larva of Caenocephus aldrichi Bradley was found boring in stems of
Holodiscus discolor (Pursh) (Rosaceae). This represents the first host plant record for the
genus. The rarity of C. a/drichi is demonstrated by the finding of 12 infested stems in 30
hours of searching patches of H. discolor. In Oregon, C. aldrichi appears to be univoltine,
with adults present from early May to early July. Observed mortality factors include
parasitism by Preromalus sp. (Hymenoptera: Pteromalidae) and possible predation by
birds.
Key Words:
The host plants of stem sawflies (Hyme-
noptera: Cephidae) are fairly well known at
the generic level. In North America there
are six genera in the Cephidae, all in the
Cephinae. Species with known hosts in the
Cephini (Calameuta, 1 sp.; Cephus, 2 spp.:
and Trachelus, 1 sp.) are stem borers in
grasses as larvae. Species with known hosts
in the Hartigiini (Caenocephus, | sp.; Har-
tigia, 6 spp.; and Janus, 4 spp.) may bore
in the stems of various angiosperms (Smith
1979, 1986). Larvae of Hartigia species ap-
pear to be restricted to the Rosaceae (roses
and blackberries) while species of Janus uti-
lize various woody dicots (Hanson 1986,
Middlekauff 1969). However, the host plant
of Caenocephus aldrichi Bradley was here-
tofore unknown. The only other species in
this genus is the Palearctic C. /unulatus
(Strobl) and its host plant is also unknown
(Schedl 1981).
Limited data are available on the biology
of adults. The species has been recorded
from British Columbia, Washington, Idaho,
Oregon, and California (Bradley 1905, Mid-
Caenocephus, Cephidae, Holodiscus, host plant.
dlekauff 1952, 1969), but specimens are very
rare in collections (D. R. Smith, pers.
comm.). The purpose of this paper 1s to re-
port the discovery of the host plant of C.
aldrichi and present brief notes on its bi-
ology.
MATERIALS AND METHODS
A search for the larval host plant was be-
gun by limiting field observations to species
of Rosaceae, Fagaceae, and Salicaceae where
geographic distributions were within the re-
corded distribution of adult C. a/drichi. Be-
cause species in the closely related genus
Hartigia are confined to Rosaceae, special
attention was devoted to members of this
family.
Field sampling was conducted by looking
for severed stems for a set period of time
that varied depending on plant density.
Typically, several thousand stems were ob-
served in an hour of searching. Stems with
severed tips were clipped and set aside until
the sampling period was over. The severed
stems were then partially dissected to verify
VOLUME 90, NUMBER 2
the presence of an insect larva. The “‘oc-
cupied” stems were placed individually in
glass tubes (20 cm x 2.5 cm) plugged with
cotton. Tubes containing infested stems were
kept outside for three months (November-
January) and were then brought into the
laboratory (22 + 2°C) for observation of
adult emergence.
When cephid larvae were discovered in
stems of Holodiscus discolor (Pursh), addi-
tional field observations and collections were
concentrated on this species. Infested stems
were collected July—October of 1986 from
the following localities in Oregon: Benton
County (Corvallis and Mary’s Peak); Curry
County (15 miles east of Port Orford); and
Wasco County (13-16 miles west of Dufur).
Adult stem sawflies, a larva, and parasit-
oids obtained from this study were depos-
ited in the Systematic Entomology Labo-
ratory, Department of Entomology, Oregon
State University.
RESULTS AND DISCUSSION
Adult cephids reared from stems of H.
discolor, ocean spray, were identified as C.
aldrichi using keys published by Middle-
kauff (1969) and comparison with speci-
mens identified by D. R. Smith in the Or-
egon State University insect collection.
The number of infested shoots was ex-
tremely low at all sites where C. aldrichi
were found. Several stands of H. discolor
were searched for a total of 16 hours without
yielding any infested stems. However, in
Wasco County (13-16 miles west of Dufur),
8 infested stems were found after a total of
6 hours of searching. Only 4 infested stems
were found at the other sites after 10 hours
of searching. Thus, a total of 12 infested
stems was found from all sites after 30 hours
of searching. The difficulty in finding stems
of H. discolor infested by C. aldrichi could
be a result of not observing the primary
host, which then would remain to be dis-
covered. However, the relative abundance
of infested stems at the Wasco Co. site sug-
gests that the association of C. aldrichi with
205
H. discolor is more than an incidental host
record.
Species of Cephidae either sever the stem
as an adult at the time of oviposition or as
a larva when stem boring is initiated. The
stems harboring larvae of C. aldrichi were
severed in a manner which resulted in a line
of girdling that was very smooth and with-
out jagged marks. In comparison, the girdle
line on severed shoots containing Janus ru-
fiventris (Cresson), in which the adult female
girdles the stem with her ovipositor, is rough
and exhibits jagged marks around the cir-
cumference of the stem (Hanson 1986). In
the absence of direct observations this sug-
gests that in C. a/drichi the shoot is probably
girdled by the larva.
Only the wider stems of H. discolor were
infested. Stem width of all growing shoots
ranged from 0.9 mm to 7.0 mm in diameter.
Infested stems ranged from 2.8—7.00 mm in
diameter when measured at mid-length of
the infested portion.
The pattern of the tunnel in the stem il-
lustrates the behavior of the larva. We noted
patterns in tunnelling that suggest the fol-
lowing: the newly hatched larva after gir-
dling the shoot tunnels toward the base in
the cambial zone for a short distance (1-3
mm), the larva then turns back (upwards)
to the girdled apex, tunnels to the midsec-
tion of the shoot, and then bores down the
center toward the base of the stem. At the
initiation of the downward tunnelling the
larva is consuming almost the entire interior
of the shoot and packing frass in the vacated
tunnel.
Patterns in the length of infested stems
suggested that a second severing of the shoot
occurs. Stems collected in July had tunnels
averaging 5 cm in length (severed apex to
location of larval head), whereas stems col-
lected in October had tunnels less than |
cm in length (newly severed apex to the base
of the prepupal chamber). These observa-
tions suggested that the last instar severs the
stem again, before forming a pupal cham-
ber. Thus, the portion of the stem contain-
206
ing the initial tunnelling falls off the plant.
Similar behavior has been documented in
other cephids (Middlekauff 1969).
Observations of emergence in the labo-
ratory indicated that adults exit the shoot
by chewing through the apical frass plug,
rather than through the stem as in other
Hartigiini.
We observed some larval-pupal mortality
factors. A larva in one of the 12 infested
stems contained larvae of a gregarious Pter-
omalus sp. (Hymenoptera: Pteromalidae;
identification by P.E.H.). These parasitoid
larvae were reared and produced six females
and two males (on July 8) one week after
collection. Two of the stems with tunnels
had irregular holes in the region of the pupal
chamber. The holes exhibited peeled edges
and the chamber lacked a cephid larva, sug-
gesting possible mortality by bird predation.
In three of the infested stems the cephid
larva had already died from unknown caus-
es and in four of the infested stems larvae
died after collection (one was preserved),
probably because they were collected too
early in the season. Thus, we obtained only
two adults (both female) out of 12 infested
stems.
Data from specimens in the Oregon State
University insect collection indicate that
adult C. aldrichi are active from early May
at lower elevations in the Willamette Valley
(about 100 m), to early July at higher ele-
vations on Mary’s Peak (about 1000 m).
Our field observations on the state of larval
development in the field indicated that this
species is univoltine in Oregon.
The only other species presently placed
in Caenocephus is the Palearctic C. lunu-
latus (Strobl), formerly known as C. jakow-
leffi Knonow (Schedl 1981). Because Hol-
odiscus is absent from the Old World, the
host plant of this species must be different
from that of C. aldrichi.
With the discovery of a host plant con-
taining the larva of C. aldrichi, it will be
possible to compare the morphology of im-
mature Caenocephus with that of other gen-
era of Hartigiini. In a key to the larvae of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Cephidae, Middleton (1917) noted that in
Hartigia the lateral area of the basal anal
lobe is setose, whereas in Janus this area is
bare. In the single preserved specimen of C.
aldrichi the lateral area of the basal anal lobe
is setose as in Hartigia. Additional collec-
tions of larval C. aldrichi are needed for
studies on variation in larval morphology.
Based on our limited observations of this
rarely collected species it appears that
Caenocephus is more similar to Hartigia
than to Janus. The following observations
suggest this relationship: host plant is in the
Rosaceae, initial stem-girdling is likely per-
formed by the larva, and the basal anal lobe
is setose in the larva.
ACKNOWLEDGMENTS
We are grateful to D. R. Smith, System-
atic Entomology Laboratory, USDA,
Washington, D.C., for sharing his personal
records on C. aldrichi with us. This is tech-
nical paper no. 8365, Oregon State Univer-
sity Agricultural Experiment Station.
LITERATURE CITED
Bradley, J. C. 1905. Caenocephus in America. Can.
Entomol. 37: 363-364.
Hanson, P.E. 1986. Biology of Janus rufiventris (Hy-
menoptera: Cephidae). Ann. Entomol. Soc. Am.
79: 488-490.
Middlekauff, W. W. 1952. Notes on two species of
California stem borers. Pan. Pac. Entomol. 23:
108-109.
1969. The cephid stem borers of California
(Hymenoptera: Cephidae). Bull. Calif. Insect Surv.,
vol. 11, 19 pp.
Middleton, W. 1917. Notes on the larvae of some
Cephidae. Proc. Entomol. Soc. Wash. 19: 174—
179.
Schedl, W. 1981. Zur Nomenklatur, Morphologie und
Verbreitung der Halmwespe Caenocephus lunu-
latus (Strobl, 1895) comb. nov. (Hymenoptera,
Cephidae). Nachrichtenblatt der Bayerischen En-
tomologen 30: 28-31.
Smith, D. R. 1979. Superfamily Cephoidea, Family
Cephidae, pp. 133-137. Jn K. V. Krombein, P. D.
Hurd, Jr., D. R. Smith, and B. D. Burks, eds.,
Catalog of Hymenoptera in America North of
Mexico, vol. 1. Smithson. Inst. Pr. Wash. D.C.
. 1986. The berry and rose stem-borers of the
genus Hartigia in North America (Hymenoptera:
Cephidae). Trans. Am. Entomol. Soc. 112: 129-
145.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 207-215
PHYTOPHAGOUS INSECT FAUNA OF BACCHARIS SAROTHROIDES
GRAY (ASTERACEAE) IN ARIZONA AND NEW MEXICO
PAUL E. BoL_ptT, WILLIAM Woops, AND THOMAS O. ROBBINS
(PEB, TOR) Grassland, Soil and Water Research Laboratory, USDA, Agricultural Re-
search Service, 808 E. Blackland Rd., Temple, Texas 76502; (WW) Western Australian
Department of Agriculture, South Perth 6151, Western Australia.
Abstract. —Phytophagous insects representing seven orders, 26 families, and 64 species
were collected on the unisexual shrub desert broom, Baccharis sarothroides Gray, a native
plant of the southwestern United States. Twenty-five species fed in two or more plant
families, six species fed only within the Asteraceae, and five species fed only on Baccharis
species. No severe foliage or stem damage due to feeding by the insects was observed.
Key Words:
Desert broom, Baccharis sarothroides
Gray (Asteraceae: Astereae), 1s an attrac-
tive, erect, unisexual, woody shrub, | to 4
m in height with broom-like branches and
resinous evergreen leaves and stems. Native
to the southwestern United States, desert
broom is common in sand or gravel riparian
washes, drainage areas, and low hills at el-
evations of 300-1500 m above sea level
(Benson and Darrow 1981, Lee et al. 1984).
The few leaves present on desert broom
are simple, linear, alternate, and up to 2 cm
long; margins are entire. Flower heads are
dioecious, discoid in shape, usually solitary,
and occur on elongate leafless twigs. Female
heads contain about 10 florets and are 6-8
mm in length; male heads are 3-4 mm in
length. Flowering is dependent on rainfall
and may occur between August and No-
vember. Leaves and stems are not palatable
to livestock. The many achenes are about
0.25 mm long, 10-nerved, glabrous, and
dispersed by wind. The white pappus is 2-
2.5 cm long and conspicuous during bloom
(McGinnies 1986, Munz and Keck 1968,
Vines 1960).
Desert broom, plant-feeding insects, ecology
Desert broom is often seen in the front
yards of residential properties and is sold
by nurseries in Arizona for landscaping in
dry environments because it is drought-re-
sistant and tolerant of temperature extremes
and saline water. Recently, a low-growing
hybrid, B. sarothroides x B. pilularis
DeCandolle, was developed at the Arizona
Agricultural Experiment Station, Tucson,
for use in xeriscaping. This new shrub,
named “Centennial,” is a compact, pros-
trate, leafy, green bush which survives sum-
mer heat of up to 45°C without wilting (Lee
et al. 1984, Thompson 1985).
Desert broom was tested on different soil
types for use in the reclamation of copper
mine waste areas in Arizona. Initially, the
plant did poorly, but in the second year,
height, vegetation yield, and ground cover
compared well with the four other shrubs
in the test (Day and Ludeke 1980). AI-
though two years 1s an insufficient period of
time for a proper evaluation, this shrub may
be one of several plants which are beneficial
for reclaiming disturbed mine soils.
Pellet (1930) considered desert broom to
208
be locally important for honey production
because it blooms in the fall when few other
flowers are available.
Insects were previously collected on des-
ert broom in Arizona from July to Septem-
ber by Meyer et al. (1979). Of the 25 species
collected, 8 were phytophagous and the re-
maining species were predators or parasites.
Other surveys for insects of Baccharis in the
United States have been made of B. hal-
imifolia L. (Bennett 1963, Palmer 1987) and
B. pilularis (Tilden 1951).
We investigated the phytophagous insects
of B. sarothroides as part ofa study of insects
associated with the genus Baccharis. Al-
though this is a beneficial shrub, it is closely
related to B. salicifolia (R&P) Pers., B. ne-
glecta Britt., and B. halimifolia, three weedy
shrubs that we are studying as potential tar-
gets for biological control. This paper is the
second in our series; the first paper (Boldt
and Robbins 1987) reported the phytoph-
agous insects of B. neglecta in Texas.
MATERIALS AND METHODS
We examined plants of B. sarothroides on
33 occasions from June 1985 to September
1987 at sites near Rye, Picacho, Gila Bend,
Tucson, Sasabe, and Dragoon, Arizona as
well as Rodeo and Lordsburg, New Mexico
(Fig. 1). Up to four of the eight sites were
visited each month of the year. At each site,
insects were handpicked, aspirated, or swept
from 10 to 20 plants. Immature insects that
were collected were reared to maturity on
excised plant material and adults found
resting on the plant material were caged on
leaf bouquets in the laboratory to confirm
their ability to feed on the plant. Male and
female flower heads were collected in bulk
from near Tucson in October 1986; some
were dissected while the remainder were held
for emergence. Detailed collection and rear-
ing records were maintained so that we could
estimate the relative frequency of each in-
sect species collected and record collection
data, plant association, and plant phenol-
ogy. We deposited voucher insect speci-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mens in the insect collection of the Temple
laboratory.
RESULTS AND DISCUSSION
The distribution of desert broom is pre-
sented in Fig. 1. This shrub is abundant in
central and southern Arizona but occurs in
the Sonoran Desert and desert grasslands
from New Mexico to California, Baja Cal-
ifornia, and Sonora, Mexico. It is also re-
corded from Sinaloa, Mexico (Benson and
Darrow 1981, Munz and Keck 1968, per-
sonal observations).
Desert broom was the host or alternate
host for 64 species of phytophagous insects
representing seven orders and 26 families
excluding those that feed exclusively on pol-
len (Table 1). At least 38 (59.4%) repro-
duced and developed to maturity on this
plant, and 12 (18.7%) of these were endo-
phagous as immatures.
Three (4.7%) species (Table 1) also feed
on two related shrubs, B. neglecta and B.
halimifolia (Boldt and Robbins 1987, Palm-
er 1987): Nysius raphanus Howard, Lygus
lineolaris (Palisot de Beauvois), and Neo-
lasioptera lathami Gagné. In addition, four
(6.2%) species: Frankiniella occidentalis
(Pergande), Hesperotettix viridis viridis
Thomas, Brochymena quadripustulata (F.),
and Clastoptera lineatocollis Stal (Table 1)
were also collected from B. neglecta in Tex-
as by Boldt and Robbins (1987). Palmer
(1987) listed one (1.6%) species, Acantho-
cephala thomasi (Uhler) from B. halimifo-
lia. Only Chrysobothris bacchari Van Dyke
(Table 1) was also listed by Tilden (1951)
on B. pilularis in California. The aforemen-
tioned insects are polyphagous except for
the gall midge, N. lathami, and the flat-
headed borer, C. bacchari.
The following insects on desert broom
were listed in the literature but not collected
by us: Dactylotum bicolor variegatum
(Scudder), Melanoplus desultorius Rehn,
Aztecacris gloriosa Hebard, Poecilotettix
pantherina (F. Walker), and Poecilotettix
sanguineus Scudder (Ball et al. 1942); Agri-
VOLUME 90, NUMBER 2
CALIFORNIA
SAROTHROIDES
BAJA
Fig. 1.
lus aurilatera Waterhouse (Hespenheide
1974), and Tragidion annulatum LeConte
(Meyer et al. 1979).
In April and May 1987, we collected sev-
eral larvae of each of two noctuids, Cucullia
laetifica Lintner and Platysenta videns
(Guenée), from leaves of desert broom
growing in pots in our nursery at Temple,
Texas. We reared five C. /aetifica and one
P. videns to adults. The only other con-
firmed host plant of C. /aetificais B. neglecta
(Boldt and Robbins 1987). Other Cucullia
feed on leaves and flowers of various As-
teraceae (R. W. Poole, pers. comm.). Pla-
tysenta videns occurs east of the Continental
Divide (Crumb 1956). We, therefore, con-
cluded that this noctuid does not normally
attack desert broom. Crumb (1956), how-
ever, collected larvae of another species on
this plant in Arizona which he identified as
209
Insect collection sites and distribution of Baccharis sarothroides in the United States and Mexico.
Platysenta sp. No. 29 but was unable to rear
them to adult for species identification.
According to published literature and
available host records for the 51 identified
species (Table 1), we determined that 25
(39.1%) were polyphagous because they fed
on plants outside the family Asteraceae.
Eleven (17.2%) of these are pests of eco-
nomically important plants and feed on des-
ert broom during the winter or spring when
their annual hosts are not available. This
percentage is similar to the 18% of identified
insects on B. neglecta which were listed as
pests by Boldt and Robbins (1987) and 1l-
lustrates the role of these shrubs in harbor-
ing pests. We listed six (9.4%) species as
oligophagous because they fed on plants
within the Asteraceae and six (9.4%) as
monophagous because they apparently fed
only on the genus Baccharis. The gall midge
210 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Phytophagous insects collected from B. sarothroides.
Insects
Orthoptera
Acrididae
Aztecacris gloriosa (Hebard)
Dactylotum bicolor vari-
egatum (Scudder)
Hesperotettix viridis viridis
(Thomas)
Melanoplus desultorius
Rehn
M. pictus Scudder
Poecilotettix pantherina (F.
Walker)
P. sanguineus Scudder
Schistocerca alutacea sho-
shone (Thomas)
Thysanoptera
Thripidae
Frankliniella minuta (Moul-
ton)
F. occidentalis (Pergande)
Heteroptera
Coreidae
Acanthocephala femorata
(F.)
A. thomasi (Uhler)
Lygaeidae
Lygaeus reclivatus (Say)
Melanopleurus belfragei
(Stal)
Nysius raphanus Howard
Ochrimnus foederatus (van
Duzee)
Miridae
Lygus desertinus Knight
L. hesperus Knight
L. lineolaris (Palisot de
Beauvois)
Parthenicus baccharidus
Knight
Rhinacloa forticornis (Reu-
ter)
Month Collected
Immature
Aug.—Oct.
June
Aug.-Oct.
Apr.—Aug.
Mar.—Apr.
Feb.-Oct.
Aug.—Sept.
Aug.—Sept.
Sept.—Jan.
Oct.—Dec.
Oct.-Dec.
Oct.-Dec.
Apr.—Sept.
Adult
Oct.—Jan.
June—Oct.
Aug.—Sept.
July-Nov.
Aug.—Oct.
Apr.-Nov.
Apr.—Aug.
Sept.—Oct.
Oct.
Feb.-Oct.
July—Sept.
July—Sept.
Oct.—Dec.
Aug.—Oct.
May-Jan.
Oct-Nov.
May-Dec.
May-Dec.
May-Dec.
Apr.—Jan.
Aug.—Nov.
Rela-
tive
quency
Ore Ar @
vs)
v@)
~ © © OF
Associated
Plant Part®
L, F, St
Host
Speci-
ficity®
PE
PE
PE
PE
PE
References
Ball et al.
1942
Ball et al.
1942
Ball et al.
1942
Ball et al.
1942
Ball et al.
1942
Ball et al.
1942
Ball et al.
1942
Ball et al.
1942
Yudin et al.
1986
Meyer et al.
1979
Meyer et al.
1979
Ward et al.
1977
Brailovsky
1982
Graham et al.
1986
Young 1986
VOLUME 90, NUMBER 2
Table 1. Continued.
Month Collected
Insects
Pentatomidae
Brochymena quadripustula-
ta (F.)
Homoptera
Acanaloniidae
Acanalonia clypeata Van
Duzee
Acanalonia fasciata Metcalf
Aphididae
Brachycauda helichrysi
(Kaltenbach)
Cercopidae
Clastoptera lineatocollis Stal
Cicadellidae
Aceratagallia sp. A
Aceratagallia sp. B
Empoasca sp.
Homalodisca lacerta (Fow-
ler)
Idiocerus sp.
Stragania robusta (Uhler)
Cixudae
Oecleus productus Metcalf
Eriococcidae
Ovaticoccus californicus
McKenzie
Flatidae
Mistharnophanita sonorana
Kirkaldy
Ormenis prob. yumana Ball
Membracidae
Hypsoprora neglecta Ball
Spissistilus festinus (Say)
Psyllidae
Trioza collaris Crawford
Coleoptera
Buprestidae
Agrilus aurilatera Water-
house
Chrysobothris bacchari
VanDyke
C. beyeri Schaeffer
Immature
Aug.—Sept.
Mar.—May
Mar., Aug.
May
Aug.—Sept.
Nov.—Jan.
Sept.—June
Sept.—June
Adult
Feb.-May
Aug.—Sept.
Aug.-Sept.
Mar.—May
May, Aug.—
Dec.
Jan., May,
July
July—Aug.
Jan.—Dec.
Aug.—Nov.
Nov.—Mar.
Aug.—Sept.
Apr.—Sept.
Aug.
May, Aug.—
Sept.
May-Nov.
Mar., Aug.—
Oct.
Aug.—Mar.
June-July
June
Rela-
live
Fre-
quency*
O
Onn
(eo)
one)
Associated
Plant Part?
St
sot
L, St
eat
est
Lest
Ags
Ae)!
Last
est
L, St
| OF se
Host
Speci-
ficitys
PE
RE
References
Gamboa and
Alcock
1973
Doering 1942
Fletcher 1937
McKenzie
1964
Mueller and
Dumas
1987
Tuthill 1945
Hespenheide
1974
Nelson et al.
1981
Werner (pers.
comm.)
DZ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Month Collected
Table 1. Continued.
Insects Immature
Cerambycidae
Stenodontes lobigenis Sept.—June
(Bates)
Tragidion annulatum Le- Oct.-Aug.
Conte
Chrysomelidae
Exema deserti Pierce May-Aug.
Systena blanda Melsh.
Curculionidae
Anthonomus Stolatus Fall
Lixus pervestitus Chittenden
Smicronyx undescribed sp.
Lepidoptera
Ctenuchidae
Ctenucha venosa Walker Aug.
Gelechiidae
Aristotelia argentifera Busck | May-Sept.
Gnorimoschema unde- Apr.—Nov.
scribed sp.
Geometridae
Anavitrinelia sp. Feb.—Mar.
Elpiste metanemaria (Hulst) | Mar.—May,
Aug.—Sept.
Lyonetiidae
Bucculatrix sp. near seorsa Dec., Apr.-
Braun May, Aug.
Pterophoridae
genus unknown Mar.—June
Diptera
Cecidomyiidae
Neolasioptera lathami Mar.—Oct.
Gagné
Tephritidae
Aciurina mexicana (Aczél) Feb.—Apr.
A. thoracica Curran Feb.—Apr.
Euarestoides acutangulus
(Thomson)
E. flavus (Adams) Oct.—Jan.
Tephritis arizonaensis Quis- — Oct.-Nov.,
enberry Mar.—Apr.
Trupanea nigricornis (Co- Oct.-Nov.
quillett)
Trupanea wheeleri Curran Oct-Nov.
Adult
July—Sept.
July—Sept.
May-Sept.
Aug.—Sept.
Aug.—Sept.
Sept.
Aug.—Sept.
Sept.-Oct.
May-Oct.
Dec.—Mar.
Mar.—Apr.
Mar.—June,
Sept.-Nov.
Jan.—Feb.
June, Aug.-
Sept.
Mar.—Oct.
Oct.—June
Feb.-July
Oct.—Feb.
Aug.—Apr.
Jan.—Dec.
Nov.
Mar., Aug.—
Nov.
Rela-
uve
Fre-
quency*
oO
Pe)
ooke)
Associated
Plant Part?
R
St,R
LF
St
St
St,G
F, St, G
Host
Speci-
ficity*
PE
PE
OE
References
Werner (pers.
comm.),
Linsley
1962
Meyer et al.
1979: Lins-
ley 1962
Werner (pers.
comm.)
Braun 1963
Diatloff and
Palmer
1986
Steyskal 1984
Steyskal 1984
Goeden and
Ricker
1986
Wasbauer
1972
Foote and
Blanc 1963
Goeden 1985
Goeden 1985
VOLUME 90, NUMBER 2
was tested for host specificity and released
in Australia as a biological control agent for
B. halimifolia (Diatloff and Palmer 1987).
The remaining five monophagous species
may also be potential biological control
agents for B. halimifolia, although they were
not tested for host specificity and their im-
pact on the plant was not assessed.
The host specificity of 29 (45.3%) species
collected on desert broom was not deter-
mined because their identification was in-
complete or host records were unknown or
notavailable. Based on the host plant record
of other species in the genus, some of these
may be found at a later date to be monoph-
agous or oligophagous. At least two of the
unidentified insects were apparently unde-
scribed.
We collected insects on desert broom dur-
ing every month of the year. The largest
number of adult species, 39 (60.9%), was
collected in both August and September
when normal daily mean temperatures were
21 and 23°C (Wallis 1977) and the least
number, 9 (14.1%), was collected in January
when normal daily mean temperature was
1O:5°C.
About 40 (62.5%) insect species, most of
which were Hemiptera and Homoptera, fed
on the leaves and small stems; 21 (32.8%)
species fed entirely or partly on the flowers
and another 12 (18.7%) fed in the stems or
both roots and stems. Various insects, such
as bees, ants, syrphids, and beetles were also
encountered feeding on the resinous exu-
date of the stems and leaves but they were
not collected.
Seventeen (26.6%) species were recorded
as rare because they were encountered at a
density of less than one per ten plants. Many
of them were identified only to genus, and
many are polyphagous insects for which
213
desert broom may not be an important plant.
Of the 15 (23.4%) species that were record-
ed as common because they were encoun-
tered at a density of more than one per plant,
seven were polyphagous species of sap-feed-
ing Hemiptera and three were species of
stem- or flower head-feeding Diptera.
At no time during our collecting did we
observe widespread foliage or stem damage
due to feeding insects, although we occa-
sionally found isolated areas of shrubs with
stem and leaf damage which we attributed
to feeding by grasshoppers. No obviously
destructive insects were encountered on
desert broom such as the chrysomelid leaf
feeder, Trirhabda bacharidis (Weber), on B.
neglecta and B. halimifolia (Boldt and Rob-
bins 1987), or 7. flavolimbata (Manner-
heim) and the gall midge, Rhopalomyia cal-
ifornica (Felt), on B. pilularis (Tilden 1951).
ACKNOWLEDGMENTS
We gratefully thank Bradford Burk (Par-
kinsonia Biological Control Laboratory,
Western Australian Department of Agri-
culture, Tucson, Arizona), J. P. Cuda and
C. J. DeLoach (Grassland, Soil and Water
Research Laboratory, USDA, Agricultural
Research Service, Temple, Texas) for their
technical assistance and E. W. Baker, D. C.
Ferguson, R. C. Froeschner, R. J. Gagné, P.
M. Marsh, D. R. Miller, S. Nakahara, A. L.
Norrbom, R. W. Poole, R. K. Robbins, T.
J. Spilman, M. B. Stoezel, R. E. White and
D. R. Whitehead (Systematic Entomology
Laboratory, USDA, Agricultural Research
Service, Beltsville, Maryland), D. R. Davis
(National Museum of Natural History,
Washington, D.C.), L. B. O’Brien (Florida
A&M University, Tallahassee), and H. R.
Burke (Texas A&M University, College Sta-
tion) for identification of insects mentioned
oes
*C, common, more than one per plant; O, occasional, less than one per plant but more than one per 10 plants:
R, rare, less than one per 10 plants.
'L, leaf; St, stem; R, root; F, flower head; Sd, seed (achenes); G, gall on stem.
*M, monophagous (apparently restricted to the genus Baccharis); O, oligophagous (apparently restricted to
the Compositae); P, polyphagous (apparently feeds on various plant families); E, economically important (see
Literature Cited).
214
in this report. Host plant records were fur-
nished by F. W. Werner (University of Ar-
izona, Tucson). We thank C. Mason, Jr.,
(University of Arizona, Tucson) for access
to herbarium records.
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Benson, L. and R. Darrow. 1981. Trees and Shrubs
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PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 216-228
THE PHYTOPHAGOUS INSECT FAUNA ASSOCIATED WITH
BACCHARIS HALIMIFOLIA L. IN THE
EASTERN UNITED STATES
W. A. PALMER AND F. D. BENNETT
(WAP) Northern American Field Station, Queensland Department of Lands, 2714
Pecan Drive, Temple, Texas 76502; (FDB) Entomology and Nematology Department,
University of Florida, Gainesville, Florida 32611 (formerly Director, Commonwealth
Institute of Biological Control, Trinidad).
Abstract. —A survey of the phytophagous insects found on Baccharis halimifolia along
the eastern seaboard of the United States was undertaken as part of an extensive program
to find biological control agents for this plant in Australia. One hundred and seventy four
phytophagous insect species were collected or were recorded in the host records of the
Division of Plant Industry, Bureau of Entomology insect collection at Gainesville. Four-
teen species were considered to be monophagous and potential biological control agents.
Lepidoptera and endophages constituted a high proportion of this group. Fifty five general
predators and 51 agricultural pests were also found on B. halimifolia.
Key Words:
Baccharis halimifolia L. (Family Astera-
ceae: Tribe Astereae: Sub-Tribe Bacchari-
nae) is a North American shrub that has
become a noxious weed in Queensland,
Australia (Stanley and Ross 1986). As part
of its effort to control this weed, the Queens-
land Department of Lands initiated a pro-
gram in 1960 to find suitable biological con-
trol agents from the New World where the
Baccharinae are native.
B. halimifolia is found along the eastern
seaboard of the United States from Florida
to Massachusetts. It was probably intro-
duced into Australia from Florida which has
a subtropical climate most closely approx-
imating that of southeast Queensland where
B. halimifolia is most troublesome. The
eastern seaboard was therefore selected as
a very appropriate area in which to survey
the insect fauna associated with this plant.
From the survey it was hoped that mono-
phagous species suitable for importation and
Biological control, survey, weed
release into Australia could be selected for
further study.
Various surveys of insect faunas on Bac-
charis have already been reported. Tilden
(1951) listed 221 insects, including 55 pri-
mary herbivores, associated with the veg-
etative parts of B. pilularis DC. F. D. Ben-
nett (unpublished) surveyed the fauna on
various species of Baccharis in Brazil. Kraft
and Denno (1982) listed the major foliage-
feeding herbivores attacking B. halimifolia
in Maryland. Palmer (1987) surveyed the
insect fauna on B. halimifolia and the closely
related B. neglecta Britton in Louisiana,
Texas, and northern Mexico and reported
133 phytophagous species, of which 11 were
considered monophagous. Boldt and Rob-
bins (1987) surveyed B. neglecta in Texas
and reported 91 phytophagous species.
B. halimifolia is a perennial, dioecious
woody shrub that grows to a height of 15
feet. It produces new growth in spring, and
VOLUME 90, NUMBER 2
the quality of the foliage in terms of nitrogen
content, moisture content, secondary chem-
icals, and toughness declines as the year pro-
gresses (Kraft and Denno 1982). It flowers
in autumn, producing a prodigious quantity
of seed (Panetta 1979). The phenology of
this plant is described in more detail by
Palmer (1987).
THE AREA AND METHODS OF SURVEY
The area between southern Florida and
Washington, D.C. was first surveyed on two
car trips of 3-4 weeks in 1962. The first trip
was made in spring when the B. halimifolia
was producing new foliage, and the second
was undertaken in October when the plants
were in full flower. Two to three sites, ap-
proximately 50 miles apart, were examined
each day on these trips. From 1982-1987
further surveying was undertaken on visits
of a few days to Miami and Gainesville,
Florida; to Charleston, South Carolina; to
Williamsburg, Virginia; and to Toms River,
New Jersey. In 1983 a site near Gainesville
was also inspected each month.
Collecting procedures were much as de-
scribed by Palmer (1987). Insects were found
by visual inspection and sweeping the fo-
liage. Inflorescences were examined under
the microscope. Immatures were reared
through to adulthood to enable them to be
accurately identified. All insects were sent
to expert taxonomists (cf. acknowledg-
ments) for their identification.
A second source of data was the collection
and files of the Bureau of Entomology, Di-
vision of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services
(DPI), in Gainesville. This collection main-
tains a catalogue of host records for all in-
sects submitted for identification. Because
the authors had no control over the collec-
tion or treatment of these data, information
from this collection is clearly marked in the
tables to distinguish it from our own col-
lections.
Identified insects were classified as mono-
phagous if restricted to Baccharis, oli-
217
gophagous if the host range was restricted
to the tribe Astereae and polyphagous if
having a wider host range. Evidence of host
range was obtained from observations dur-
ing the course of the survey, consultations
with acknowledged experts, examination of
host data attached to specimens in major
insect collections, information in texts such
as Arnett (1985), Slater and Baranowski
(1978), Arnett et al. (1980), Smith and Smith
(1978), Baranowski and Slater (1986), and
Borror et al. (1981), and, in some instances,
formal host testing.
Insects were classified as endophagous if
they were found feeding on internal tissues
of Baccharis and ectophagous if they fed
externally on Baccharis. They were classi-
fied as pest species if mentioned as such in
Arnett (1985).
Monophagous species were considered
potential agents for biological control of B.
halimifolia and their potential was rated
subjectively by the authors and objectively
by applying the formula of Goeden (1983).
An insect can score a maximum of 79 points
by this formula and is classified as effective,
partially effective or ineffective if it scores
>50, 20-50, or <20, respectively.
RESULTS
The phytophagous fauna (excluding pol-
len and nectar gatherers) found on B. hal-
imifolia are shown in Table 1. One hundred
and eight species were collected and a fur-
ther 66 species were obtained from the DPI
files. The Acari, Hemiptera, Homoptera,
Lepidoptera, Coleoptera and Diptera were
represented by 5 (3% of total species), 20
(11%), 71 (41%), 22 (13%), 43 (25%) and 13
(7%) species, respectively.
Fourteen species (8% of the total species)
were considered to be monophagous. Three
species were considered oligophagous, and
the remainder were either polyphagous or
host range unknown. Of the monophagous
species, 7 (50% of the monophagous species)
were Lepidoptera and 8 (57%) were endoph-
agous for at least part of their life cycle.
218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Phytophagous species collected on B. halimifolia along the eastern seaboard of the United States.
Location Insect-Host Rela- Pest
Species! (State) tionship to Baccharis Specificity? Status?
Acari
Eriophyidae
*4ceria nr. baccharices Kiefer Fl ectophagous “ts
*Tegonotus acidotus (Keifer) Fl ectophagous *2
*Tegonotus nr. baccharis (Keifer) Fl ectophagous B2
Tegonotus undescribed sp. Fl ectophagous m7
Tetranychidae
*Paratetranychus sp.
Hemiptera
Coreidae
Acanthocephala confraterna (Uhler) Ga-Fl ectophagous *
Acanthocephala femorata (F.) SC-FI ectophagous =
Acanthocephala terminalis (Dallas) Fl ectophagous =
Catorhintha guttula (F.) Fl ectophagous U
*Euthochtha galeator (F.) Fl ectophagous *
Leptoglossus phyllopus L. NC-FI ectophagous = i
Merocoris typhaeus Fab. Fl ectophagous :
Lygaeidae
*Ochrimnus lineoloides Slater FI ectophagous yi
Ochrimnus mimulus (Stal) Fl ectophagous ns
*Palagonatus divergens Distant Fl
Miridae
Adelphocoris rapidus (Say) NC ectophagous * ~
Lopidea hesperus (Kirkaldy) Fl ectophagous s
Slaterocoris pallipes (Knight) NY-FI ectophagous xe
Taylorilygus pallidulus (Blanchard) Ga-Fl ectophagous *
Pentatomidae
*Euschistus crassus Dallas Fl ectophagous *
Euschistus servus Say SC-Fl ectophagous a x
*Loxa sp. Fl ectophagous
Mormidea sp. Fl ectophagous
Tingidae
Corythucha baccharidis Drake Fl ectophagous ax
*Corythucha marmorata (Uhler) Fl ectophagous x
Homoptera
Acanaloniidae
*4Acanalonia latifrons Walker Fl ” ectophagous Wi
Aleyrodidae
*Bemesia berbicola Cockerell Fl ectophagous *
*Paraleyrodes naranjae Dozier Fl ectophagous x
Aphididae
Aphis coreopsidis (Thomas) SC-Fl ectophagous
Aphis gossypii Glover Fl ectophagous
Macrosiphum sp. Fl ectophagous
Myzus persicae (Sulzer) Fl ectophagous
Toxoptera aurantil (Fonscolombe) Fl ectophagous
Uroleucon eupatoricolens (Patch) Fl ectophagous
Uroleucon gravicornis (Patch) Fl ectophagous ac
Cercopidae
*4Aphrophora sp. Fl ectophagous
Clastoptera obtusa Say Fl ectophagous
Clastoptera xanthocephala Germar NJ ectophagous
VOLUME 90, NUMBER 2 219
Table 1. Continued.
Location Insect—Host Rela- Pest
Species! (State) tionship to Baccharis Specificity? Status?*
Cicadellidae
*Empoasca kraemeri Ross & Moore Fl ectophagous * *
*Empoasca sp. Fl ectophagous
*Graminella nigrifrons (Forbes) Fl ectophagous = *
*Graphocephala coccinea (Forster) Fl ectophagous ae
*Graphocephala versuta (Say) Fl ectophagous
Gyponana sp. Fl ectophagous
*Paraphlepsius sp. Fl ectophagous
*Penthimia nr. americana Fitch Fl ectophagous *
*Ponana sp. Fl ectophagous
Oncometopia nigrifrons (Walker) Fl ectophagous Gs
*Scaphytopius sp. Fl ectophagous
Cixiidae
*Bothriocera sp. Fl ectophagous
*Myndus crudus Van Duzee Fl ectophagous =
Myndus pallidus Caldwell Fl ectophagous
Oliarus sp. NJ ectophagous
Coccidae
Ceroplastes ceriferus (F.) Fl ectophagous * *
*Ceroplastes cirripediformis Comstock Fl ectophagous Ys *
*Ceroplastes floridensis Comstock Fl ectophagous as *
Coccus hesperidum L. Fl ectophagous bs =
*Coccus longulus (Douglas) ial ectophagous <
Coccus viridis (Green) Fl ectophagous z 5
*Eucalymnatus tessellatus (Signoret) Fl ectophagous *
Kilifia acuminata (Signoret) Fl ectophagous *
Kilifia elongatus (Signoret) Fl ectophagous *?
*Parasaissetia nigra (Nietner) Fl ectophagous 5 ‘3
Protopulvinaria pyriformis (Cockerell) FI ectophagous bs =
*Pulvinaria innumerabilis (Rathvon) Fl ectophagous i *
Pulvinaria psidii Maskell lal ectophagous * a
Pulvinaria urbicola Cockerell Fl ectophagous * Z
Saissetia coffeae (Walker) Fl ectophagous + i
*Saissetia miranda (Cockerell & Parrott) Fl ectophagous ag =
Saissetia neglecta DeLotto all ectophagous * ce
Saissetia oleae (Olivier) Fl ectophagous * *
Delphacidae
Stobaera pallida Osborn NJ-Fl ectophagous a
Diaspididae
Abgrallaspis cyanophylli (Signoret) Fl ectophagous * *
Aonidomytilus solidaginis (Hoke) Fl ectophagous i
Hemiberlesia lataniae (Signoret) Jel ectophagous s =
Melanaspis similacis (Comstock) Fl ectophagous ag
*Pinnaspis strachani (Cooley) Fl ectophagous * =
*Pseudaonidia trilobitiformis (Green) Fl ectophagous - oe
Rhizaspidiotus dearnessi (Cockerell) Ga ectophagous * ‘
*Velataspis dentata (Hoke) Fl ectophagous i *
Flatidae
Anormenis septentrionalis (Spinola) Fl ectophagous * -
Cyarda melichari Van Duzee Fl ectophagous 3 “a
Fulgoridae
Cyrpoptus reineckei Van Duzee SC ectophagous
Poblicia fuliginosa Olivier Ga ectophagous
220 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Continued.
Location Insect-Host Rela- Pest
Species! (State) tionship to Baccharis Specificity? Status?
Membracidae
*4cutalis tartaren nigrinervis Fowler Fl ectophagous
Acutalis tartaren semicrema (Say) Fl ectophagous
*Campylenchia latipes (Say) Fl ectophagous
*Micrutalis sp. Fl ectophagous
*Spissistilus festinus (Say) Fl ectophagous
Umbonia crassicornis (A. and S.) Fl ectophagous
Vanduzeea arquata Say Fl ectophagous =
Ortheziidae
Orthezia insignis Brown Fl ectophagous i x
Pseudococcidae
Dysmicoccus sp. Fl ectophagous
*Dysmicoccus bispinosis Beardsley Fl ectophagous U
*Planococcus citri (Risso) Fl ectophagous +
Pseudococcus sorghiellus Forbes Ga-Fl ectophagous *
Lepidoptera
Arctiidae
*Estigmene acrea (Drury) Fl ectophagous =
Cochylidae
Lorita baccharivora Pogue SC-Fl ectophagous bi
Coleophoridae
Coleophora sp. Va-Fl ectophagous
Cossidae
Prionoxystus piger (Grote) Fl endophagous +e
Prionoxystus robiniae (Peck) Fl endophagous * *
Gelechiidae
Aristotelia ivae Busck SC-Fl ectophagous ase
Dichomeris serrativittella Zeller Fl ectophagous U
Gnorimoschema sp. Fl endophagous
Geometridae
Anacamptodes defectaria (Guenée) NC ectophagous -
* 4navitrinella pampinaria (Guenée) 1s) ectophagous is =
*Eusarca fundaria (Guenée) Fl ectophagous U
Itame varadaria (Walker) SC-Fl ectophagous a
Lyonetiidae
Bucculatrix ivella Busck NJ-Fl endo and ecto =t%
Noctuidae
Spodoptera ornithogalli (Guenée) Fl ectophagous BS na
Spragueia onagrus (Guenée) Fl U
Psychidae
*Cryptothelia sp. Fl
Pyralidae
Glyphodes floridalis (Fernald) Fl ectophagous U
Pterophoridae
Oidaematophorus balanotes (Meyrick) NJ-Fl endophagous bday
Tortricidae
Choristoneura parallela (Robinson) NC ectophagous - *
Epiblema discretivana (Heinrich) SC-Fl endophagous gs
Epiblema nr. scudderiana (Clemens) NC-Fl endophagous =
Sparganothis sulfureana (Clemens) Fl ectophagous =
Anthribidae
*Toxotropis floridanus Leng Fl ectophagous
OT
VOLUME 90, NUMBER 2 221
Table 1. Continued.
Location Insect-Host Rela- Pest
Species! (State) tionship to Baccharis Specificity? Status*
Buprestidae
*Chrysobothris chrysoela (Illiger) Fl endophagous
Chrysobothris femorata (Olivier) Fl endophagous * od
Cerambycidae
Amuniscus perplexus (Haldeman) Ga-Fl endophagous rie +
*4ncylocera bicolor (Olivier) Fl *
*4nelaphus inermis (Newman) Fl *
*Leptura sp. Fl
Sternidius rusticus (LeConte) NJ U
Typocerus zebra Olivier SC U
Chrysomelidae
*4ltica ludoviciana Fall Fl ectophagous *
Anomoea laticlavia (Forster) SC ectophagous =?
*Bassareus brunipes Olivier Fl U
*Chlamisus sp. Fl
*Chrysomela scripta F. Fl U
Colaspis recurva Blake Va U
Cryptocephalus pumilis Haldeman SC-Fl ectophagous *
Diabrotica balteata LeConte Fl ectophagous * *
Diabrotica undecimpunctata howardii Barber Ga-Fl ectophagous * =
Diachus auratus (F.) SC ectophagous
*Disonychya conjugata F. Fl ectophagous bg
*Exema gibba F. Fl ectophagous U
Exema neglecta Blatchley SC-Fl ectophagous =
*Pachybrachys sp. Fl ectophagous
Paria aterrima Olivier SC-FI ectophagous U
*Triachus cerinus LeConte Fl ectophagous U
Trirhabda bacharidis (Weber) NJ-Fl ectophagous zee
Curculionidae
Apion metallicum Germar Fl ectophagous * *
Apion sp. Fl ectophagous
Artipus floridanus Horn Fl ectophagous = *
*Baris sp. Fl
*Centrinaspis albotectus Casey Fl =
*Chalcodermus aeneus Bohemann Fl = *
Curculio sp. Fl
*Diaprepes abbreviatus (L.) ail * a
Epicaerus formidolosus Boheman Fl ectophagous =
*Nicentrus grossulus Casey Fl =
Notolomus basalis LeConte Fl ectophagous -
Pachnaeus opalus (Olivier) Fl ectophagous bs
Rhodobaenus tredecimpunctatus (Illiger) Fl ectophagous “
Sitophilus oryzae L. Fl ectophagous = =
Tanymecus lacaena (Herbst) Fl ectophagous be
Scarabaeidae
*Pachystethus marginatus F. Fl i
Popillia japonica Newman Va ectophagous is se
Diptera
Agromyzidae
*4mauromyza maculosa (Malloch) Fl endophagous =
*Liriomyza trifolii (Burgess) Fl endophagous “2
Melanagromyza sp. Fl endophagous
NR
Ne
tN
Table 1. Continued.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Species!
Nemorimyza posticata (Meigen)
Phytobia sp.
Cecidomyiidae
Contarinia nr. perfoliata
Dasineuria undescribed sp.
Neolasioptera baccharicola Gagné
Neolasioptera lathami Gagné
Neolasioptera undescribed sp.
Prodiplosis undescribed sp.
Tephritidae
Paroxyma sp.
Tephritis subpura (Johnson)
'* = Record from DPI collection card file.
Location Insect-Host Rela- Pest
(State) tionship to Baccharis Specificity? Status?
SC-Fl endophagous x
SC-Fl endophagous
Md ectophagous U
Md ectophagous
Va endophagous aa
NJ-Fl endophagous =
Md ectophagous
Md ectophagous
Ga ectophagous
NC-Fl endophagous a
2 *** — Monophagous (host-plants apparently restricted to the genus Baccharis); ** = oligophagous (host-
plants apparently restricted to the tribe Astereae); * = polyphagous (having a wider host range than above two
categories); *? = specificity unknown but very likely polyphagous; U = specificity unknown.
3* = Pest species.
Conversely, 8 of 18 (44%) endophagous
species were monophagous.
Only 2 of the monophagous species, P.
piger and N. baccharicola, had a limited
geographic distribution. The other 12 species
were found in at least 2 states and 6 species
were found throughout the survey area. Ten
of the 14 species (72%) were found west of
the Mississippi River by Palmer (1987). By
contrast only 27 of the total number of
species (16%) were common to this survey
and that of Palmer (1987).
Nearly one third of the phytophagous
species (51 species) were pests of agricul-
tural or ornamemental plants. These in-
cluded well known pests such as the brown
stinkbug, Euchistus servus (Say); the green
peach aphid, Myzus persicae (Sulzer); the
black citrus aphid, Toxoptera aurantii (Fon-
scolombe); the green scale, Coccus viridis
(Green); the green shield scale, Pulvinaria
psidii Maskell; the carpenterworm, Priono-
xystus robiniae (Peck), the yellow-striped
armyworm, Spodoptera ornithogalli (Gue-
née); and the southern corn rootworm, Dia-
brotica undecimpunctata howardi Barber.
Non-phytophagous insects such as polli-
nators, predators, parasitoids, nectar gath-
erers, and casual associates that were col-
lected or reared during the survey are listed
in Table 2. This list includes 55 predatory
species.
NOTES ON THE More IMPORTANT SPECIES
The phenologies of 7rirhabda bacharidis
Weber, Amniscus perplexus (Haldeman),
Oidaematophorus balanotes (Meyrick),
Bucculatrix ivella Busck, Aristotelia ivae
Busck, Epiblema discretivana (Heinrich),
Neolasioptera lathami Gagné, Tephritis
subpura (Johnson), Ochrimnus mimulus
(Stal), and Stobaera pallida Osborn were de-
scribed by Palmer (1987). These species were
all commonly encountered along the eastern
seaboard. In Florida, however, the phenol-
ogies of 7. bacharidis and O. balanotes were
different from the previous description
(Palmer 1987). Here larvae of T. bacharidis
were commonly found in the autumn and
early spring, suggesting a partial second gen-
eration or some populations being asyn-
chronous. Similarly, O. balanotes was not
discretely univoltine in Florida. A survey in
February revealed a range of immature
stages from early instars to pupae.
The black mirid, S/aterocoris pallipes
(Knight), was abundant. It occurs further
south, however, than Wheeler (1981) de-
scribed with the southern limit of its range
at Gainesville, Florida.
VOLUME 90, NUMBER 2 223
Table 2. Parasitoids, predators, and incidental visitors recorded, reared or collected on B. halimifolia during
the course of the survey.
Species! Habit
Acari
Bdellidae
*Bdellodes longirostris (Hermann) general predator
Phytoseiidae
*Typhlodromalus peregrinus (Muma)
Passalozetidae
*Passalozetes sp.
Tydeidae
*Lorryia formosa Cooreman
*Tydeus nr. munsteri Meyer and Ryke
Araneae
Anyphaenidae
*4ysha sp.
Araneidae
*4draneus mimiatus (Walckenaer)
*4rgiope sp.
*Conepeira mineatus (Walckenaer)
*Neoscona sp.
Clubionidae
*Clubiona maritima L. Koch
*Trachelas volutus Gertsch
Salticidae
*Hentzia ambigua (Walckenaer)
*Hentzia mitrata Hentz
Theridiidae
*4nelosimus studiosus (Hentz)
*4nelosimus textrix (Walckenaer)
*Theridion flavonotatum (Becker)
Thomisidae
*Misumenops oblongus (Keyserling)
Thysanoptera
*Diceratothrips sp.
*Leptothrips mali (Fitch)
Hemiptera
Anthocoridae
Orius insidiosus (Say)
Nabidae
Nabis capsiformis Germar
Pentatomidae
Euthyrhynchus floridanus (L.)
*Podisus maculiventris (Say)
Stiretrus anchorago (F.)
Phymatidae
Phymata fasciata fasciata (Gray)
Phymata fasciata mystica Evans
Reduviidae
*Apiomerus spissipes (Say)
Pselliopus cinctus F.
Zelus longipes (L.)
Zelus cervicalis Stal
Zelus longipes (L.)
general predator
incidental
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
general predator
224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Continued.
en Un IID SSSSSSSSSS
Species!
Habit
ne CEE yt SISSISSSSSSSISISSSSSSSSSSSSSSSSS SS
Neuroptera
Chrysopidae
Chrysopa spp.
Lepidoptera
Phycitidae
Laetilea coccidivora Comstock
Coleoptera
Cantharidae
Cantharis sp.
Chauliognathus marginatus (F.)
Discodon sp.
Coccinellidae
Adalia bipunctata (L.)
*4zya orbigera Mulsant
Coleomegilla maculata (DeGeer)
*Cryptolaemus montrouzeiri Mulsant
Cycloneda sanguinea (L.)
Exochomus childreni Mulsant
Hippodamia convergens Guerin
Hyperaspis signata Olivier
* Microwelsea sp.
Olla y-nigrum (Mulsant)
Scymnus creperus Mulsant
Scymnus fraternus LeConte
Elateridae
Ampedus luteolus (LeConte)
*Melanotus communis (Gyllenhal)
Scarabaeidae
Trigonopeltastes delta (Forster)
Diptera
Asilidae
Asilus sp.
*Ommatius tibialis Say
Bibionidae
Plecia nearctica Hardy
Chamaemyiidae
Leucopis americana Malloch
Micropezidae
*Taeniaptera trivatta Macquart
Otitidae
*Euxesta notata (Wiedemann)
Platystomatidae
Rivellia steyskali Namba
Sciomyzidae
Dictya sp.
Syrphidae
Pseudodoros clavatus (F.)
Tabanidae
*Chrysops flavidus Wiedemann
Tabanus imitans Stone
Tachinidae
Lixophaga sp.
aphid predators
coccid predator
pollen feeder
pollen feeder
pollen feeder
aphid predator
soft scale predator
aphid predator
mealybug predator
aphid predator
soft scale predator
aphid predator
soft scale predator
diaspine scale predator
aphid predator
aphid predator
aphid predator
incidental
incidental
incidental
general predator
general predator
incidental
aphid predator
incidental
incidental
incidental
incidental
aphid predator
incidental
incidental
incidental
SE
VOLUME 90, NUMBER 2
Table 2. Continued.
Species!
Hymenoptera
Aphelinidae
Centrodora cercopiphagus (Milliron)
Coccophagus sp. |
Coccophagus sp. 2
Aphiidae
*Diaeretiella sp.
Lysiphlebus testaceipes (Cresson)
Bethylidae
Parisarola sp.
Braconidae
Apanteles undescribed sp.
Apanteles epinotiae Viereck
Apanteles forbesi Viereck
Agathis texana Cresson
Bucculatriplex sp.
Chelona sp.
Chelonus (Microchelonus) sp.
Macrocentrus cerasivoranae Viereck
Macrocentrus delicatus Cresson
Macrocentrus pallister Degant
Microgaster mediata Cresson
Mirax texana Muesebeck
Opius undescribed sp.
Optus undescribed sp.
Ceraphronidae
Lygocerus sp.
Chalcididae
Spilochalcis sanguineventris (Cresson)
Cynipidae
Gonaspis potentillae Bass
Eupelmidae
* Anastatus sp.
Eupelmus sp.
Eupelmus sp.
Eupelmus sp.
Eupelmus sp.
Eupelmus sp.
Eulophidae
Achrysocharella sp.
Chrysocharis parksi Crawford
Cirrospilus girualti Peck
Derostenus sp.
Tetrastichus minutus (Howard)
Eurytomidae
Eudecatoma quercilanae (Fitch)
Eurytoma sp.
Formicidae
*Crematogaster ashmeadi Mayr
*Crematogaster atkinsoni Wheeler
*Dolichocerus pustulatus Mayr
*Dorymyrmex pyramicus (Rogor)
*Hypoclinea mariae Forel
to
Habit
egg parasite of Clastoptera
parasite of Coccus hesperidum
parasite of Pulvinaria urbicola
aphid parasite
aphid parasite
parasite of Epiblema discretivana
parasite of Bucculatrix ivella
parasite of Lepidoptera defoliator
parasite of Lepidoptera defoliator
parasite of Lepidoptera
parasite of Bucculatrix ivella
parasite of Oidaematophorus balanotes
parasite of Oidaematophorus balanotes
parasite of Oidaematophorus balanotes
parasite of Lepidoptera defoliator
parasite of Bucculatrix ivella
parasite of agromyzid
parasite of agromyzid
parasite of Bucculatrix ivella
parasite of Exema neglecta
parasite of Exema neglecta
parasite of agromyzid
parasite of Epiblema discretivana
parasite of Tephritis subpura
parasite of Neolasioptera lathami
parasite of agromyzid
parasite of agromyzid
parasite of Bucculatrix ivella
parasite of agromyzid
parasite of Coleomegilla maculata
incidental
general predator
nn
226
Table 2. Continued.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Species!
Habit
*Monomorium floricola (Jerdon)
*Pseudomyrma brunnea F. Smith
*Pseudomyrma pallida F. Smith
*Wasmannia auropunctata (Rogor)
Ichneumonidae
Brachycyrtus pretiosus Cushman
Eiphosoma mexicana Cresson
Labena grallator Say
Temelucha sp.
Trogomorpha trogiformis (Cresson)
Mutillidae
Dasymutilla cypris BI.
Platygasteridae
Platygaster baccharicola (Ashmead)
Trichasis sp.
Pteromalidae
Heteroschema sp.
Sphecidae
Sceliphron caementarium Dru.
Vespidae
Polistes annularis L.
general predator
general predator
parasite of Chrysopa
parasite of Amniscus perplexus
parasite of Oidaematophorus balanotes
general predator
parasite of Neolasioptera lathami
parasite of Neolasioptera lathami
parasite of Exema neglecta
general predator
general predator
'* = Record from DPI collection card file.
The cossid, Prionoxystus piger (Grote),
caused considerable damage to the shrubs
by its stem-boring activity. This was a uni-
voltine species, with moth activity in spring
and larvae present in the stems throughout
the year. It was found only ina very limited,
frost-free area to the south of Miami, sug-
gesting that it may be a tropical, immigrant
species from the Caribbean Islands. It has
been previously reported from Cuba (Grote
1865).
The cochylid, Lorita baccharivora Pogue,
isa multivoltine species that was commonly
encountered from South Carolina to Flor-
ida. Larvae tied terminal and surrounding
leaves together with silken threads to form
tubes in which they lived. This action caused
growth to be arrested, and the growing points
to die, as reported by Diatloff and Palmer
(1988, in press).
The case-bearing chrysomelid Exema ne-
glecta Blatchley, was also commonly en-
countered from South Carolina to Florida.
Both larvae and adults fed on the plant.
PROSPECTS FOR BIOLOGICAL CONTROL
Trirhabda_ bacharidis (W. Haseler, un-
published), Oidaematophorus balanotes (W.
Haseler, unpublished), Aristotelia ivae
(Diatloff and Palmer 1988, in press), Buc-
culatrix ivella (Palmer and Diatloff 1987),
Lorita baccharivora (Diatloff and Palmer
1988, in press), Neolasioptera lathami (Diat-
loff and Palmer 1987), Amniscus perplexus
(Palmer, unpublished), S/aterocoris pallipes
(Palmer, unpublished), Stobaera pallida
(Palmer, unpublished), and /tame varadaria
(Palmer, unpublished) have been proven
host specific and have been introduced into
Australia. Trirhabda bacharidis and A. ivae
have been established in the field in Queens-
land but they have not contributed to ef-
fective control except in localized areas.
Oidaematophorus balanotes and L. bac-
charivora are at present being released and
establishment is anticipated. Bucculatrix
ivella, A. perplexus, and I. varadaria are
undergoing final testing in Australia prior
to their release. Neolasioptera lathami, S.
VOLUME 90, NUMBER 2
pallida, and S. pallipes have not yet been
successfully cultured in the Australian quar-
antine facilities. The remaining monopha-
gous species will be further tested for host
specificity in the future.
The monophagous species were rated by
the formula of Goeden (1983) and also sub-
jectively by the authors, based on their North
American experience with the insects (Ta-
ble 3). The two methods of assessment were
not in close agreement, although both in-
dicated a number of promising species. 47-
niscus perplexus, B. ivella, T. bacharidis, and
O. balanotes were given good scores by both
methods. All 14 species received a score of
>20 by the Goeden formula indicating that
they might be at least partially effective
agents and worthy of further study.
DISCUSSION
In order to find all the insects on the plant,
we found it essential to use both sweeping
and visual inspection. Baccharis halimifolia
is a tall bush growing well above surround-
ing grasses and herbs and therefore can be
swept with little risk that the sample will be
contaminated with arthropods from other
plants. Sweeping proved to be the best
method for capturing small active species
and caterpillars present in low numbers. On
the other hand, it was essential to inspect
the plants visually in order to collect en-
dophages and tightly adhering insects such
as coccids.
Despite differences in sampling proce-
dures and time allocated for survey, the size
of the insect fauna is similar to that found
on B. pilularis (Tilden 1951) and on B. hal-
imifolia and B. neglecta west of the Missis-
sipp1 by Palmer (1987). However, in one
respect, this survey differed from the others;
a much larger number of species of scale
insects was taken, all in Florida. This may
be due in part to Florida’s subtropical cli-
mate and proximity to the Caribbean Is-
lands from which many tropical species have
become established.
The survey emphasized the importance
D7,
Table 3. The potential effectiveness for biological
control of the monophagous species as predicted by
the formula of Goeden (1983) and by the authors’ sub-
jective assessment (with a poor candidate scoring | and
a superior prospect scoring 5).
Authors”
Goeden’s Assess-
Species Formula ment
Amniscus perplexus 47 5
Bucculatrix ivella 45 5
Prionoxystus piger 37 5
Trirhabda bacharidis 45 5
Oidaematophorus balanotes 53 4
Aristotelia ivae 49 3
Lorita baccharivora 51 3
Neolasioptera lathami 47 3
Tephritis subpura 40 3
Itame varadaria 44 2
Slaterocoris pallipes 30 2
Stobaera pallida 41 2
Epiblema discretivana 36 1
Neolasioptera baccharicola 37 1
of searching for endophages, as a very high
proportion of these were monophagous as
indicated also by Palmer (1987). Not only
is there a high probability that an endophage
will be monophagous, but their endopha-
gous habit may protect them from many
general predators and parasites in the coun-
try of release.
The survey also indicated that B. hal-
imifolia harbours a rich insect fauna occu-
pying a diverse range of niches. As B. hal-
imifoliais acommon plant along the eastern
seaboard, it may be ecologically important
to its habitat and to nearby human agricul-
tural endeavours. This is suggested by the
number of species of general predators as-
sociated with it and by the number of ag-
ricultural pests that either feed or seek shel-
ter on it. It may therefore play an important
role as an alternate host for these insects.
ACKNOWLEDGMENTS
We thank the Systematic Entomology
Laboratory (SEL), Agricultural Research
Service, USDA, and the Department of
Entomology, National Museum of Natural
228
History (NMNH) for providing such a fine
service for us and other collectors.
We thank the following taxonomists for
their identifications and advice: R. L. Brown,
Mississippi Entomological Museum (Lepi-
doptera: Tortricidae); E. V. Cashatt, Illinois
State Museum (Lepidoptera: Pterophori-
dae); J. A. Chemsak, UC Berkeley (Coleop-
tera: Cerambycidae); H. Cromroy, Univer-
sity of Florida (Acari: Eriophyidae); D. R.
Davis, NMNH (Lepidoptera: Lyonetiidae);
D. C. Ferguson, SEL (Lepidoptera: Geo-
metridae, Pterophoridae, Pyralidae); R. H.
Foote, SEL (Diptera: Tephritidae); R. J.
Gagné, SEL (Diptera: Cecidomyiidae); R.
D. Gordon, SEL (Coleoptera: Scarabaeidae,
Coccinellidae); T. J. Henry, SEL (Hemip-
tera); R. W. Hodges, SEL (Lepidoptera: Ge-
lechoidea); J. P. Kramer, SEL (Homoptera:
Fulgoridae, Cicadellidae, Membracidae,
Cercopidae); D. R. Miller, SEL (Homop-
tera: Coccoidea); M. Pogue, NMNH (Lep-
idoptera: Cochylidae); R. W. Poole, SEL
(Lepidoptera: Noctuidae); F. C. Thompson,
SEL (Diptera: Syrphidae); T. J. Spilman,
SEL (Coleoptera: Cerambycidae, Elater-
dae); M. B. Stoetzel, SEL (Homoptera:
Aphididae); R. E. White, SEL (Coleoptera:
Chrysomelidae, Cerambycidae); and D. R.
Whitehead, SEL (Coleoptera: Curculioni-
dae).
We also thank the taxonomists at DPI (H.
Denmark, H. Weems, F. Mead, and R.
Woodruff) for allowing us to use host rec-
ords from card files in that collection, for
identifications and for advice.
We also thank G. Diatloff, D. Green, D.
Harbeck, W. Haseler, S. Passoa, and V.
Krischick who collected some of the species
that we have reported.
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QM84007. 623 pp.
Tilden, J. W. 1951. The insect associates of Baccharis
pilularis De Candolle. Microentomology 16: 149-
188.
Wheeler, A. G. 1981. The distribution and seasonal
history of S/aterocoris pallipes (Knight) (Hemip-
tera: Miridae). Proc. Entomol. Soc. Wash. 83: 520-
323:
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 229-243
SPECIES OF AUSTRALIAN TELENOMINAE
(HYMENOPTERA: SCELIONIDAE) OF
A. P. DODD AND A. A. GIRAULT
NORMAN F. JOHNSON
Department of Entomology, The Ohio State University, 1735 Neil Avenue, Columbus,
Ohio 43210
Abstract. —The species of Telenominae described by A. P. Dodd and A. A. Girault with
types deposited in Australian collections are reviewed. The genera Dissolcoides, Neoteleia,
and Platytelenomus are synonymized with Telenomus (new synonymies); Archiphanurus
is synonymized with Paratelenomus (new synonymy). Telenomus ophiusa and T. sac-
charalis are transferred to Paratelenomus (new combinations); Telenomus elpenor and T.
omphale are transferred to Psix (new combinations); Dissolcoides exsertus, D. flavinervus,
Neoteleia punctata, and Platytelenomus planus are transferred to Telenomus (new com-
binations); Telenomus biproruli, T. darwinensis, T. eetion, T. egeria, T. ephyra, T. erigone,
T. obliteratus, T. oecleoides, T. oecleus, T. oedipus, T. oeneus, T. oenone, T. oenopion, T.
ogyges, T. oreas, T. orontes, T. otho, and T. wilsoni are transferred to Trissolcus (new
combinations); Telenomus oaxes is transferred to Embidobia (new combination); and
Telenomus orestes is transferred to Gryon (new combination). Lectotypes are designated
for Dissolcus atriscapus, Trissolcus coriaceus, Telenomus darwinensis, Telenomus eetion,
Telenomus egeria, Telenomus oecleus, Telenomus oedipus, and Telenomus ogyges. The
type material is missing for Phanurus longicornis, P. depressus, Telenomus diemenensis,
and Neotelenomus magniclavatus.
Key Words: _ types, synonymies, combinations
Our knowledge of the Telenominae (Hy-
menoptera: Scelionidae) of Australia is
based primarily upon the pioneering work
of Alan P. Dodd (1895-1981). Several fac-
tors, however, make it difficult to use his
species to place them in the proper context,
at least as it is understood today. I am con-
vinced that the generic concepts within the
Telenominae are in need of significant mod-
ification. It is premature to offer such a re-
published works. Generic concepts within
the subfamily have substantially changed,
and Dodd placed great emphasis upon color
and the relative lengths and widths of an-
tennomeres as diagnostic characters. Solely
on the basis of the original descriptions (no
figures were published), I have found it im-
possible to determine to which present-day
genera Dodd’s species belong and to eval-
uate the genera he described. I thus found
it necessary to examine the types of each
vision, but in the meantime I believe it is
important to indicate how Dodd’s species
fit within the genera presently recognized
(see Masner 1980, Kozlov and Kononova
1983, Johnson 1984), so as to indicate which
type-specimens are relevant for future re-
visionary work at the species level.
I discuss below the telenomine types de-
scribed by Dodd and A. A. Girault that are
deposited in Australian institutions (with
acronyms used in the text in parentheses):
230
Australian National Insect Collection, Can-
berra (ANIC); Department of Primary In-
dustries, Queensland, Indooroopilly (DPIQ);
National Museum of Victoria, Melbourne
(NMV): Queensland Museum, Brisbane
(QM); and the South Australian Museum,
Adelaide (SAM). These include several
species described from Indonesia and Fiji.
A small number of types of Australian
species are deposited in the British Museum
(Natural History) and the National Mu-
seum of Natural History, Washington; the
status of these has been discussed by Masner
(1961) and Masner and Muesebeck (1968).
The species of Australian Scelioninae have
already been treated (see Galloway 1976,
Austin 1981, and Galloway and Austin
1984). The condition of each specimen or
series is briefly summarized. I have cited
verbatim the label data accompanying spec-
imens between quotation marks so as to
assist recognition of the types. The abbre-
viation ““NQ” in the labels stands for North
Queensland, where both Dodd and Girault
conducted much of their field work. The
town of Nelson is now known as Gordon-
vale. The fore wings of slide mounts are
often quite faded, and in many I was unable
to find them although the label stated that
they should be present. The type depository
is indicated following the citation of the
original description for each species. I have
designated lectotypes only for those species
in the genus 7rissolcus in which I am con-
ducting revisionary work. Many of Dodd’s
species were placed in the genera Baeoneu-
rella, Phanurus, and Neotelenomus, names
which were later synonymized with Eumi-
crosoma (the first) and Telenomus. In most
cases the Australian species were not ex-
plicitly transferred to Telenomus; I have ac-
cepted these transfers as having been made
implicitly and have not used the designation
“new combination” for them.
SUBFAMILY TELENOMINAE
Archiphanurus Szabo
See Paratelenomus Dodd.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Baeoneura Foerster
See Eumicrosoma Gahan.
Baeoneurella Dodd
See Eumicrosoma Gahan.
Dissolcoides Dodd
See Telenomus Haliday.
Dissolcus Ashmead
See Telenomus Haliday.
Eumicrosoma Gahan
Eumicrosoma bellum (Dodd)
Baeoneurella bella Dodd, 1913b: 336. QM
Holotype 2 on slide; “TYPE Hy/1628;
Queensland Museum; Baeoneurella bella
Dodd °.” Condition: All tagmata slightly
crushed and broken; antennae detached.
Eumicrosoma elongatum (Dodd),
Baeoneura elongata Dodd, 1913a: 166.
SAM
Holotype @ on slide I11115; ““Baeoneu-
rella elongata Dodd ¢ type.”’ Condition: head
and mesosoma crushed.
Eumicrosoma giraulti (Dodd)
Baeoneura giraulti Dodd, 1913c: 176. SAM
Holotype 2 on slide 11440; ““Baeoneura
giraulti Dodd 2 type; sweeping in forest,
Pentland, NQ, 4th Jun. 13, A. A. Giraulti.”
Condition: head and mesosoma crushed.
Eumicrosoma nigrum (Dodd),
Baeoneurella nigra Dodd, 1914b: 124.
Holotype ¢ on slide 12191; “Baeoneurella
nigra Dodd ° type; 2191.’ Condition: slide
broken in half; cover slip hanging over edge
and cracked; head detached, crushed; meso-
soma and metasoma slightly crushed.
VOLUME 90, NUMBER 2
Eumicrosoma pulchrum (Dodd),
Baeoneurella pulchra Dodd, 1914b: 124.
SAM
Holotype 2 on slide 12190; ‘“Baeoneurella
pulchra Dodd 2 type, B. giraulti Dodd;
2190.” Condition: mounted on slide with a
2 of E. giraulti; holotype of E. pulchrum is
closest to the center of the cover slip; head
and mesosoma crushed.
Neoteleia Dodd
See Telenomus Haliday.
Neotelenomus Dodd
See Telenomus Haliday.
Paratelenomus Dodd
Paratelenomus bicolor (Dodd)
Telenomus bicolor Dodd, 1914c: 251. SAM
Holotype 2 on slide 111169; *“Telenomus
bicolor Dodd 2 type.” Condition: head and
mesosoma crushed.
This species, although remarkable for its
color pattern (black head and golden body),
is clearly congeneric with the type species
of the genus 4rchiphanurus Szabo, viz. A.
graeffei (Kieffer). Thus, the last valid telen-
omine genus described by Szabo (1975) falls
as ajunior synonym of Paratelenomus Dodd
(new synonymy).
Paratelenomus ophiusa (Dodd),
NEw COMBINATION
Telenomus ophiusa Dodd, 1913d: 84. SAM
Holotype 2 on slide I1 1176; ““Telenomus
ophiusa Dodd 2 type; 11176." Condition:
tagmata separated, all badly crushed.
Paratelenomus saccharalis (Dodd),
NEw COMBINATION
Telenomus saccharalis Dodd, 1914f: 293.
QM
Syntype 4, 2 on slide; ‘““Telenomus sac-
charalis Dodd, 6 2 type, From Pentatomid
eggs on sugarcane, Java; Hy 2059; TYPE.”
231
Condition: 2 with all tagmata separated,
crushed; ¢ has slipped with the balsam over
and on top of the edge of the cover slip and
is only partially covered by the medium.
Phanuromyia Dodd
Phanuromyia rufobasalis Dodd
Phanuromyia rufobasalis Dodd, 1914g: 121.
SAM
Holotype 2 on slide I11027; *‘Phanuro-
myia rufobasalis Dodd 2 type.’ Condition:
head and mesosoma crushed.
This species clearly represents the same
species group as genus /ssidotelenomus Pé-
lov, and possibly both 4radoctonus Masner
and Phlebiaporus Kozlov. A short time ago
I would have freely synonymized all under
the name 7e/enomus. I now believe that this
species group will warrant generic recogni-
tion and therefore I have elected not to
transfer P. riufobasalis.
Platytelenomus Dodd
See Telenomus Haliday.
Psix Kozlov and Lé
Psix elpenor (Dodd),
New COMBINATION
Telenomus elpenor Dodd, 1914h: 4. SAM
Holotype 2 with mesosoma and metaso-
ma on point; ““Telenomus elpenor Dodd 2
type.” Condition: good. Head, antennae,
fore wing on slide I1 1095; **Telenomus el-
penor Dodd ¢ type; 111095.” Condition:
head badly crushed, cover slip cracked over
head.
Psix glabriscrobus (Girault)
Telenomus glabriscrobus Girault, 1926b:
138. ANIC
See Johnson and Masner, 1985: 46-47.
Psix olympus (Dodd)
Telenomus olympus Dodd, 1913c: 166.
SAM
Holotype 2 on slide 11418; ““Telenomus
232
olympus Dodd ¢ type, sweeping on edge of
jungle, Nelson, NQ, 5.iv.13 (A. P. Dodd).”
Condition: badly crushed. Also under the
cover slip is a specimen of Telenomus. The
recognition of the specimen of Psix as rep-
resenting 7. olympus is based upon another
specimen in ANIC identified by Dodd (see
Johnson and Masner 1985: 49).
Psix omphale (Dodd),
New ComBINATION
Telenomus omphale Dodd, 1913c: 166.
SAM
Holotype 2 on point, ““Telenomus om-
phale Dodd, 2 type.’ Condition: good; an-
tennae, both without radicle, forewing on
slide 11419, **Telenomus omphale Dodd, ?
type, antennae, forewings, From Pentatom-
id eggs in forest, Nelson, NQ, Apr. 13, A.
P. Dodd.”
Telenomus Haliday
Telenomus acares Johnson
Neotelenomus minimus Dodd, 1913c: 172,
[preoccupied by minimus Ashmead,
1893]. SAM
Telenomus acares Johnson, 1984: 6 (re-
placement name).
Holotype 2 on slide 11433: ‘*Neoteleno-
mus minimus Dodd 2 type; On window,
Nelson, NQ, 2nd.x11.12, A. P. Dodd.” Con-
dition: crushed.
Telenomus aegeus Dodd
Telenomus aegeus Dodd, 1914g: 124. SAM
Holotype 2 on slide I1 1031; ““Telenomus
aegeus Dodd 2 type.” Condition: head and
mesosoma crushed.
Telenomus aegicerophilus (Dodd)
Neotelenomus aegicerophilus Dodd, 1914h:
11. SAM
Holotype 2 on slide I11141; “Neotelen-
omus aegicerophilus Dodd @ type; 11141.”
Condition: crushed, especially mesosoma.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Telenomus ajax Dodd
Telenomus ajax Dodd, 1914g: 125. SAM
Holotype 2 on slide I1 1032; *“Telenomus
ajax Dodd 2 type.” Condition: head and
mesosoma badly crushed.
Telenomus anthereae (Dodd)
Neotelenomus anthereae Dodd, 1913c: 171.
SAM
One ¢, two 2 syntypes on slide 11429;
*“Neotelenomus anthereae Dodd 4, 2 types;
From egg of Antherea janetta, Nelson, NQ,
May, 13, A P Dodd.” Condition: all crushed;
male genitalia exserted and clearly visible.
Nixon (1937) synonymized Neoteleno-
mus with Telenomus on the basis of his con-
viction that the characters defining a genus
should be applicable to both sexes. In this
case, the only character used to distinguish
Neotelenomus was the 10-merous female
antenna. Nixon’s interpretation that, aside
from this character, Neotelenomus was in-
distinguishable from Telenomus was based
not On an examination of the type species,
N. anthereae, but on Dodd’s original de-
scription and his knowledge of other species
with the same reduction in antennomeres.
I can now confirm that Nixon’s synony-
mization is correct.
Telenomus atratus Johnson
Neotelenomus niger Dodd, 1913c: 172
[preoccupied by niger (Dodd), 1913c:
158]. SAM
Telenomus atratus Johnson, 1984: 12 (re-
placement name).
Twelve syntype 2 on card (with 3 chal-
cidoids); ““Kuranda, Qld., Mch 04, F. P.
Dodd; Neotelenomus niger Dodd 2 types.”
Condition: generally good, but very dirty.
One 2 syntype on slide 11432; ““Neotelen-
omus niger Dodd ¢ type; Kuranda, NQ,
March, 04, F. P. Dodd.” Condition: crushed.
Telenomus australis (Dodd)
Neotelenomus australis Dodd, 1913d: 86.
SAM
VOLUME 90, NUMBER 2
Holotype 2 on slide I11140; ““Neotelen-
omus australis Dodd ° type.”’ Condition:
mesosoma and head crushed and broken.
Telenomus beatus (Dodd)
Neotelenomus beatus Dodd, 1913d: 85.
SAM
Holotype 2 on slide I11142; ‘“Neotelen-
omus beatus Dodd ¢ type.’ Condition: end
of slide broken off; specimen badly crushed.
Telenomus caesaris (Girault)
Neotelenomus caesaris Girault, 1939: 149.
ANIC
Syntypes on card; ‘4, 2 Neotelenomus
caesaris Gir, Types.”’ Condition: four 2, one
4in good condition; also with bits and pieces
of several broken specimens. There 1s also
a second unlabelled pin with five ? and one
6 that appear to belong to the same series.
Telenomus carnifex Johnson
Neotelenomus ovivorus Dodd, 1913c: 172
[preoccupied by ovivorus (Rondan1),
1870]. SAM
Telenomus carnifex Johnson, 1984: 7 (re-
placement name).
Syntype 4, 2 on slide 11430; ‘*Neotelen-
omus ovivorus Dodd, 4 and 2 types, Nelson,
May, 13, Dodd.” Condition: ¢ with head
and mesosoma crushed; 2 entirely crushed.
Telenomus corniger Johnson
Phanurus longicornis Dodd, 1913c: 160,
[preoccupied by /ongicornis Ashmead,
1901].
Telenomus corniger Johnson, 1984: 8 (re-
placement name).
Type missing from SAM. The unit tray
refers to slide 11409, but it is missing from
the slide collection.
Telenomus depressus (Dodd)
Phanurus depressus Dodd, 1914h: 8.
Type missing from SAM.
233
Telenomus diemenensis Dodd
Telenomus diemenensis Dodd, 1914g: 123.
Type missing from SAM. Unit tray refers
to slide I11030; this is missing from the
slide collection and the label on the inside
of the box’s lid questions whether the slide
was ever received.
Telenomus doddi Johnson
Telenomus giraulti Dodd, 1914d: 161.
[preoccupied by giraulti (Dodd), 1913c].
QM
Telenomus doddi Johnson, 1984: 9 (replace-
ment name).
Holotype 2 on slide; “TYPE Hy/2057;
Queensland Museum; Scelionid, Teleno-
mus giraulti Dodd 2.’ Condition: head de-
tached; mesosoma on its side, slightly
crushed laterally.
Telenomus eleleus Dodd
Telenomus eleleus Dodd, 1914h: 5. SAM
Holotype ¢ on slide I1 1170; **Telenomus
eleleus Dodd 2 type; I11170.° Condition:
crushed, tagmata separated.
Telenomus emersoni Girault
See Telenomus olsenni Johnson.
Telenomus endymion Dodd
Telenomus endymion Dodd, 1914h: 6. SAM
Holotype 2 on point; “Telenomus en-
dymion Dodd @ type.” Condition: good.
Antennae (and possibly also the fore wings)
on slide 111096; “I11096; Telenomus en-
dymion Dodd ¢ type.” Condition: good;
radicles still attached to head.
Telenomus eteocles Dodd
Telenomus eteocles Dodd, 1914h: 5. SAM
Holotype 2 on slide 111171; ““Telenomus
eteocles Dodd 2 type; 111171.” Condition:
tagmata separated, head and mesosoma
crushed.
Telenomus exsertus (Dodd),
NEw COMBINATION
Dissolcoides exsertus Dodd, 1913a: 179.
SAM
Holotype 2 with mesosoma and meta-
soma on point; “Dissolcoides exsertus Dodd
2 type; Pentland, Queensland.”’ Condition:
deeply embedded in glue, barely visible.
Head on slide I1 1059; *“‘Dissolcoides exsert-
us Dodd 2 type; head, antennae, forewings;
sweeping grass in forest, Pentland, NQ, 15th
[?] Jan 13, A. A. Girault.’” Condition: head
crushed, A10-A11 of right antenna missing;
left antenna detached, near edge of cover
slip.
Dodd apparently erected the genus Dis-
solcoides to accommodate what he per-
ceived was a species that combined impor-
tant characters of several of Ashmead’s
genera (1893). With the head detached it is
now difficult to visualize the habitus of this
wasp that so struck his attention, but it ap-
pears to me to be a fairly typical species of
Telenomus (new synonymy).
Telenomus eximius (Dodd)
Neotelenomus eximius Dodd, 1914b: 121.
SAM
Holotype 2 on point; ““Neotelenomus ex-
imius Dodd 2 type; 12186.” Condition: body
deeply embedded in glue. Antennae and
possibly fore wings on slide 12186; “‘Neo-
telenomus eximius Dodd ° type: forewings,
antennae.” Condition: one antenna crushed.
Telenomus flavescens Dodd
Telenomus flavescens Dodd, 1914h: 4. SAM
Holotype ¢ on slide I1 1172; *“Telenomus
flavescens Dodd ¢ type.” Condition: head
separated from body; mesosoma and head
crushed.
Telenomus flavineryus (Dodd)
New ComBINATION
Dissolcoides flavinervus Dodd, 1914c: 253.
SAM
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Holotype 2 on point; “Dissolcoides flav-
inervus Dodd 2 type; Herbert R.”’ Condi-
tion: deeply embedded in glue, otherwise
good. Antennae (and possibly fore wings)
on slide 111060; **Dissolcoides flavinervus
Dodd 2 type; antennae forewings.”
Condition: antennae broken off from head
beyond radicles, one clava slightly crushed.
Telenomus giraulti (Dodd)
Phanurus giraulti Dodd, 1913c: 159. SAM
Two syntypes, 4 and 2 on separate slides,
both coded 11403; ““Phanurus giraulti Dodd
é type, Nelson, NQ, 13.11.13, A P Dodd”;
“‘Phanurus giraulti Dodd 2 type, Forest Nel-
son, NQ, 10.vii.12, A A Girault.’’ Condi-
ton: both with head and mesosoma crushed.
Telenomus giraulti Dodd, 1914d
See Telenomus doddi Johnson.
Telenomus gloriosus Dodd
Telenomus gloriosus Dodd, 1913d: 84. SAM
Holotype 2 on slide 111173; ‘*Telenomus
gloriosus Dodd 2 type; 11173.’ Condition:
head and mesosoma crushed; A8-A11 of
both antennae missing.
Telenomus hackeri (Dodd)
Phanurus hackeri Dodd, 1913b: 337. QM
Holotype 2 on slide; “TYPE Hy/1629;
Queensland Museum; Phanurus_hackeri
Dodd 2.” Condition: tagmata separated, all
badly crushed.
Telenomus hilli (Dodd)
Phanurus hilli Dodd, 1914b: 119. SAM
One 2, 3 2syntypes on slide I2 180; ““Phan-
urus hilli Dodd 4 + 2 types, 2180. Con-
dition: all badly crushed.
Telenomus javensis Dodd
Telenomus javensis Dodd, 1914e: 163. QM
Syntype 2 on point; “TYPE Hy/2060; Te-
lenomus javensis Dodd 2 type.” Condition:
deeply embedded in glue. Syntype 2 on slide
VOLUME 90, NUMBER 2
[together with the holotype of Telenomus
vandergooti Dodd]; ““TYPE/2060 2061;
Scelionid, Queensland Museum, 2 Teleno-
mus javensis D. 2060 T. vandergooti D. 2
2061.” Condition: mesosoma crushed; head
is separated and has slipped to the edge of
the cover slip beneath a drop of balsam.
Telenomus laticeps (Dodd)
Neotelenomus laticeps Dodd, 1914h: 10.
SAM
Holotype 2 on slide I1 1143; ““Neotelen-
omus laticeps Dodd 2 type; 111143.’ Con-
dition: mesosoma and metasoma_ badly
crushed; head has slipped beyond the edge
of the cover slip; one antenna, with A2-A9
is beyond the head, along the edge of the
slide; second antenna not found.
Telenomus leai (Dodd)
Neotelenomus leai Dodd, 1913c: 172. SAM
Holotype 2 on slide 11431; *‘Neoteleno-
mus leai Dodd 2 type; King Island, Bass
Strait, Tasmania, A. M. Lea.” Condition:
head and apex of metasoma broken off; all
badly crushed.
Telenomus longicornis (Dodd)
See Telenomus corniger Johnson.
Telenomus longicorpus (Dodd)
Phanurus longicorpus Dodd, 1913c: 160.
SAM
Two syntype 2 mounted on separate slides
both coded 11406. ““Phanurus longicorpus
Dodd 2 type, sweeping forest, Nelson, Feb
[other handwriting indistinct].’’ Condition:
head detached, crushed; mesosoma crushed.
“Phanurus longicorpus Dodd 2 type, sweep-
ing forest, Nelson, NQ, 13.11.12, A A Gi-
rault.”” Condition: head detached, all tag-
mata crushed.
Telenomus longipennis (Dodd)
Phanurus longipennis Dodd, 1913c: 160.
SAM
235
Holotype 2 on slide 11407; ‘““Phanurus
longipennis Dodd 2 type, sweeping in forest,
Ingham, 14.1.13, A. P. Dodd.’ Condition:
head detached, mesosoma crushed.
Telenomus magniclayatus (Dodd)
Neotelenomus magniclavatus Dodd, 1914b:
122.
Type missing from SAM. It should be
mounted on slide 12187, but this is missing
from the slide collection.
Telenomus minimus (Dodd)
See Telenomus acares Johnson.
Telenomus montanus (Dodd)
Phanurus montanus Dodd, 1913c: 159.
SAM
Holotype 2 on slide 11404, “Phanurus
montanus Dodd ¢ type [with specific name
fumipennis crossed out], on window, Her-
berton, NQ (3000 feet), 28.xu1.11, A. A. Gi-
rault.”” Condition: mesosoma crushed.
Telenomus necopinatus (Dodd)
Phanuromyia necopinata Dodd, 1916: 32.
Holotype 2, mesosoma and metasoma on
point; “Phanuromyia necopinata Dodd °
type.” Condition: head missing. Antennae,
fore wing on slide I11158; ““Phanuromyia
necopinata Dodd 2 type.’ Condition: rad-
icles missing.
Telenomus nelsonensis (Dodd)
Phanurus nelsonensis Dodd, 1913c: 160.
SAM
Holotype 2 on slide I1405; ‘““Phanurus
nelsonensis Dodd 2 type, sweeping in forest,
Nelson, NQ, 14.vi.12, A A Girault.”’ Con-
dition: good.
Telenomus niger (Dodd)
Phanurus niger Dodd, 1913c: 158. SAM
Holotype 2 on slide 11402; **“Phanurus ni-
ger Dodd 2 type, Nelson, NQ, 24th Dec, 12,
236
on window of laboratory porch.” Condi-
tion: head and mesosoma crushed.
Telenomus niger (Dodd), 1913c
See Telenomus atratus Johnson.
Telenomus nigricorpus (Dodd)
Phanurus_ nigricorpus Dodd, 1913c: 160.
SAM
Holotype 2 on slide 11408; **Phanurus ni-
gricorpus Dodd 2 type, Nelson, Jan 12, A
A Girault.”’ Condition: crushed, barely cov-
ered by balsam.
Telenomus ocnus (Dodd)
Telenomus ocnus Dodd, 1914b: 120. SAM
Holotype 2 on slide 12183; ““Telenomus
ocnus Dodd 2 type, head, antennae, fore-
wings, 2183.” Condition: head crushed;
mesosoma and metasoma missing.
Telenomus odyssea (Dodd)
Telenomus odyssea Dodd, 1913c: 162. SAM
Holotype 2 on slide 11410; **Telenomus
odyssea Dodd 2 type, sweeping in forest,
Nelson, 3.1x.12, A. A. Girault.”” Condition:
head detached, mesosoma crushed.
Telenomus oeagrus Dodd
Telenomus oeagrus Dodd, 1913c: 163. SAM
Holotype 2 on slide 11411: ““Telenomus
oeagrus Dodd, ? type, sweeping jungle along
streamlet, Babinda, NQ, 26.x.11, A. A. Gi-
rault.”” Condition: mesosoma crushed.
Telenomus oechalia Dodd
Telenomus oechalia Dodd, 1913d: 83. SAM
Holotype 2 on slide I1 1175; ““Telenomus
oechalia Dodd ¢ type.” Condition: badly
crushed.
Telenomus oeta Dodd
Telenomus oeta Dodd, 1914c: 252. SAM
Holotype 2 on slide I1 1174; **Telenomus
oeta Dodd 2 type.” Condition: head de-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tached, all tagmata crushed, mesosoma es-
pecially so; balsam barely covers specimen.
Telenomus olsenni Johnson
Telenomus emersoni Girault, 1932: 6 [in
Gordh et al., 1979: 298; preoccupied by
emersoni (Girault), 1916]. QM
Telenomus olsenni Johnson, 1984: 12 (re-
placement name).
One wing, one antenna on slide; ““Telen-
omus emersoni Gir. Type °.’’ Condition:
good; remainder of specimen missing.
Telenomus ophion Dodd
Telenomus ophion Dodd, 1913c: 167. SAM
One syntype specimen on card, sex un-
certain; ““Telenomus ophion Dodd 4 type.”
Condition: fair, antennae, forewings miss-
ing. Slide 11420 with head of 2, one antenna
attached, complete; left antenna with radi-
cle, A10O-A11 missing; head of é with one
antenna attached, second detached: pro-
pleuron, fore legs, a fore wing and hind wing
present; “Telenomus ophion Dodd, 4, 2
types; From Pentatomid eggs, Nelson, NQ,
May, 12, A. A. Girault.”
Telenomus opis Dodd
Telenomus opis Dodd, 1913d: 84. SAM
Holotype 2 on slide I1 1177; **Telenomus
opis Dodd 2 type; 11177.’ Condition: head,
propleura, prosternum and fore legs sepa-
rated; entire body crushed.
Telenomus orithyia Dodd
Telenomus orithyia Dodd, 1913c: 180. SAM
Holotype 2 on slide 11451; ““Telenomus
orithyia Dodd 2 type; sweeping in jungle,
Nelson, NQ, 15.v.13, A. P. Dodd.” Con-
dition: head and mesosoma badly crushed.
Telenomus ormenis Dodd
Telenomus ormenis Dodd, 1913c: 181. SAM
Holotype 2 on slide 11452; ““Telenomus
ormenis Dodd 2 type; sweeping on edge of
jungle, Kuranda, NQ, 18.v.13, A P Dodd.”
VOLUME 90, NUMBER 2
Condition: head detached, mesosoma
crushed.
Telenomus orodes Dodd
Telenomus orodes Dodd, 1913c: 181. SAM
Holotype 2 on slide 11453; *“Telenomus
orodes Dodd 2 type; sweeping on edge of
jungle, Kuranda, NQ, 18.v.13 (A P Dodd).”
Condition: crushed.
Telenomus orpheus Dodd
Telenomus orpheus Dodd, 1913c: 181. SAM
Holotype 2 on slide 11454; *“Telenomus
orpheus Dodd 2 type; sweeping foliage of
lantana, Mackay, NQ, 11.x.11 (A A Gi-
rault).”’ Condition: crushed.
Telenomus osiris Dodd
Telenomus osiris Dodd, 1913c: 180. SAM
Holotype 2 on slide 11450; **Telenomus
osiris Dodd 2 type; sweeping forest and jun-
gle, Nelson, NQ, 3.1x.12, A. A. Girault.”
Condition: tagmata detached, mesosoma
broken, head and anterior part of mesosoma
have slipped beyond the edge of the cover
slip.
Telenomus ossa Dodd
Telenomus ossa Dodd, 1914b: 119. SAM
Holotype 2 on slide 12181; *“Telenomus
ossa Dodd 2 type; 2181. Condition:
crushed.
Telenomus ovivorus (Dodd)
See Telenomus carnifex Johnson.
Telenomus oxycareni Girault
Telenomus oxycareni Girault, 1934: 2
[Gordh et al. 1979: 307]. QM
Two syntype 2 (with identical labels);
“Telenomus oxycareni Gir., 2, Type.”’ Con-
dition: one 2 in good condition; second with
head missing and mesosoma crushed.
Telenomus pallidicornis (Dodd)
Neotelenomus pallidicornis Dodd, 1913d:
86. SAM
238i,
Holotype 2, mesosoma and metasoma on
point; ‘“‘Neotelenomus pallidicornis Dodd &
type; Cairns.”’ Condition: deeply embedded
in glue. Head, fore wing on slide I11147;
“Neotelenomus pallidicornis Dodd 2 type
head, forewings.” Condition: head crushed,
fore wings not visible.
Telenomus pallidithorax (Dodd)
Neotelenomus pallidithorax Dodd, 1914h:
10. SAM
Holotype 2 on slide I11149; **Neotelen-
omus pallidithorax Dodd 2 type; 111149.”
Condition: crushed.
Telenomus pallidiventris (Dodd)
Neotelenomus pallidiventris Dodd, 1913d:
86. SAM
Holotype 2 on slide I11146; ‘‘Neotelen-
omus pallidiventris Dodd 2 type.” Condi-
tion: head and mesosoma crushed; speci-
men very pale.
Telenomus parvulus (Dodd)
Neotelenomus parvulus Dodd, 1914h: 12.
SAM
Holotype 2 on slide 111144; **Neotelen-
omus parvulus Dodd 2 type; I1 1144. Con-
dition: head detached; it and mesosoma
crushed; right antenna missing.
Telenomus planus (Dodd),
New COMBINATION
Platytelenomus planus Dodd, 1914a: 126.
SAM
Holotype 2: mesosoma and metasoma on
point; “‘Platytelenomus planus Dodd 2 type;
Cairns.’ Condition: good. Head, fore wing
on slide 111072; “Platytelenomus planus
Dodd ¢ type, head, forewings.’ Condition:
head slightly crushed, fore wing not visible.
After having examined the type species,
I consider the genus Platytelenomus Dodd
to be a junior synonym of Telenomus (new
synonymy). 7el/enomus planus 1s closely re-
lated to the species I have grouped together
238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
in the floridanus species group as indicated
by the elongate, depressed body and, more
importantly, the elongate clavomeres
(Johnson 1984). All species in these group,
so far as is now known, are parasites of the
eggs of lygaeids. Platytelenomus, when cor-
rectly understood, is not closely related to
the depressed species of Telenomus that
parasitize the flattened eggs of various moths
(see Fergusson 1983). If at some time in the
future it is thought appropriate to recognize
the floridanus group as a genus, the names
Hemisius Westwood and Dissolcus Ash-
mead have priority.
Telenomus pseudoclavatus (Dodd)
Neotelenomus pseudoclavatus Dodd: 1913d:
87. SAM
Holotype 2 on slide I1 1145; ‘‘Neotelen-
omus pseudoclavatus Dodd 2 type.” Con-
dition: head detached; all tagmata crushed.
Telenomus pulcherrimus Dodd
Telenomus pulcherrimus Dodd, 1914b: 121.
SAM
Holotype 2 on slide 12185; ““Telenomus
pulcherrimus Dodd 2 type 2185.” Condi-
tion: mesosoma, head broken.
Telenomus pulchricornis (Dodd)
Neotelenomus pulchricornis Dodd, 1914h:
9. SAM
Holotype 2 on slide I1 1148; ““Neotelen-
omus pulchricornis Dodd 2 type; 111148.”
Condition: head detached; all tagmata, es-
pecially mesosoma, crushed.
Telenomus punctatus (Dodd),
NEw COMBINATION
Neoteleia punctata Dodd, 1913a: 169. SAM
Holotype 6, mesosoma and metasoma on
point; “Neoteleia punctata Dodd ¢ type;
Cairns.” Condition: good. Head on slide
111070; ‘‘Neoteleia punctata Dodd 4 type,
head, forewings.” Condition: head crushed;
fore wings not visible.
This is a fairly large species that may be
most closely related to Phanuromyia. How-
ever, until freshly collected material is iden-
tified and the cephalic characters can be
carefully examined, it falls within the, ad-
mittedly, very broad definition of Teleno-
mus. I prefer to synonymize Neote/eia under
that name (new synonymy) than to main-
tain it as a distinct genus. This species can
probably be identified by its large size,
sculptured frons, presence of episternal fo-
veae, very elongate T2, and the elongate
basal flagellomeres.
Telenomus sidneyi Girault
Telenomus sidneyi Girault, 1932: 5 [Gordh
et al. 1979: 297]. QM
One leg, two fore wings on slide; “Scelion-
id, Type 2, Telenomus sidneyi Gir. wing
{remaining handwriting unclear]. The rest
of the specimen is missing.
Telenomus simulans (Dodd)
Neotelenomus simulans Dodd, 1914h: 11.
SAM
Holotype 2 on slide I11150; ‘‘Neotelen-
omus simulans Dodd @ type; I1 1150. Con-
dition: head detached, slightly broken;
mesosoma crushed.
Telenomus spodopterae Dodd
Telenomus spodopterae Dodd, 1914e: 164.
QM
Four syntype 2 on slide; “TY PE Hy/2062;
Queensland Museum; Telenomus spodop-
terae Dodd 2.”’ Condition: two 2 with meso-
soma broken, otherwise good: one 2 with
head widely separated from mesosoma,
metasoma also detached but close to meso-
soma, mesosoma broken; one 2 with meso-
soma crushed, head and metasoma widely
separated from mesosoma; clavomeres of
all specimens distorted.
Telenomus vandergooti Dodd
Telenomus vandergooti Dodd, 1914e: 164.
QM
VOLUME 90, NUMBER 2
Holotype 2 on slide [together with syntype
of Telenomus javensis Dodd]; “TY PE/2060
2061; Scelionid, Queensland Museum, 2
Telenomus javensis D. 2060 T. vandergooti
D. 2 2061.” Condition: tagmata detached,
head and mesosoma crushed.
Trissolcus Ashmead
Trissolcus atriscapus (Girault)
Dissolcus atriscapus Girault, 1926a: 1
[Gordh et al. 1979: 200]. DPIQ
Lectotype (here designated) 2 on point;
“Dissolcus atriscapus Gir. Type 42.”° Con-
dition: head missing; most of the second
specimen on the point has been lost, only
legs remain glued to the tip.
Trissolcus beenleighi (Girault)
Dissolcus beenleighi Girault, 1932: 5 [in
Gordh et al. 1979: 297]. QM
Holotype 2 on point; “Dissolcus been-
leighi Gir., 2, Type; 29.x11.1925, Beenleigh,
Forest.” Condition: metasoma detached;
antennae and wings from left side of body
missing.
Trissolcus biproruli (Girault),
New ComBINATION
Telenomus biproruli Girault, 1926b: 137.
QM
Holotype 2 on point: “7e/enomus bipro-
ruli Gir., 2, Type.’ Condition: good.
Trissolcus coriaceus Dodd
Trissolcus coriaceus Dodd, 1915: 451. SAM
Lectotype 2 (here designated) and para-
lectotype 2 on point; “Trissolcus coriaceus
Dodd 2 types.”’ Condition: lectotype at apex
of point with head and anterior half of
mesosoma embedded in glue; paralectotype
with mesosoma broken between mesotho-
rax and metathorax; otherwise both in good
condition. Two antennae on slide I5177;
“Trissolcus coriaceus 2 type.”
239
Trissolcus darwinensis (Dodd),
New COMBINATION
Telenomus darwinensis Dodd, 1914h: 7.
SAM
Lectotype 4, paralectotype 2 on point:
“Telenomus darwinensis 2, 4 types.”’ Con-
dition: good. Male antennae, female head
and antennae on slide I1 1099; **Telenomus
darwinensis Dodd 4, 2 types.’” Condition:
head badly crushed, male antennae with
scape broken, still attached to head.
Trissolcus eetion (Dodd),
NEw COMBINATION
Telenomus eetion Dodd, 1914h: 3. SAM
Lectotype 2 on point; ““Telenomus eetion
Dodd 2 type.” Condition: dirty, otherwise
good; both antennae present. Paralectotype
male, female antennae, fore wing on slide
111093; “111093, Telenomus eetion Dodd
42 types.” Condition: female antenna good;
male antenna broken.
Trissolcus egeria (Dodd),
NEw COMBINATION
Telenomus egeria Dodd, 1914h: 4. SAM
Lectotype 2 ““Telenomus egeria Dodd 2
type.’ Condition: head missing. Paralecto-
type 2 on slide 111094; ““Telenomus egeria
Dodd ° type.” Condition: head crushed and
broken, mesosoma crushed, A6—A1 1 of right
antenna, A8-A11 of left antenna missing.
Trissolcus ephyra (Dodd),
NEw COMBINATION
Telenomus ephyra Dodd, 1914h: 7. SAM
Holotype 2 on point; *“Telenomus ephyra
Dodd 2 type.” Condition: body deeply
embedded in glue. Antennae, fore wings on
slide 111097; ‘“‘Telenomus ephyra Dodd ¢
type; 111097.” Condition: antennae crushed,
without radicles.
Trissolcus erigone (Dodd),
New ComBINATION
Telenomus erigone Dodd, 1914h: 8. SAM
240
Holotype 2 on point; ““Telenomus erigone
Dodd @ type.” Condition: body deeply
embedded in glue. Antennae (and possibly
the fore wings) on slide 111098; ‘*Teleno-
mus erigone Dodd ¢ type; I11098.” Con-
dition: radicles missing (still attached to
head), antennae slightly crushed.
Trissolcus euander (Dodd)
Telenomus euander Dodd, 1914h: 7. SAM
Holotype 2 mesosoma and metasoma on
point; ““Telenomus euander Dodd @ type.”
Condition: good. Head, antennae (and pos-
sibly fore wings) on slide 111092; **Telen-
omus euander Dodd 2 type; 111092.” Con-
dition: head crushed.
Trissolcus flaviscapus Dodd
Trissolcus flaviscapus Dodd, 1916: 32. SAM
Holotype 2 on point; “Trissolcus flavis-
capus Dodd ¢ type.”’ Condition: fore wings
missing, otherwise good. Antenna on slide
111180; “Trissolcus flaviscapus Dodd 2 type;
11180.” Condition: broken off from head
above radicle.
Trissolcus obliteratus (Dodd),
New ComMBINATION
Telenomus obliteratus Dodd, 1914g: 122.
SAM
Holotype 2 on point; ““Telenomus obliter-
atus Dodd 2 type.” Condition: only one hind
wing remaining, left antenna broken off just
above radicle, right antenna with A2-A11
missing, otherwise good. Slide 11029 with
nothing visible (it may hold the wings, but
I could not find them). The label in the unit
tray states that the type of this species was
not received. I found the specimen in the
tray with Trissolcus oecleoides.
Trissolcus oecleoides (Dodd),
New ComBINATION
Telenomus oecleoides Dodd, 1914g: 122.
SAM
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Holotype 2 on point, “Telenomus oec-
loides [sic] Dodd 2 type.”’ Condition: good.
Antennae (without radicles) on slide I1 1028,
*Telenomus oecloides [sic] Dodd @ type.”
Trissolcus oecleus (Dodd),
New COMBINATION
Telenomus oecleus Dodd, 1913c: 163. SAM
Lectotype 4 (here designated): ‘‘Teleno-
mus oecleus Dodd type.’? Condition:
mounted on point, A7—-A12 ofleft antennae,
A10-A12 of right antenna missing; lower
half of body, most of head embedded in
glue. One paralectotype 2; ““Telenomus oec-
leus Dodd type.” Condition: mounted on
point, antennae, wings missing; deeply
embedded in glue. The paralectotype, al-
though clearly labelled as a type by Dodd,
is not conspecific with the male. The choice
of the lectotype is based upon a series of
specimens in DPIQ identified by Dodd as
T. oecleus that are conspecific with the male.
There are also two slides with the code 11412
in SAM that bear labels identifying them as
types. One has a fore wing and two complete
male antennae and bears the label ““Telen-
omus oecleus Dodd, 4 type, forewing, an-
tenna, reared from Pentatomid eggs, Ku-
randa, NQ, 3/ix/04, F P Dodd”; the second
has the label ‘“‘Telenomus oecleus Dodd, 2
type, forewings and antennae, sweeping edge
of jungle, Kuranda, NQ, 20.x1.12, A P
Dodd,” presumably from the point-made
female.
Trissolcus oedipus (Dodd),
New ComBINATION
Telenomus oedipus Dodd, 1913c: 164. SAM
Three specimens glued to a card; “Ho-
bart, Tas: Lea; Hobart, Tas: Lea; Teleno-
mus oedipus Dodd 2 types.’ Condition: |
2 lectotype (here designated) on left (viewed
from above with pin at bottom) with line
beneath, in good condition; two paralecto-
types: 1 2 in middle, metasoma missing; |
specimen on right, probably a 2, head miss-
VOLUME 90, NUMBER 2
ing. One 2 paralectotype on slide I1413:
“Telenomus oedipus Dodd 2 type, Hobart,
Tasmania, A M Lea.” Condition: head and
mesosoma badly crushed.
Trissolcus oeneus (Doda),
NEw COMBINATION
Telenomus oeneus Dodd, 1913c: 164. SAM
Holotype 2 on card; “‘King I, Tas: Lea,
Telenomus oeneus Dodd, 2 type.” Condi-
tion: body almost completely covered by
glue; mesosoma broken, mesonotum cov-
ering head; foretibia and tarsus, A2-A9 of
one antenna, A7-A11 of another on slide
11414: ““Telenomus oeneus Dodd, & type,
antenna, forewings, King Is., Bass Strait, A.
M. Lea.”
Trissolcus oenone (Dodd),
NEw COMBINATION
Telenomus oenone Dodd, 1913c: 165. SAM
Holotype 2 on card, “Cairns district, A.
M. Lea, Telenomus oenone Dodd, ¢ type.”
Condition: covered in glue, antennae, fore-
wing on slide 11415, label: ‘““Telenomus oe-
none Dodd, 2 type, forewing antennae,
Cairns district, NQ, A M Lea.”
Trissolcus oenopion (Dodd),
NEw COMBINATION
Telenomus oenopion Dodd, 1913c: 165.
SAM
Holotype 2 on slide 11416; ““Telenomus
oenopion Dodd ¢ type forewing, antenna,
From foliage of a lemon tree, Roma, Q..,
6.x.11, A. A. Girault.”” Condition: one an-
tennae, broken, and distal half of fore wing
only on slide, no body present.
Trissolcus ogyges (Dodd),
New ComMBINATION
Telenomus ogyges Dodd, 1913c: 166. SAM
Lectotype 2 and 2 paralectotype 2 on slide
11417; “Telenomus ogyges Dodd 2 type,
sweeping Cape River, Pentland, NQ, Jan.
241
13, A. A. Girault.”” Condition: lectotype
(here designated) near edge of cover slip,
head attached to mesosoma, mesosoma
crushed.
Trissolcus oreas (Dodd),
NEw ComBINATION
Telenomus oreas Dodd, 1913c: 180. SAM
Holotype 2 on slide 11449; *““Telenomus
oreas Dodd 2 type: Sweeping in jungle, Nel-
son, NQ, 15.v.13, A P Dodd.” Condition:
head and mesosoma badly crushed.
Trissolcus orontes (Dodd),
NEw COMBINATION
Telenomus orontes Dodd, 1914b: 120. SAM
Holotype 2 on slide 12181; **Telenomus
orontes Dodd ¢° type; 2182. Condition:
crushed.
Trissolcus otho (Dodd),
New ComBINATION
Telenomus otho Dodd, 1914c: 252. SAM
Holotype 2 on point, “Telenomus otho
Dodd 2 type; Cairns.’ Condition: tagmata
detached, otherwise good. Fore wing, an-
tennae on slide 111100, ““Telenomus otho
Dodd ¢ type, forewing antennae”; antennae
crushed.
Trissolcus wilsoni (Dodd),
New ComBINATION
Telenomus wilsoni Dodd, 1930: 28. NMV
Holotype °; “Eltham, V., F. E. Wilson,
May, 1927; HOLOTYPE T-1420 Teleno-
mus wilsoni Dodd [red museum label]; Te-
lenomus wilsoni Dodd, °, Holotype [Dodd’s
handwritten label]; F. E. Wilson Collec-
tion.” Condition: good; right mid leg be-
yond coxa, hind leg beyond femur missing;
left legs hidden beneath body.
SUBFAMILY SCELIONINAE
Embidobia Ashmead
Embidobia oaxes (Dodd),
New COMBINATION
Telenomus oaxes Dodd, 1914b: 120. SAM
Holotype @ on card, ““Telenomus oaxes
Dodd ¢ type; 12184.” Condition: good. Head
on slide 12184, ““Telenomus oaxes Dodd, ?
type, head, forewings, 2184. Condition:
head badly crushed; fore wings not found.
Gryon Haliday
Gryon orestes (Dodd),
New COMBINATION
Telenomus orestes Dodd, 1913c: 167. SAM
Holotype 4 on slide 11421; ““Telenomus
orestes Dodd 4 type, on window, Herberton
(3000 ft) NQ, 28.11.11, A A Girault.”” Con-
dition: head and mesosoma crushed.
ACKNOWLEDGMENTS
I gratefully acknowledge the help and hos-
pitality of I. D. Naumann (Canberra), J.
Cardale (Canberra), G. Gross (Adelaide), A.
D. Austin (Adelaide), I. D. Galloway (In-
dooroopilly), E. C. Dahms (Brisbane) and
G. Monteith (Brisbane) during my trip to
Australia; to K. Walker (Melbourne) for the
loan of material; and to L. Masner (Ottawa)
and J. B. Whitfield (Columbus) for review
of the manuscript. This material is based
upon work supported by the National Sci-
ence Foundation under Grant No. BSR-
8516579.
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PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 244-247
TELENOMUS SPECIES (HYMENOPTERA: SCELIONIDAE)
ASSOCIATED WITH THE EGGS OF
ZYGAENIDAE (LEPIDOPTERA)
NORMAN F. JOHNSON AND FERDINANDO BIN
(NFJ) Department of Entomology, The Ohio State University, 1735 Neil Avenue,
Columbus, Ohio 43210; (FB) Istituto di Entomologia Agraria, Universita di Perugia, Borgo
XX Giugno, 06100 Perugia, Italy.
Abstract. —Two species of Te/lenomus are known to parasitize the eggs of Zygaenidae.
Telenomus argus n. sp. has been reared from the eggs of the vineyard pest Theresimima
ampelophaga Bayle-Barelle in Israel. Telenomus zygaenae Kieffer was reported to attack
the eggs of Zygaena lonicerae (Scheven); it is compared with 7. argus and its host rela-
tionships discussed.
Key Words:
The new species of Telenomus (Hyme-
noptera: Scelionidae) described below has
been reared from the eggs of Theresimima
(Ino) ampelophaga Bayle-Barelle (Lepidop-
tera: Zygaenidae) in Israel. This moth is one
of the few zygaenids of any economic im-
portance: it feeds on Vitis spp. in countries
around the Mediterranean Basin and in oth-
er warm areas of the western Palearctic (Ba-
lachowsky 1972). We therefore present this
description in order to provide biological
control workers with a name for the para-
sitoid. In addition, we discuss 7. zygaenae
Kieffer, the only other Te/enomus recorded
from this family of moths. The morpholog-
ical terminology used follows that discussed
in Johnson (1984).
Telenomus argus, NEW SPECIES
Fig. |
Length 0.47-0.54 mm (n = 20 males, 20
females). Small, but typical species of the
T. californicus complex (see Johnson 1984
for characters of that taxon).
Head: hyperoccipital carina absent, ver-
tex broadly rounded onto occiput; preocel-
Parasitoid, biological control
lar pit (Bin and Dessart 1983) absent; frons
smooth medially, orbital bands present
ventrally, effaced dorsally; frons width >
eye height; lower frons with curved wrinkles
flanking clypeal range and antennal inser-
tions; labrum articulated with clypeus, not
fused, mandibles tridentate, teeth subequal
in size.
Mesosoma: notauli absent; disk of scu-
tellum smooth; dorsellum as long laterally
as medially, rugulose above, striate below;
episternal foveae, metapleural carina ab-
sent; mesopleural carina absent, mesopleur-
al scrobe not sharply defined anteriorly; ace-
tabular field small; acetabular carina simple,
not crenulate.
Metasoma: T1 with one pair of sublateral
setae; T2 transverse, smooth beyond basal
foveae; male genitalia (Fig. 1) with laminae
volsellares distinctly separated, strongly
melanized; aedeagal lobe moderately elon-
gate, slightly narrowed apically, apex trun-
cate or weakly excised; penis valves not well
differentiated; digiti each with two large dig-
ital teeth, teeth subequal in length.
Diagnosis: Telenomus argus is most eas-
VOLUME 90, NUMBER 2 245
0.2 mm
Figs. 1-2. Male genitalia, ventral view. 1, Telenomus argus. 2, Telenomus zygaenae
246 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ily recognized by the characters of the male
genitalia, in particular the possession of two
equally large teeth on each digitus and the
well-developed, but distinctly separated
laminae volsellares. Other species with only
a single pair of teeth on each digitus may
be easily separated by other genitalic char-
acters. Telenomus lobatus Johnson & Bin
has an extremely long aedeagal lobe, longer
than the remainder of the aedeago-volsellar
shaft (Johnson and Bin 1982). In both 7.
ampullaceus Johnson & Bin and T. turbatae
Nixon the aedeago-volsellar shaft is dis-
tinctly wider than the aedeagal lobe (John-
son and Bin 1982, Nixon 1937). Telenomus
sciron Nixon has small, delicate digital teeth,
in contrast to the long, thick structures in
T. argus (Nixon 1935). Telenomus guang-
dongensis Chen & Liao appears to have a
very broad aedeago-volsellar shaft and to
have the laminae volsellares closely ap-
proximated medially (Wu et al. 1979).
The genitalia of 7. zygaenae (Fig. 2), also
reportedly reared from the eggs of a zygae-
nid, are very similar to 7. argus, but may
be distinguished by the presence of three
teeth per digitus, with the mesal tooth dis-
tinctly smaller than the other two (Fig. 2).
We have found the number of these struc-
tures to be variable only in those species
with small teeth; among those with long,
stout digital teeth (the ca/lifornicus group,
arzamae group, dalmanni group, see John-
son 1984) the number seems to be constant
within a species.
The species described here keys out to 7.
etiellae Kozlov in Kozlov and Kononova
(1983). The latter, however, is known from
only four female specimens from the lower
Volga region of the Soviet Union. Its dis-
tinguishing features, viz, short funicular
segments, long fringe of setae along the pos-
terior margin of the hind wing, and short
striae at the base of T2, characterize a num-
ber of small species that parasitize the eggs
of Lepidoptera. Telenomous argus may be
conspecific with 7. etiellae, but this deter-
mination must await the discovery of either
males of the latter or useful diagnostic char-
acters in the females.
Material.—Holotype male: Israel: Jeru-
salem; 5.vii.1978; Y. Eisenstein; ex There-
simima ampelophaga; deposited in the Ca-
nadian National Collection of Insects,
Arachnids and Nematodes (Ottawa, Ontar-
io). Paratypes: 12 males, 18 females with
same data as holotype; 20 males with same
locality data, collected 9.vii.1981, deposit-
ed in the authors’ collections, CNC, and
British Museum (Natural History).
Discussion. — This tiny species, like many
other Te/enomus that parasitize the eggs of
Lepidoptera, is difficult or impossible to
identify on the basis of external morphol-
ogy. The male genitalia provide by far the
best characters for separating it from others.
Unfortunately, the male genitalia of very
few Palearctic Telenomus have been de-
scribed or figured; many species, in fact, are
known only from females. Thus it has been
impossible for us to be completely assured
that this species has not already been de-
scribed. We present it as a new species an-
ticipating that when the identities of the
many Jelenomus described in the last 150
years are finally established, its proper sta-
tus can be clearly and easily recognized.
Comments on Host Associations.— 7e-
lenomus argus was reared from its host in
two years, 1979 and 1981, but its economic
importance has yet to be assessed (D. Ger-
ling, in litt.). The parasite may be confined
to Israel or the eastern Mediterranean Basin
as no other rearings have been reported de-
spite the relatively wide distribution and
food-plant specificity of its host.
Telenomus zygaenae was reportedly
reared from Zygaena lonicerae (Scheven) in
Denmark (Kieffer 1913), but this needs to
be confirmed. The host name written on the
labels of the type material of 7. zygaenae is
“Zygaena filip.”, standing for fillipendulae
(L.). Neither the egg mass nor a description
of it is available, so this conflict between the
published information and the label data
cannot be directly resolved.
VOLUME 90, NUMBER 2
ACKNOWLEDGMENTS
We thank Dr. D. Gerling (Tel Aviv Uni-
versity) for offering us these Telenomus and
to Dr. B. Petersen (Zoologisk Museum, Co-
penhagen) for making the type material of
T. zygaenae available; and to J. B. Whitfield
for comments on the manuscript.
LITERATURE CITED
Balachowsky, A. S. 1972. Superfamille des Zygae-
noidea. Entomologie appliquée a |’Agriculture.
Tome II, pp. 1059-1634. Masson et C. Edit, Paris.
Bin, F. and P. Dessart. 1983. Cephalic pits in Proc-
totrupoidea Scelionidae and Ceraphronoidea (Hy-
menoptera). Redia 66: 563-575.
Johnson, N. F. 1984. Systematics of Nearctic Telen-
omus: classification and revisions of the podisi and
phymatae species groups (Hymenoptera: Scelion-
idae). Bull. Ohio Biol. Surv. 6(3). 113 pp.
247
Johnson, N. F. and F. Bin. 1982. Species of Telen-
omus (Hym., Scelionidae), parasitoids of the stalked
eggs of Neuroptera (Chrysopidae & Berothidae).
Redia 65: 189-206.
Kieffer, J. J. 1913. Zwei neue Hymenopteren aus
Danemark. Entomol. Meddel. 4: 378-380.
Kozlov, M. A. and S. V. Kononova. 1983. [Telen-
omine fauna of the USSR (Hymenoptera, Scelion-
idae, Telenominae).] Determinations of the Fauna
of the USSR, vol. 136. Publishing House “‘Nau-
ka,” Leningrad. 336 pp.
Nixon, G. E. J. 1935. A revision of the African Te-
lenominae (Proctotrupoidea, fam. Scelionidae).
Trans. R. Entomol. Soc. Lond. 83: 73-103.
1937. New Asiatic Telenominae (Hym.,
Proctotrupoidea). Ann. Mag. Nat. Hist. (10) 20:
113-127.
Wu Yen-ju, Chen Tai-lu, Liao Ting-shi, and Her Jyuhn-
hua. 1979. [Descriptions of six new species of
Telenomus (Hymenoptera: Scelionidae).] Acta
Zootaxon. Sinica 4: 392-398.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 248-255
EFFECTS OF MALATHION AND DIAZINON EXPOSURE ON FEMALE
GERMAN COCKROACHES (DICTYOPTERA: BLATTELLIDAE)
AND THEIR OOTHECAE
J. D. HARMON AND M. H. Ross
Department of Entomology, Virginia Polytechnic Institute and State University, Blacks-
burg, Virginia 24061.
Abstract. — Female German cockroaches, Blattella germanica (L.), were exposed to mal-
athion and diazonon by tarsal contact. The effects of exposure on hatch of oothecae and
on newly-emerged nymphs are compared in resistant and susceptible strains. Insecticide
exposure increased the frequency of oothecal drop over that which occurred naturally in
the susceptible but not in the malathion or diazinon-resistant strains. The time from
exposure to hatch was decreased when oothecae hatched on a treated surface in all except
oothecae of the diazinon-resistant strain. Exposure to diazinon, but not malathion, de-
creased nymphal emergence. Nymphs of the susceptible strain that emerged on treated
surfaces were unable to shed the embryonic cuticle, whereas 18 to 23% of resistant strain
nymphs were upright and moving freely. Results are compared to similar experiments
with propoxur.
Key Words: Insecticides, behavior
Many studies of the German cockroach,
Blattella germanica (L.), have dealt with in-
secticide resistance. Behavioral responses to
insecticides have received less attention.
German cockroaches are repelled by certain
insecticides (Ebeling et al. 1967, 1968, Ebel-
ing et al. 1966), but little is known con-
cerning possible modifications of the repel-
lency response following several decades of
selection pressure from insecticides. Bret and
Ross (1985, 1986) reported evidence that
behavioral modifications have indeed ac-
companied the development of insecticide
resistance. Dispersal and grooming behav-
ior in adults of a susceptible strain exposed
to propoxur vapors differed from that of
adults in a propoxur-resistant field strain.
Irritability and repellency are widely rec-
ognized responses to insecticides (Lock-
wood et al. 1984), but in B. germanica, a
third response has been reported. Insecti-
cide exposure may cause a female to drop
her ootheca prematurely (Woodbury 1938,
Parker and Campbell 1940, Russell and
Frishman 1965, van den Heuvel and Shenk-
er 1965, Chadwick and Evans 1973, Muller
and Coch 1975, Barson and Renn 1983,
Harmon and Ross 1987). In two of the fore-
going studies, comparisons were made be-
tween resistant and susceptible strains. Rus-
sell and Frishman (1965) reported a higher
frequency of premature drop among sus-
ceptible than chlordane-resistant females
when exposed to vapors from dichlorvos
resin strips. The ages of the oothecae were
unknown. Harmon and Ross (1987) also
found a higher retention of oothecae by re-
sistant than susceptible females. They used
females of a propoxur-resistant field strain
and a laboratory susceptible strain (VPI) that
VOLUME 90, NUMBER 2
carried oothecae in Stage XII of embryonic
development (Tanaka 1976), that is, oothe-
cae that were due to hatch within 48 to 72
h after selection.
When oothecae are within 2 to 3 days of
hatch and conditions approximate those
typical of German cockroach infestations
(room temperature; r.h. usually above 30%),
hatch and nymphal survival depend pri-
marily on whether the ootheca falls on a
treated or untreated surface. Van den Heu-
vel and Shenker (1965), Barson and Renn
(1983) and Harmon and Ross (1987) found
treatment of females bearing mature oothe-
cae had little effect on hatch or productivity
of the ootheca unless it dropped on a treated
surface. On the other hand, when oothecae
hatched on a treated surface, nymphal
emergence and survival, when studied, were
affected. Decreases in the latter effects due
to hatch on a propoxur-treated surface were
less in a propoxur-resistant than in a sus-
ceptible strain (Harmon and Ross 1987).
Reported here are experiments on the ef-
fects of malathion and diazinon on females
carrying mature oothecae. The purpose was
two-fold: first, to compare the effects on fe-
males of resistant strains to those of sus-
ceptible strains; secondly, to compare the
effects of two organophosphates (malathion
and diazinon) to those of propoxur, a car-
bamate.
MATERIALS AND METHODS
Ootheca-bearing females were from three
strains: the VPI strain, our standard sus-
ceptible laboratory strain; the Carver strain,
a field strain resistant to malathion, pro-
poxur, and pyrethrins, collected in Gaines-
ville, Fla., in 1983 (Cochran, pers. com-
mun.); and the Lynn Haven strain, a
diazinon-resistant strain collected in Lynn
Haven, Va., in 1983 (Cochran, pers. com-
mun.).
Cockroach rearing conditions and test
procedures were like those of the propoxur
experiments (Harmon and Ross 1987). In
brief, females carrying 20-24 day-old first
249
oothecae were lightly anesthetized with CO,
and examined under low power of a bin-
ocular microscope to determine the stage of
egg development, using Tanaka’s (1976) de-
velopmental table. Females carrying Stage
XII oothecae were selected for test pur-
poses, that is, females carrying oothecae due
to hatch within 48 to 72 h. Only those 0o-
thecae with normal-appearing embryos in
each compartment or, at most, no more than
three undeveloped eggs, were used. The fe-
males were held for 24 h and then exposed
by tarsal contact to filter paper treated with
either malathion or diazinon (Cochran
1973). The filter paper (Whatman #1) was
cutin 15 x 15 cm squares, placed on a glass
plate, and impregnated evenly with a 3 ml
mixture of risella oil and trichloroethylene
(1:2 v/v) and either 0.08 ml of technical
grade malathion (0.356 ul/cm?’) or 0.015 ml
of technical grade diazinon (0.067 pl/cm?).
Dosages were selected by preliminary ex-
perimentation so as to give an approximate
mortality of 65% among VPI strain females
following an exposure period of 100 min.
The purpose was to make the results com-
parable to prior work with propoxur, where
65% of the VPI strain females died as a
result of a 100 min exposure to propoxur-
treated filter paper (Harmon and Ross)
1987).
Groups of 10 to 12 ootheca-bearing fe-
males were confined by a glass chimney on
the treated papers or on unused filter paper
(controls) after the papers had dried for 1
h. Initial tests with the TCE/risella oil mix-
ture showed that, as expected, neither le-
thality nor oothecal drop were affected. TCE
evaporates rapidly and the possibility that
risella oil, basically a mineral oil, would af-
fect the cockroaches was highly unlikely. The
experiments were begun between 9:00 and
11:00 h. Oothecal drop during exposure was
recorded. Following exposure, females that
dropped oothecae were separated from those
that retained oothecae. The latter were
placed in separate jars for subsequent ob-
servations, as follow: retention of oothecae
250
at 24 h after exposure; hatch of oothecae;
time from the end of the exposure period
to hatch; number of embryos remaining in
the ootheca; number of nymphs that
emerged; and the number that were alive
and whether they were upright and/or fully
pigmented 24 h after hatch. Like observa-
tions were made on oothecae that dropped
during the exposure period. They were kept
on the treated papers but were placed in
individual jars with a water source 5 to 7
cm above the bottom of the jar that main-
tained a high humidity (85-90% r.h.) within
the jar, ensuring the oothecae did not dry
out. The purpose was to assess the effects
of a treated surface on hatch and nymphal
survival. Additional data were obtained by
manually detaching oothecae of like stage
to those carried by females at the time of
exposure and placing them in individual jars
on treated papers as described above or, in
the case of the controls, on clean filter paper.
Female mortality was recorded at 72 h after
exposure. At that time, mortality was com-
plete and could be compared to that in the
propoxur experiment.
Percentages of oothecal hatch, nymphal
emergence, nymphal survival, and freely-
moving nymphs were calculated. Nymphs
were counted as live if pulsation was oc-
curring in the dorsal blood vessel. This in-
cluded nymphs that were alive, but that were
unable to free themselves from the embry-
onic cuticle. Percentage survival was based
on the number of live nymphs among those
that emerged successfully in order to distin-
guish between death that occurred within
the oothecae (reduced emergence) from that
due to emergence on a treated surface.
Data were analyzed using the Statistical
Analysis Service (SAS) and the Bio-medical
Data Processing (BMDP) statistical soft-
ware available on the VPI & SU IBM main-
frame computer. The best statistical model
to analyze each type of data was determined
by loglinear models (BMDP). Percentage
premature drop of oothecae was analyzed
by a least significant differences (LSD) pair-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
wise comparison procedure. The associa-
tion between mortality and premature drop
was analyzed by establishing two-way fre-
quency tables. The null hypothesis was that
post treatment mortality among females that
retained their oothecae did not differ sig-
nificantly from that of females that dropped
their oothecae. Percentage hatch was ana-
lyzed using a one-way ANOVA (F = 19.79;
df = 1075: and P > 0.0001) followed by a
LSD comparison test. Times from exposure
to hatch were analyzed using a general linear
models procedure (F = 134.55; df = 893;
and P > 0.0001) followed by a LSD com-
parison test. Means were calculated for each
treatment group and sorted in ascending or-
der. Significance groupings were assigned
from LSD tests. Estimates of percentage
nymphal emergence, survival, and freedom
of movement were arc-sine transformed and
then analyzed as above with a general linear
models procedure and LSD pairwise com-
parison.
RESULTS
None of the VPI strain females died dur-
ing exposure to untreated filter paper and,
in the two resistant field strains, only one
(3%) died in each strain (Table 1). Mortality
due to malathion exposure of VPI strain
females did not differ significantly from that
due to diazinon exposure, as expected due
to pre-selection of dosages. Resistance in
the Carver strain reduced mortality from
malathion to approximately half that of the
VPI strain. No mortality occurred following
exposure of Lynn Haven strain females to
diazinon.
Premature drop of oothecae during the
exposure period occurred only in the mal-
athion experiment. It was limited to 3 0o-
thecae in one strain and 4 in the other (Table
1). No oothecae were dropped by the control
females or those in the experiment with di-
azinon. A tendency towards higher mortal-
ity of females that dropped oothecae than
of those that retained oothecae was evident.
Mortality was significantly higher among the
VOLUME 90, NUMBER 2
Table 1.
Stage XII: oothecae (oot).
251
Effects of malathion and diazinon on mortality and oothecal drop in female B. germanica carrying
————————————
Control Data?
% Mortality
oot Drop during oot Drop 24 h after
22 Dropped Exposure Exposure
oot during
Strain No. of 22 All Exposure Retained oot No. % No %
om
VPI 27 Oa - Oa 0 Oa 4 15 be
Carver 34 3a — 3a 0 Oa 7 Pie
Lynn Haven 30 3a _ 3a (0) Oa 2) 17 be
Malathion Treatment
VPI 100 56¢ 66b DIG 3 3a 28 28c
Carver 102 24b 50 a4 23b 4 4a 17 17 be
Diazinon Treatment
VPI 100 63¢ _ 63¢ 0 (0) 26 26¢
Lynn Haven 48 Oa = Oa 0 0 4 8a
* Means within a column followed by the same letter are not significantly different (P > 0.05).
» Control females were exposed to untreated filter paper.
© 72 h after exposure to untreated or treated filter paper.
4 Significantly higher than mortality of females that retained their oothecae (23%) (P > 0.05).
former when results from the two strains
were pooled (P > 0.05).
Normal drop and hatch of oothecae was
expected to occur in the period from 48 to
72 h after selection. Oothecae 24 h after
exposure were aged 48 h plus 100 min (time
of exposure of females to malathion, diaz-
inon, or clean filter paper). At this time,
control females had begun to drop their
oothecae. The frequency of drop by control
females ranged from 15 to 21% in the three
strains (Table 1). Percentage drop by VPI
strain females was increased significantly
over that of unexposed females when the
females had been exposed to either mala-
thion or diazinon. The difference between
naturally occurring drop and that found
among insecticide-exposed VPI strain fe-
males gives a rough estimate of drop due to
the insecticide, i.e. 13% were dropped pre-
maturely due to malathion and 11% due to
diazinon. In the Carver strain, drop follow-
ing exposure to malathion did not differ sig-
nificantly from that in the controls. Reten-
tion of oothecae by Lynn Haven strain
females exposed to diazinon was signifi-
cantly longer than that by control females.
Table 2 summarizes the effects of mala-
thion and diazinon on hatch of oothecae
and on the newly-hatched nymphs. “Days
to hatch” refers to the time between the end
of the exposure period and hatch (opening)
of the ootheca. In the control data, results
were closely similar. The only notable vari-
ations were that detached oothecae of VPI
strain females took longer to hatch than
those that hatched naturally (attached) and
hatch time of the latter was significantly less
than in the Carver and Lynn Haven strains.
In the malathion experiment, VPI and
Carver strain oothecae hatched more quick-
ly on the treated surface (oothecae dropped
or detached) than on an isecticide free sur-
face (Table 1, control data and oothecae at-
tached to malathion-exposed females).
Hatch time was also decreased when VPI
strain oothecae hatched on a diazinon treat-
ed surface.In one instance only, the time
required for hatch on a treated surface was
not decreased over that in the control data.
That was among detached Lynn Haven
strain oothecae.
In general, the number of oothecae that
hatched (opened to permit nymphal emer-
252
Table 2.
emergence and survival.*
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Effects of malathion and diazinon on hatch of Stage XII B. germanica oothecae (oot) and on nymphal
Control Data
oot Hatched
Nymphs from Hatched oot
Days to % % % Freely
Strain oot Hatch? n % Emergence Survival: Moving*
VPI attached 1.9b Dy 100a 96a 100a 100a
detached 2 Nic 36 92a 95a 100a 100a
Carver attached 2.7 cd 32 100a 96a 100a 100a
detached 2.3\¢ 29 85b 90 b 100a 100a
Lynn Haven attached 22d'C 30 100a 95a 99a 100a
detached 2:2'¢ 30 100a 89b 100a 100a
Malathion Treatment
VPI attached 1.9b 84 86b 94a 100a 100a
dropped O0.3a 3 100a 92a 97a 2d
detached 1.5Sb 101 98a 90 b 94b Od
Carver attached 2.36 95 94a 91 ab 100a 100 a
dropped 1.0a 1 100 a 93a 100a 54b
detached 1.8b 89 94a 83c¢ 95 ab 23K
Diazinon Treatment
VPI attached 31d 80 80 b 83c¢ 100 a 100a
dropped ~ 0) = — — —
detached I5'b 102 97a 83¢ 87b Od
Lynn Haven attached 2.5¢ 40 85b 80c¢ 100 a 100 a
dropped 7 1
detached 23G 46 93a 89 be 99a 8c
* Means within a column followed by the same letter are not significantly different (P > 0.05).
» Days from end of exposure period (24 h and 100 min after oot stage identification) to hatch.
© % of emerged nymphs that were alive 24 h after emergence.
4% of emerged nymphs that were free of the embryonic cuticle and moving about freely 24 h after emergence.
gence) was not decreased due to manual de-
tachment, except for possible injury to a few
in the Carver strain (Table 2). In the ex-
perimental groups, small but significant re-
ductions occurred among oothecae retained
by malation-treated VPI strain females and
by diazinon-treated VPI and Lynn Haven
strain females (attached oothecae). Some of
the VPI and a few of the Carver strain 00-
thecae were from dead females, but death
had no perceptible effect on hatch. Once the
nymphs had forced open the oothecae at the
time of hatch, most emerged successfully.
At least 95% emerged from oothecae of the
untreated control females, except for 90 and
89% from detached Carver and Lynn Haven
oothecae, respectively. Handling of the lat-
ter may have injured a few of the eggs. A
tendency towards reduced emergence was
evident in the malathion experiment, but it
was only in the diazinon experiment that
treatment had a clear effect on emergence.
It was reduced among attached and de-
tached oothecae of both the VPI and Lynn
Haven strains.
Nymphs were scored as living at 24 h after
exposure of the parent females to untreated
or treated filter paper if pulsation was ob-
served in the dorsal blood vessel. On this
basis, survival was below 99% only in sit-
uations where nymphs emerged on a treated
surface (Table 2, dropped or detached).
Slight but significant reductions occurred
among detached oothecae of VPI and Car-
ver strain females exposed to malathion, al-
though the results were similar for the two
VOLUME 90, NUMBER 2
strains (94 and 95% survival, respectively).
The greatest decrease (87% survival) was
among VPI strain nymphs that hatched on
a diazinon-treated surface. Survival of Lynn
Haven strain nymphs was not affected. Dead
nymphs, as well as survivors, were fully pig-
mented.
The main difference between the effects
of malathion and diazinon on newly-hatched
nymphs of susceptible and resistant strains
was in the ability of nymphs that hatched
on a treated surface to free themselves from
the embryonic cuticle (Table 2). These
nymphs were still alive at 24 h after emer-
gence. Entanglement of VPI strain nymphs
on the malathion and diazinon treated sur-
faces was complete except for 2% of those
that emerged from oothecae that dropped
on a malathion-treated surface. The per-
centage of Carver strain and Lynn Haven
strain nymphs that freed themselves was
significantly higher than in the VPI strain.
The nymphs of the resistant strains were
upright and could have escaped the treated
surface if permitted to do so.
DISCUSSION
Estimates of times between exposure of
females to untreated filter paper and hatch
may have been affected by slight differences
in oothecal development at the time of se-
lection. Nevertheless, it was apparent that
oothecae carried by field strain females took
longer to hatch than those of the VPI strain.
This difference may have been related to
either laboratory culture or modifications of
resistant field strains. Although Muller and
Coch (1975) found that manual detachment
increased hatch time, a similar phenome-
non was observed here only in the VPI strain.
Hatch on a treated surface decreased hatch
time in all experiments except that with the
Lynn Haven strain. Another difference in
this strain was that diazinon treatment
caused a longer-than-normal retention of
oothecae. More extensive studies on field
strains are needed before it can be deter-
mined whether these special characteristics
to
wn
is)
are related to the development of diazinon
resistance.
The only effect of insecticide exposure that
could be attributed to exposing females
rather than to hatch on a treated surface was
a slight reduction in the number of oothecae
that hatched. Only the Carver strain females
did not show this effect. Perhaps it was only
when oothecae were retained that there was
suficient time for toxic substances to be
passed from the females to the developing
embryos.
Emergence was reduced slightly when
oothecae hatched on a treated surface. Diaz-
inon had a stronger effect than malathion,
especially in the susceptible strain. Possibly
vapors from diazinon penetrated the oothe-
ca more deeply than those of malathion,
both during exposure of ootheca-bearing fe-
males and when oothecae were dropped or
placed on a diazinon-treated surface. Law-
son (1949) found that diazinon was the only
one of several insecticides tested that caused
a reduction in emergence.
A basis was laid for comparing the effects
of propoxur (Harmon and Ross 1987) to
those of two organophosphates, malathion
and diazinon, by selecting dosages that
caused similar mortality in VPI strain fe-
males during the same period (100 min).
The primary difference was that premature
drop of oothecae due to propoxur was more
immediate than that due to the two organo-
phosphates. Delay may have been due to
the slower action of malathion and diazinon
than of propoxur (Matsumura 1985).
Oothecae dropped due to exposure to pro-
poxur would be more likely to fall ona treat-
ed surface, as observed in shipboard exper-
iments (Ross and Bret 1986). Whether
oothecae would indeed be dropped pre-
maturely depends on whether or not the
strain was resistant. A second difference be-
tween the effects of propoxur and those re-
ported here was that nymphs that hatched
on a propoxur-treated surface died before
pigmentation was complete, whereas dead
nymphs in the present experiments were
254
fully pigmented. This difference may also
be attributed to a slower action of the two
organophosphates than of propoxur.
The results of the experiments with mal-
athion and diazinon, considered in con-
junction with the earlier study on propoxur,
indicate clearly that premature drop of
oothecae is a phenomenon associated with
susceptibility. It is decreased or, indeed, may
disappear entirely among females resistant
to chlordane (Russell and Frishman 1965),
propoxur, malathion, and diazinon. In ad-
dition, Lawson (1949) found a higher mor-
tality among females that dropped oothecae
prematurely than among those that retained
them and Muller and Coch (1975) reported
that premature drop only occurred when the
dosage of an insecticide was sufficient to
cause the knock-down condition. Mortality
and knockdown are, of course, indications
of susceptibility. The relative frequency of
premature drop by females of a resistant
strain compared to those of a susceptible
strain may well provide an indication of the
frequency of genetically-susceptible females
present in the resisistant strain, providing
it is not completely homozygous for resis-
tance-conferring genes. Perhaps most insec-
ticides, regardless of differences in mecha-
nisms of resistance, affect the nervous system
of susceptible females in a way that leads to
premature drop. In any case, the ability of
resistant strain females to retain their oo-
thecae following insecticide exposure is
clearly an advantageous characteristic.
When females bearing mature oothecae are
dispersed due to an insecticide treatment,
they have more opportunity to find a situ-
ation (harborage) favorable to oothecal hatch
and survival of newly-hatched nymphs.
When immature oothecae are retained, the
likelihood of their maturing successfully is
enhanced.
The only oothecae that could be distin-
guished on an individual basis as being
dropped prematurely were those aborted
during the exposure period. Although only
seven were dropped, mortality of their par-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ent females was higher than that of females
that retained oothecae—a finding that can
be attributed to the association of prema-
ture drop with suceptibility noted above.
Many of the nymphs that emerged on di-
azinon or malathion treated surfaces were
unable to shed the embryonic cuticle—a phe-
nomenon that was reported previously fol-
lowing exposure to a variety of insecticides
(Lawson 1949, Killough 1958, Harmon and
Ross 1987). The present study supports Kil-
lough (1958) who concluded that entangle-
ment would not occur unless an insecticide
came in contact with nymphs during emer-
gence. Entanglement was nearly complete
in the susceptible strain but not the mala-
thion or diazinon resistant strains. On the
basis that entanglement would eventually
have resulted in death, mortality of suscep-
tible strains nymphs exceeded that of the
resistant nymphs. The frequency of suscep-
tible nymphs that were dead at the time of
observation (24 h after hatch) did not differ
markedly from that of resistant strain
nymphs, although a tendency towards higher
mortality of susceptible nymphs occurred
in the malathion experiment and, in the di-
azinon experiment, the difference was sig-
nificant.
Whether drop would be sufficiently rapid
to cause oothecae to fall on a treated surface
would depend on both resistance charac-
teristics of the target population and on the
choice of insecticides. If oothecae did hap-
pen to hatch on a treated surface, the effects
of malathion or diazinon on resistant strain
nymphs would probably be less severe than
those of propoxur (Harmon and Ross 1987).
At least some of the nymphs that survived
for 24 h on the treated surfaces were free of
the embryonic cuticle and could have left
the treated surface if permitted to do so.
Nymphs that were still alive after a similar
exposure to propoxur were either entangled
in the cuticle or were in the knockdown con-
dition. We conclude that oothecal retention
by resistant females, as well as decreased
mortality, adds to the difficulty of control-
VOLUME 90, NUMBER 2
ling insecticide resistant populations of the
German cockroach.
ACKNOWLEDGMENTS
Weare grateful to D. G. Cochran for help-
ful criticism of the manuscript and to the
National Pest Control Association for par-
tial support of the research.
LITERATURE CITED
Barson, G. and N. Renn. 1983. Hatching from oothe-
cae of the German cockroach (Blattella germani-
ca) under laboratory culture conditions and after
premature removal. Entomol. Exp. Appl. 34: 179-
185.
Bret, B. L.and M.H. Ross. 1985. Insecticide-induced
dispersal of the German cockroach, Blattella ger-
manica (L.) (Orthoptera: Blattellidae). J. Econ.
Entomol. 78: 1293-1298.
1986. Behavioral responses of the German
cockroach, Blatella germanica (L.) (Orthoptera:
Blattellidae) to a propoxur formulation. J. Econ.
Entomol. 79: 426-430.
Chadwick, P. R. and M. Evans. 1973. Laboratory
and field tests with some pyrethroids against cock-
roaches. Intl. Pest Control 15: 11-16.
Cochran, D. G. 1973. Inheritance of malathion re-
sistance in the German cockroach. Entomol. Exp.
Appl. 16: 83-90.
Ebeling, W., D. A. Reierson, and R. E. Wagner. 1967.
Influence of repellency on the efficacy of blatti-
cides. II. Laboratory experiments with German
cockroaches. J. Econ. Entomol. 60: 1375-1390.
1968. Influence of repellency on the effects
of blatticides. III. Field experiments with German
cockroaches, with notes on three other species. J.
Econ. Entomol. 61: 751-761.
Ebeling, W., R. E. Wagner, and D. A. Reierson. 1966.
Influence of repellency on the efficacy of blatti-
cides. I. Learned modifications of behavior of the
German cockroach. J. Econ. Entomol. 59: 1374—
1388.
Harmon, J. D. and M. H. Ross. 1987. Effects of pro-
poxur exposure on females of the German cock-
roach, Blattella germanica and their oothecae.
Entomol. Exp. Appl. 44: 269-275.
Killough, R. A. 1958. Susceptibility of the oothecae
of the American cockroach, Periplaneta ameri-
cana Linn., to various insecticides. M.S. Thesis,
Purdue Univ., West Lafayette, IN.
Lawson, F. A. 1949. Structural and insecticidal stud-
ies on cockroach egg cases. Ph.D. Dissertation,
Ohio State University, Columbus, Ohio. 54 p.
Lockwood, J. A., T. C. Sparks, and R. N. Story. 1984.
Evolution of insect resistance to insecticides: A
reevaluation of the roles of physiology and be-
havior. Bull. Entomol. Soc. Am. 30: 41-51.
Matsumura, F. 1985. Toxicology of Insecticides. Ple-
num Press, N.Y. 598 pp.
Muller, von P. and F. Coch. 1975. Abwurfund schupt
der ootheken von Blattella germanica (L.) unter
einwirkung von insektiziden. Z. ges. Hyg. 21: 899-
903.
Parker, B. M. and F. L. Campbell. 1940. Relative
susceptibility of the ootheca and adult female Ger-
man cockroach to liquid household insecticides.
J. Econ. Entomol. 116: 57-68.
Ross, M. H. and B. L. Bret. 1986. Effects of propoxur
treatment on populations containing susceptible
and resistant German cockroaches (Orthoptera:
Blatellidae). J. Econ. Entomol. 79: 338-349.
Russell, M. P. and A. M. Frishman. 1965. Effective-
ness of dichlorovos in resin strips for the control
of the German cockroach, Blattella germanica. J.
Econ. Entomol. 58: 570-572.
Tanaka, A. 1976. Stages in the embryonic develop-
ment of the German cockroach, Blattella german-
ica Linne. (Blattaria: Blattellidae). Kontyu 44: 512-
525.
van den Heuvel, M. J. and A. M. Shenker. 1965.
Cockroach control using non-persistent insecti-
cides. Intl. Pest Control 7: 10-11.
Woodbury, E. N. 1938. Test methods on roaches.
Soap Sanitation Chem. 14: 89-90, 107, 109.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 256-257
NOTE
Hyadaphis tataricae (Homoptera: Aphididae):
10 Years After its Introduction into
North America
The arrival of alien species of insects on
the North American continent is a common
phenomenon, whether occurring acciden-
tally or purposefully. The aphid Hyadaphis
tataricae (Aizenberg) is one such recently,
accidentally-introduced immigrant (Voeg-
tlin. 1981. Proc. Entomol. Soc. Wash. 83(2):
361-362); it has not only established itself
but also has rapidly expanded its geograph-
ical range. This aphid has a fairly narrow
host range within the honeysuckles, feeding
only on those within the Lonicera tatarica
complex (Voegtlin. 1982. Great Lakes Ento-
mol. 15(3): 147-152). Species and cultivars
of this complex of honeysuckles have been
planted extensively across Canada and the
northern United States, and there have
probably been few insects that found a more
favorable environment waiting for them on
this continent.
Over the past 10 years, as H. tataricae has
continued to expand its range, we have re-
Fig. 1. Distribution of Hyadaphis tataricae in North America. Years indicated on the map represent first
verification of the collection of the aphid in the region, state or province.
VOLUME 90, NUMBER 2
ceived many samples for identification and
verification. As can be seen from Fig. 1,
there has been a continual expansion west-
ward, and the aphid has now crossed the
Rocky Mountains, at least into Idaho (S. E.
Halbert, pers. commun.). The aphid prob-
ably can be found in Montana,Utah and
several of the north-eastern states as well,
but to date we have no positive records from
them. An interesting aspect of the spread
has been the apparent limit in the expansion
of the aphid’s range southward. This could
be a function of the distribution of its host
plants, which are not widely planted in the
middle to southern regions of the United
States, or of unfavorable climate. Grigorov
(1965. Gradinor. Lozar. Nauk. Sofia 2(4):
493-501) noted that H. tataricae did poorly
in hot weather in Bulgaria.
The impact of this aphid has been felt
most keenly in the north central areas of the
United States and adjacent parts of Canada
where honeysuckles have been widely used
as ornamentals and in shelterbelts. Nurser-
ies, which catered to the demand for hon-
eysuckles and grew, almost exclusively, cul-
tivars related to L. tatarica, have suffered
severe losses with the result that these cul-
tivars have pretty much disappeared from
the horticultural trade. The Landscape Ar-
boretum of the University of Minnesota has
257
developed and released a cultivar that ap-
pears to be resistant to H. tataricae but has
retained most of the desirable horticultural
characteristics of the species in the L. ta-
tarica complex (Pellet et al. 1985. J. Envi-
ron. Hort. 3(2): 79-81).
The invasion by this newly introduced
aphid species has not been considered a
problem by everyone. It is well known that
the species in the L. tatarica complex escape
cultivation and become problems in native
landscapes. Initially it was hoped that H.
tataricae might effect some natural control
of these escaped, weedy honeysuckles. Un-
fortunately, this has not happened. How-
ever the intensity of attack, the dwarfing and
folding of terminal leaves, and the resulting
witches’ broom undoubtedly influences
growth rate and seed set and thus may slow
the plant’s proliferation.
We would like to acknowledge and thank
those who sent in samples for verification
and provided distributional information.
David Voegtlin, [//inois Natural History
Survey, 607 E. Peabody, Champaign, IIli-
nois 61820; Manya B. Stoetzel, Systematic
Entomology Laboratory, Agricultural Re-
search Service, U.S. Department of Agri-
culture, Beltsville, Maryland 20705.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 258-259
NOTE
A Gynandromorph of Hockeria rubra (Ashmead)
(Hymenoptera: Chalcididae)
A gynandromorph is an individual which
has both male and female characters. This
condition is often reported in the literature
and occurs in many insect orders. Gynan-
dromorphism is commonplace in Drosoph-
ila and ants. In ants, a specialized termi-
nology has been developed to denote
particular types of gynandromorphs (Don-
isthorpe. 1929. Zool. Anz. 52: 92-96; Berndt
and Kremer. 1982. Experientia 38: 798-
799). Gynandromorphism is believed to be
caused by fertilization anomalies and
meiotic abnormalities during embryogensis
though the definitive causes, at least in ants,
remain unknown (Jenkins. 1979. Genetics,
Second edition, Houghton Mifflin Co., Bos-
ton, MS; Jones and Phillips. 1985. Proc.
Entomol. Soc. Wash. 87(3): 583-586; ibid.,
1982).
While examining specimens at the United
States National Museum of Natural History
(USNM) for a revision of the genus Hock-
eria Ashmead in North America, I found a
gynandromorph of Hockeria rubra (Ash-
mead). I take this opportunity to describe
this gynandromorph because the condition
is believed to be rare in this family and may
be the first such record.
Typically, females of H. rubra are red or
orange with slender filiform antennae, pat-
terned wings, and are about 7 mm long (4—
10 mm). Males are black, have robust fili-
form antennae, clear wings, and are about
5 mm long (3-6 mm). The gynandromorph
was reared from the Western Grapeleaf
Skeletonizer, Harrisina brillians B & McD.
(Lepidoptera: Zygaenidae), a pest in which
the larvae defoliate grapes (Vitis spp.) and
two ornamental vines, Virginia Creeper
(Parthenocissus quinquefolia) and Boston
Ivy (P. tricuspidata) in the southwestern
United States and Mexico (Stern et al. 1981.
Western Grapeleaf Skeletonizer, pp, 140-
146. In Flaherty et al. [eds.]. Grape Pest
Management. Univ. Calif. Publ., Berkeley,
CA). Hockeria rubra ranges throughout the
United States and Mexico (Burks. 1979.
Chalcididae, pp. 860-879. In Krombein et
al. [eds.]. Catalog of Hymenoptera in Amer-
ica North of Mexico. Vol I. Smithson. Inst.
Press., Wash., D.C.; Halstead, in prep.).
The gynandromorph “EX. Harrisina bril-
lians pupa, coll. Chihuahua, Mexico, 10/10/
51, O. J. Smith, #A,” is as follows (see Figs.
1-4): 3 mm. Right antenna (2), scape long
and thin, 12 as long as wide, flagellum
slender and filiform, flagella 2 as long as
wide; scape, pedicel and flagella 1-2 orange,
remainder black. Left antenna (¢), scape
short and stout, 7 x as long as wide, flagel-
lum robust and filiform, flagella 1.7 = as
long as wide; antenna black except for scape
which is dark orange-brown. Right half of
occiput (dorsal view) black anteriorly (6),
orange posteriorly (2); left half of occiput
orange anteriorly (2), black posteriorly (é).
Right lateral ocellus elliptical, situated
slightly posterior to vertex (4); left lateral
ocellus round, situated on vertex (2). Right
gena orange (2), left gena black (4) with cen-
tral area orange (2). Right frons black dor-
sally (6), remainder orange (2); left frons or-
ange ventrally and dorsally (2), black
centrally (3). Pubescence on eyes similar to
that of females, less pubescent than on males.
Right eye situated more dorsad, giving a
tilted assymetry to the face in frontal view.
Pronotum, forelegs, and right hindleg or-
ange (2). Left hindleg black except for apex
of tibia and tarsus which are orange-brown
(3). Right hindfemur narrowly ovoid (2): left
hindfemur broadly ovoid (é). Forewings
clear (4). Remainder of thorax and abdomen
black (@).
VOLUME 90, NUMBER 2
\
Figs. 1-4.
\
Hockeria rubra (Ashmead). 1, Head, frontal view, of male; side view of antenna. 2, Head, frontal
view, of gynandromorph (stipuled = male, black color; nonstipuled = female, orange color). 3, Head, frontal
view, of female; side view of antenna. 4, Head, dorsal view, of gynandromorph (stipuled = male, black color;
nonstipuled = female, orange color). Scale lines 1.0 mm.
I thank D. J. Burdick, Department of Bi-
ology, California State University Fresno,
Fresno; N. J. Smith, Fresno County Agri-
cultural Commissioner’s Office, Fresno,
California; and R. D. Haines, Tulare Coun-
ty Agricultural Commissioner’s/Sealer’s Of-
fice, Visalia, California for comments on
earlier drafts of this manuscript. I thank an
anonymous reviewer for comments on a lat-
er draft of this manuscript. I thank also E.
E. Grissell, Systematic Entomology Labo-
ratory USDA-ARS, % USNM, Washing-
ton, D.C. for loaning the specimen, and the
Kings River Conservation District, Fresno,
California for the use of word processing
equipment.
Jeffrey A. Halstead, Department of Biol-
ogy, California State University Fresno,
Fresno, California 93740. Present address:
2110 N. Hayes, Fresno, California 93722.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 260-261
NOTE
Note on the Habitat of Prerostichus (Pseudomaseus) tenuis (Casey)
(Coleoptera: Carabidae) with Six New State Records
The carabid species Pterostichus (Pseu-
domaseus) tenuis (Casey) was recently re-
described and removed from synonymy with
P. luctuosus (Dejean) (Bousquet, Y. and J.
G. Pilon. 1983. Redescription of Prerosti-
chus (Pseudomaseus) tenuis (Casey), a valid
species (Coleoptera: Carabidae). Coleopter-
ists Bulletin 37: 389-396). The authors in-
cluded range maps and listed the states and
provinces from which they had seen spec-
imens (in Canada: Alberta, Manitoba, On-
tario, Quebec, Nova Scotia, Newfoundland;
in the United States: Colorado, Illinois,
Michigan, New York, New Hampshire,
Maine, Massachusetts, Rhode Island, New
Jersey). They were not, however, aware of
any differences in habitat between the two
species. They noted that P. /uctuosus is a
marsh species and speculated that the hab-
itat of P. tenuis might be the same. Most of
the specimens of the latter collected by the
authors were taken in hibernation under
rocks and logs in a deciduous forest near a
large marsh. In a later paper (Bousquet Y.
1985. Morphologie comparée des larves de
Pterostichini (Coleoptera: Carabidae): de-
scriptions et tables de déterminations des
espéces du Nord-est de ’ Amérique du Nord.
Le Naturaliste Canadien 112: 191-251),
Bousquet repeated these observations and
stated that the precise habitat of P. tenuis
remained to be discovered.
Although diagnostic differences between
P. tenuis and P. luctuosus were not widely
known until recently (Bousquet and Pilon
1983), I had been aware of them for many
years through conversations with Yves
Bousquet. Specimens from different locali-
ties and habitats were therefore kept care-
fully segregated and labelled. These speci-
mens verify that Prerostichus luctuosus is a
marsh species occurring in a variety of wet-
land habitats, a fact also noted by Larochelle
(Larochelle, A. 1975. Les Carabidae du
Québec et du Labrador. Bulletin de la Dé-
partement de Biologie du Collége Bourget,
Rigaud 1: 1-155). The sixty or so specimens
of P. tenuis (collected at thirteen localities
in Six states) indicate that this species has a
much more limited habitat and is probably
found exclusively in Sphagnum.
All of the specimens of Pterostichus tenuis
listed below were taken by treading Sphag-
num, both in large bogs and in smaller, more
isolated patches. In Pine Swamp on the
Maryland-West Virginia border, beetles
were taken from several small (ten square
feet) patches of Sphagnum on the wet grassy
shores of a pond. Other carabid species
abundant in the Sphagnum were Agonum
mutatum G. & H. and Pterostichus patruelis
(Dejean). At the Linn Run State Park in
Pennsylvania, Pterostichus tenuis was taken
in a large open bog with sundews (Drosera)
and pitcher plants (Sarracenia purpurea),
again in the company of Agonum mutatum
and Prerostichus patruelis. The specimens
of P. tenuis from Shelburne Pond, Vermont,
were taken in a similar large open bog, along
with specimens of Agonum darlingtoni Lin-
droth (a bog species). In no instances were
Pterostichus tenuis and P. luctuosus taken
together, though this should be expected to
occur occasionally. The areas of Sphagnum
preferred by P. fenuis are often very close
to marsh habitats preferred by P. /uctuosus.
There follows a detailed list of new lo-
calities and dates of collection of Prerosti-
chus tenuis. Specimens were collected by me
unless otherwise indicated. They are de-
posited in the Carnegie Museum of Natural
History, except the Ohio specimens (Harry
J. Lee private collection) and the Wisconsin
specimens collected by Walter Suter (Rob-
VOLUME 90, NUMBER 2
ert C. Graves private collection). Collector
and number of specimens are listed in pa-
rentheses after each date. All specimens were
collected by treading Sphagnum. These lo-
calities represent six new state records not
included in Bousquet and Pilon (1983). Ca-
naan Mountain at Canaan Heights, West
Virginia, is the southernmost locality in the
eastern United States from which P. tenuis
has thus far been collected. I thank Dr. Rob-
ert E. Acciavatti for his efforts in collecting
the West Virginia specimens and Dr. John
E. Rawlins for suggestions and improve-
ments on this paper.
MARYLAND. Garrett Co.: Pine Swamp,
2 km. SE Cranesville (West Virginia), July
6, 1986 (Acciavatti & Davidson, 8).
OHIO. Ashtabula Co.: Grand River Ter-
races, Morgan Township, September 27,
1987 (Lee, 2).
PENNSYLVANIA. Westmoreland Co.:
Linn Run State Park, 10 km. SE Rector,
July 27, 1981 (11), May 13, 1984 (1), July
24, 1986 (2).
VERMONT. Addison Co.: 2 km. N.
Starksboro, July 19, 1977 (1). Chittenden
261
Co.: Colchester Bog, May 30, 1976 (1); Shel-
burne Pond, Shelburne, April 14, 1974 (4),
April 12, 1975 (3), April 22, 1975 (2), July
4, 1975 (2). Franklin Co.: Fletcher, June 22,
1975 (1). Grand Isle Co.: Isle la Motte, June
18, 1975 (1).
WEST VIRGINIA. Tucker Co.: Back-
bone Mountain, 6 km. N. Hendricks, 975
m, August 15, 1986 (Acciavatti, 5); 1.5 km.
NW Canaan Heights, 1130 m, August 11,
1986 (Acciavatti, 1); Canaan Mountain at
Canaan Heights, 1130 m, July 29, 1986 (Ac-
clavatti, 6).
WISCONSIN. Kenosha Co.: Van Halter
Bog, Silver Lake, Salem, September 9, 1972
(Suter, 1), June 20, 1974 (Suter, 2), June 21,
1975 (Suter, 1). Racine Co.: Kneeland Bog,
April 23, 1966 (Suter, 1); King Bog, 5 km.
SW Dover, October 23, 1970 (Suter, 1), Au-
gust 11, 1971 (Suter, 1).
Robert L. Davidson, Section of Inverte-
brate Zoology, Carnegie Museum of Natural
History, 4400 Forbes Avenue, Pittsburgh,
Pennsylvania 15213.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 262-263
Book REVIEW
Bees and Their Keepers. Richard F.
Trump, 1987. Iowa State University
Press, Ames, Iowa. 171 pp. $17.95 Hard-
cover.
Bees and Their Keepers is a delightful book
presenting the science of beekeeping in a
novel manner: from the perspective of the
bee as well as the beekeeper. The book is
written in a simple, clear style, and is easily
readable by both the novice and the expert.
The book is illustrated by 69 excellent black
and white photographs, most apparently by
the author (absence of photo credits and
figure numbers are weaknesses).
The twenty chapter headings are thought-
provoking, eye catching ““come-ons” such
as “Those Wild Bees,” ““How to Tell a Bee
From the Keeper,” “A Sampling of Nec-
tars’ and ““Gum Robbers” to name a few.
Despite the “down-home” jargon, each
chapter is carefully organized to logically
present a major beekeeping topic to a nov-
ice, or to entertain as well as educate the
more experienced beekeeper. In most chap-
ters he begins with a historical introduction
to the development of a method or practice,
which is usually accompanied by anecdote,
commentary and personal or classroom ex-
perience.
Being a native of the midwest (Missouri,
Illinois), Richard Trump has made the book
strongly oriented to beekeeping in that re-
gion, especially Iowa and Iowa State Uni-
versity. Many comments are made about
Walter Rothenbuhler, Vic Thompson, Frank
Pellett, and others who are familiar to most
beekeepers as well as to many established,
professional beekeepers such as Glen and
Lloyd Stanley, Henry Hansen, and to sev-
eral hobbyists, such as Ann Garber, Paul
Goossen and many others—names that will
not be noticed outside Iowa, but by anec-
dote become recognized perhaps as the peo-
ple who make beekeeping such an impor-
tant part of agriculture in the U.S. today.
Of particular interest are anecdotes refer-
ring to the development of the American
foulbrood-resistant, “Brown” strain of hon-
ey bees in Iowa by Dr. Rothenbuhler. Mr.
Trump comments on observations on the
behavior of the bees of this and susceptible
strains (some based on research by Mr.
Trump) and the eventual loss of the strain.
Other interesting and noteworthy com-
ments are made on honey plants of the re-
gion (and elsewhere), importation of exotic
plants and weeds and their comparison as
nectar sources to native prairie plants, and
many comments on ecology throughout the
book.
The value of this book is its potential for
many uses: it can be an introduction to bee-
keeping for almost any novice from high
school to retirement; it can serve as a “mo-
tivating reference” for college students; es-
tablished beekeepers can read it for a source
of information and popular reading; and ex-
perts can use it as a reference, especially to
historical notes, but also to some of Mr.
Trump’s novel ideas and different perspec-
tive.
As a teacher of apiculture I greatly ap-
preciate Mr. Trump’s numerous classroom
anecdotes and even the use of student ques-
tions and replies as chapter introductions.
Mr. Trump’s experience and excellence as
a biology teacher are obvious throughout
the book. I believe other educators may
benefit from some of his ideas as I have.
A major drawback of the book is the in-
complete reference list: Mr. Trump pre-
sented many interesting points in the text
that led me to want to read more but some
citations were missing. Also I would have
liked a little more information on bee dis-
eases, and bee mites, etc., especially with
the coming of Africanized Bees and Varroa
VOLUME 90, NUMBER 2
mite to the U.S. Some modern key refer-
ences, such as Morse and Cooper’s “*Ency-
clopedia of Beekeeping,” and Morse’s ‘‘Dis-
eases and Pests of the Honey Bees’ were
absent from the list of selected books.
One error in biology on page 62 implies
that all adult bee protein is derived directly
from the food glands; actually, most adult
protein is derived directly from digested
pollen and the protein secreted from food
glands is passed on mostly to the larvae and
the queen (this may have been an error of
syntax). A few other minor errors in usage
were noted, such as the use of the term grub
for larvae (I personally prefer the use of
New MEMBERS FOR 1987
Nezha Aouad
Gerald T. Baker
Vitor Osmar Becker
Paul E. Boldt
Jeffrey R. Brushwein
John A. Chemsak
Stephen L. Clement
James C. Cokendolpher
Marcia Souto Couri
Gary Dodson
Lance A. Durden
Lester E. Ehler
Sonia Maria Lopes Fraga
J. Kenneth Grace
Harlan J. Hendricks
Aileen Nien-Hwa Hsu
Herb Jacobi
John Jenkins
James B. Johnson
Joseph W. Kamp
Scott F. Larcher
Ted C. MacRae
263
“grub” for the larvae of Coleoptera), and an
occasional incomplete sentence or misspell-
ing.
But despite these few minor errors, I thor-
oughly enjoyed reading the book: I learned
several new facts, the change in style and
point of view were refreshing, and Mr.
Trump is obviously well acquainted with
the literature of beekeeping. For the modest
price of $17.95 I believe that every serious
beekeeper should have a copy.
James W. Amrine Jr., Div. Plant Sci.-
Entomol., West Virginia Univ., Morgan-
town, WV.
Donald D. Miller
Charles Mitter
John D. Oswald
Analia C. Paggi
Robert R. Parmenter
Daniel M. Pavuk
Foster Forbes Purrington
Donald Lambert Jesse Quicke
Jay Aaron Rosenheim
John Schulte
Cristopher K. Starr
Daniel Strickman
Michael Joseph Sharley
Robert B. Trumbule
David B. Wahl
D. M. Wood
Gregory Zolnerowich
Total new members for 1987: 39
Total membership as of 8 December 1987:
580
Submitted by Geoffrey B. White, Member-
ship Chairman, 10 December 1987.
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 264-266
SocteTY MEETINGS
934th Regular Meeting—October 1, 1987
The 934th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Thomas E. Wallen-
maier in the Naturalist Center, National
Museum of Natural History, at 8 p.m. on
October 1, 1987. Twenty-nine members and
eleven guests were present. Corresponding
Secretary R. G. Robbins read the minutes
of the May meeting. R. J. Gagné distributed
reprints of the Society’s revised Bylaws,
published in Proc. Entomol. Soc. Wash. 89:
625-631. Membership Chairman G. B.
White read the names of the following ap-
plicants for membership: Vitor Osmar
Becker, Planaltina, Brazil; Lester E. Ehler,
University of California, Davis; John Jen-
kins, Michigan State University; Charles
Mitter, University of Maryland, College
Park; John D. Oswald, Cornell University;
Daniel M. Pavuk, Ohio State University,
Wooster; Donald Lambert Jesse Quicke,
University of Sheffield, England; John
Schulte, Imlay City, Michigan; Christopher
K. Starr, Department of Entomology,
Smithsonian Institution, Washington, D.C.;
Marcia Souto Couri, Museu Nacional, Rio
de Janeiro, Brazil; and Sonia Maria Lopes
Fraga, also of the Museu Nacional, Rio de
Janeiro.
President Wallenmaier noted that the cost
of the annual banquet was still under study.
He also asked that ESW’s final meeting of
this year be held on the 10th of December
so as not to conflict with the national meet-
ing of the Entomological Society of Amer-
ica.
R. J. Gagné announced the death of Hon-
orary Member Frederick W. Poos. Presi-
dent Wallenmaier called for nominations
for anew Honorary Member, who must be
unanimously approved by the Executive
Committee and by two-thirds vote of mem-
bers present at a regular meeting, in accor-
dance with Article III, Section 6, of the By-
laws. T. J. Spilman suggested that someone
write an obituary for Dr. Poos.
R. J. Gagné reminded the membership
that a nominating committee for a new slate
of officers should be formed as soon as pos-
sible. Any member who regularly attends
meetings or has a little spare time to devote
to the Society should feel free to volunteer
for a position as an officer. President Wal-
lenmaier also called for volunteers to bring
refreshments to Society meetings.
President Wallenmaier displayed a newly
published manual, Microlepidoptera from
the Sandy Creek and Illinois River Region,
by George L. Godfrey, Everett D. Cashatt,
and Murray O. Glenn, Illinois Natural His-
tory Survey Special Publication 7. J. M.
Kingsolver exhibited a new reference for co-
leopterists, The Insects and Arachnids of
Canada. Part 15. The Metallic Wood-Bor-
ing Beetles of Canada and Alaska. Coleop-
tera: Buprestidae, by Donald E. Bright, Pub-
lication 1810, Biosystematics Research
Centre, Ottawa, Ontario. Corresponding
Secretary R. G. Robbins distributed copies
of a revised manual on Lyme disease pub-
lished by Pfizer Central Research, Groton,
Connecticut.
The speaker for the evening was David
W. Inouye, Department of Zoology, Uni-
versity of Maryland. His talk was entitled
“Mutualistic Relationships between Ants
and Plants: Examples from the Rocky
Mountains, Europe, and Australia.”
Visitors and guests were introduced and
the meeting was adjourned at 9:05 p.m.,
after which refreshments were served.
Richard G. Robbins, Corresponding Sec-
retary
VOLUME 90, NUMBER 2
935th Regular meeting— November 5, 1987
The 935th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President T. E. Wallenmaier in
the Naturalist Center, National Museum of
Natural History, at 8 p.m. on November 5,
1987. Twenty-three members and four
guests were present. Minutes of the October
meeting were read and approved. Member-
ship Chairman G. B. White noted that the
Executive Committee approved the election
of Dale Parrish to emeritus status.
President Wallenmaier discussed a few
items from the recent Executive Committee
meeting. The recent death of F. W. Poos left
a vacancy in the group of three honorary
members allowed by the Bylaws. The Ex-
ecutive Committee voted unanimously to
elect another honorary member and select-
ed C. W. Sabrosky, who has been active in
the Society and is a regular attendee at the
meetings. R. J. Gagné presented a brief re-
view of Dr. Sabrosky’s career and moved
that he be selected as an honorary member.
The motion was seconded and passed. Pres-
ident Wallenmaier presented to the mem-
bers a proposed change in the Bylaws to
delete the word “June” in Art. VIII, Sec. 1,
which will allow a more accurate numbering
of the regular society meetings. The pro-
posed change will be voted upon at the next
meeting.
D. R. Smith presented the following slate
of officers for next year selected by the Nom-
inating Committee: President-Elect, F. C.
Thompson; Treasurer, N. E. Woodley; Ed-
itor, H. G. Larew; Recording Secretary, R.
G. Robbins; Corresponding Secretary, J. M.
Kingsolver; Custodian, A. M. Wieber; Pro-
gram Chairman, W. E. Steiner, Jr.; Mem-
bership Chairman, G. B. White.
A brief discussion was held concerning
the banquet expenses and lost checks. G.
Wood said that all expenses have been cov-
ered from his personal funds and that any
who wish to may send replacement checks
to him.
265
R. J. Gagné displayed a copy of the recent
issue of the Proceedings which includes a
colored plate accompanying an article on
ticks.
J. H. Fales reported on the current heavy
northward flight into Maryland of the pierid
butterfly Phoebis sennae eubule, the cloud-
less sulfur, which is the first major migration
of this butterfly since 1975. It was first ob-
served on July 8 at Plum Point, Calvert
County, and Fales noted that there appear
to be two color forms, one perhaps being a
migratory form.
R. G. Robbins reported on the Hoogstraal
Fund to support the tick collection and re-
search at the Smithsonian’s Museum Sup-
port Center, pointing out errors in the an-
nouncement in the Bulletin of the
Entomological Society of America. He also
displayed a new text, The Ixodid Ticks of
Uganda, by Matthysse and Colbo, pub-
lished by the Entomological Society of
America.
W. E. Bickley noted that the Asian tiger
mosquito, Aedes albopictus, has recently
been found in Maryland.
The speaker for the evening was Melanie
Odlum, University of Maryland. Her talk
was entitled ““An update on the Africanized
bees”
Visitors and guests were introduced and
the meeting was adjourned at 9:45 pm, after
which refreshments were served.
Paul M. Marsh, Recording Secretary
936th Regular Meeting—December 10,
1987
The 936th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Wallenmaier in the
Naturalist Center, National Museum of
Natural History, at 8:10 p.m. on December
10, 1987. Twenty-five members and five
guests were present. Minutes of the previous
meeting were read and approved.
Annual reports of officers were given by
the Treasurer, Editor, Membership Com-
266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mittee, Corresponding Secretary, Program
Chairman, and Custodian. President Wal-
lenmaier thanked the officers for a job well
done during this year.
D. R. Smith presented the slate of nom-
inees for new officers for 1988. President
Wallenmaier called for further nominations
of which there were none. A motion was
made and seconded that the slate be ac-
cepted as presented. The motion was unan-
imously passed.
President Wallenmaier again discussed the
proposed Bylaw change to delete the word
“June” from Art. VIII, Sec. 1. It was moved,
seconded, and passed to approve the change.
R. G. Robbins displayed a mosaic tick
wall plaque given as a gift to H. Hoogstraal
by an Egyptian colleague.
The speaker for the evening was Dr.
George K. Roderick, Department of Ento-
mology, University of Maryland. His talk
was entitled “Stocks, bonds, and commod-
ities: avoidance of risk for planthoppers.”
D. W. S. Sutherland reported on the En-
tomological Society of America program to
establish a national insect. Final results of
a survey show that the monarch butterfly
was the preferred insect. A brochure will be
prepared to support a bill to be put before
Congress in the near future.
P. M. Marsh announced the recent death
of long-time member and honorary presi-
dent C. F. W. Muesebeck.
President Wallenmaier thanked Mignon
Davis and the Hospitality Committee for
their efforts this year and asked for more
volunteers to bring refreshments during the
coming year. President Wallenmaier then
passed the gavel to incoming President G.
Wood. Following the introduction of visi-
tors, the meeting was adjourned at 9:20 pm.
Paul M. Marsh, Recording Secretary
REPORTS OF OFFICERS
Treasurer’s Report
SUMMARY FINANCIAL STATEMENT FOR 1987
Special
General Publications Total
Fund Fund Assets
Assets: November 1, 1986 $5295 1271 $64,909.43 $80,461.14
Total Receipts for 1987 62,389.06 TGA3932 69,528.38
Total Disbursements for 1986 64,101.50 5,997.62 70,099.12
Assets: October 31, 1987 13,839.27 66,051.13 79,890.40
Net Changes in Funds $ 1,712.44— 1,141.70 570.74—
Norman E. Woodley, Treasurer
CORRESPONDING SECRETARY'S SUMMARY OF
Major ACTIVITIES FOR CALENDAR
YEAR 1987
Letters of welcome were mailed to 32 new
members. Our computerized membership
list tonight stands at 580 (for the record, the
figure was 592 at this time last year). In
response to changes in the tax code, seven
individuals purchased life memberships,
raising to 20 the number of members in this
category. Three of our members were wel-
comed to Emeritus status: Victor E. Adler
of Laurel, Maryland; Roger O. Drummond
of Kerrville, Texas; and Dale W. Parrish of
Camp Springs, Maryland. Eight letters were
written thanking our guest speakers. Some
two dozen thank-you letters were sent to
members who contributed to our Special
Publication Fund when paying their dues.
An equal number of letters were written to
members and other persons who had re-
quested information about the organiza-
tion, functions, or publications of our So-
ciety. The postage costs of this Office totaled
$25.00.
Richard G. Robbins, Corresponding Sec-
retary
EpItor’s REPORT
This report is my fourth and final one as
Editor of the Entomological Society of
Washington. Much worth noting happened
to the Proceedings in these last four years,
and taking leave is a good occasion for a
review.
Regional journals such as ours compete
for members with a quality journal. To
compete favorably, a journal has to look
good, appear on schedule, carry a large
enough assortment of articles to interest the
readership, have a negligible backlog to en-
courage submission of good manuscripts,
and maintain a respectable scientific qual-
ity.
As to appearance, we are fortunate that
our printer, Allen Press, prints scientific
journals as a large part of its business, so
PROC. ENTOMOL. SOC. WASH.
90(2), 1988, pp. 267-268
we enjoy the benefits of that company’s con-
siderable expertise. We pay for quality pa-
per, through which one cannot read print
on the reverse side of the page. My first issue
as Editor was the Centennial issue, and I
took Manya Stoetzel’s suggestion that the
color of the cover be changed to mark the
occasion. I ordered blue, which turned out
to be so handsome that I kept it to the pres-
ent. I encouraged authors who wrote well
to send more articles. I was usually suc-
cessful in getting authors to lead a manu-
script with a topical paragraph to interest
the general reader. The journal is not only
a permanent record; it should be topical
enough so that an intelligent, well-rounded
entomologist will want to read at least the
first paragraph of every article.
In the not so remote past, an energetic
editor convinced the Executive Committee
to budget for 200 pages per number. That
number of pages makes possible a large
number of articles in a wide range of sub-
jects and may be the major reason for our
continuing growth in membership. With the
1987 volume we made another change, sug-
gested to us by Allen Press: we began a two-
column per page format, in large part to be
able to get more print into the same 200
pages.
For many years we have had a written
policy of not accepting manuscripts that
would print to more than 15 pages: how-
ever, the rule was set aside when editors
found they needed the longer manuscripts
to fill an issue. When I became Editor,
though, we had a backlog of 12-15 months,
so I refused overlong manuscripts unless
they were of a general nature. I feared we
would lose those papers of immediate in-
terest that make a journal interesting to read.
I refused two other categories of papers also:
mere lists of insects found in a very restrict-
ed area and some descriptions of new
species, of which more later. The backlog
became reduced in due course to 5-7
months. That is the lower limit ofa backlog.
A manuscript usually requires one month
for review and revision and then has to be
268
sent to press four months before it is pub-
lished. The new policy did not by itself re-
duce the backlog; some of the reduction was
certainly due to competition from journals
that have resumed timely publication dur-
ing the last several years.
The Proceedings is a refereed journal, but
I did not send out manuscripts for the pro
forma two reviews when I was confident one
would do. Some manuscripts were sent out
to three reviewers, when that was deemed
necessary. Reviewers were almost always
willing to review a paper on request and
were generally supportive and sympathetic
to the authors. Authors were mostly happy
for the criticisms. I should mention some
author services that are not widely known.
Each member-author is subsidized by
members’ dues for one-third the cost of an
article. Approximately 10% of an issue is
made up of fully-subsidized papers because
the authors are students or retired members
without funds, or authors who are not paid
to do research. Manuscripts are published
according to date of receipt, not acceptance.
In that way, an author is not unduly penal-
ized for a slow review.
Each editor invests the office with some
individuality, especially in our Society,
where the editor probably enjoys more in-
dependence than do editors of any other
regional North American journal. An editor
who does all the work should not have it
any other way. I did not worry about uni-
formity of format among papers for such
items as collection dates of specimens, so
long as data were uniform within an article.
But I did attempt to achieve uniformity of
some terminology, especially in anatomical
terms used in Diptera, for which there is
now a lingua franca. I am averse to “hu-
morous” scientific names, so refused manu-
scripts in which they were coined. Some such
names are clever, but most are silly, and I
think the practice lends taxonomy to ridi-
cule. I refused manuscripts that featured one
or more new species descriptions that did
not enlarge the generic concept and were
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
based on one or a few specimens of one sex
caught in flight. I believe such descriptions
add nothing essential to our sum of knowl-
edge.
Several titles were suggested for our
Memorr series. The topics of only two were
general enough to consider seriously, and
one was eventually accepted. It became
Memoir No. 13 by Marsh, Shaw, and Whar-
ton, a generic key to adult North American
Braconidae.
I still have in my possession the original
drawings for papers whose authors did not
request return. I will discard them on June
1, 1988, unless an author requests them by
then.
I take this opportunity to thank individ-
uals and institutions for help in producing
the Proceedings: Arly Allen and the com-
petent, helpful, and courteous staff of Allen
Press; the Publications Committee, com-
posed of D. R. Smith, T. J. Spilman, and
G. C. Steyskal, for their support and good
counsel; I thank G. C. Steyskal additionally
for checking the scientific names for each
issue; my fellow Society Officers, particu-
larly V. Blackburn, R. Robbins, G. White,
and N. Woodley, who happily and com-
petently provided information under their
purview; the Systematic Entomology Lab-
oratory for its support of this volunteer, un-
paid work with some secretarial help in typ-
ing letters, dispensation from reviewing
in-house manuscripts, and use of the copy
machine and telephone; the U.S. Postal Ser-
vice and the Smithsonian Institution mail-
room staff for generally speedy delivery and
for never having lost a piece of mail of which
I am aware. Finally, I thank the authors for
submitting manuscripts and the reviewers
for their invaluable help. My communica-
tions with the authors and reviewers were
the most gratifying part of the editorship.
And I enjoyed keeping abreast of general
entomology.
Respectfully submitted, Raymond _ J.
Gagne, Editor
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Cynipid Galls of the Eastern United States, by Lewis H. Weld oe
Cynipid(Galls ofthe Southwest. (by) Lewis He Weld Ch Rien An Tere ie ete tek Aen Ee
Both papers;on eynipid! pallss stk see lee sih TEETER EL eh ee ee
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman............. A Wa PRs Es
Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R.
Traver
A Short History of the Entomological Society of Washington, by Ashley B. Gurney.
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C.
VESTA teak LUE oh A EEE LEY EE LEER ER RL
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
SCE SANE SL ee UE OPEN ULA PGT LE, RNS ees AEE se eae SR ee a
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939_..
No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
SAPO LO As rele e eL eR U EAE eee Bt US ee a
No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
DAB IRID Too nus bate bane pear Mee Wes OLD Re UE OR Oe PER OUD as CO ee ele UE ee Ci UN Lee
No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952.0.
No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp.
NO 577 ee EE LN PAE) Be EN ee SU
No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
Mabanasiy2 90 PON LOGO ue Washer LULA C IAT Ue CR EAM LEO ANE |
No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
APY Ke AR AE AY QE RE A OA BI Rm See IDE SLICE ML ES 8 UN a Lie A OPA
No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cera-
topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 eee
No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200
TOP PHO SI! BU LAD ALe OAe LEP PCE PLL IE Se NE Bole ee yh a) EE
No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982.00...
No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp.
BO SAME TULA PCs ere sR LEM BUC PE AOR aU E MUL AULL OLAS OR AAR ULE ES ae MAREE ES igs
No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
GED Wil OA) erect bE SUR DAS ALCOR OT ELY PSUR Re AOE PE ela EL oe
No. 13, An Identification Manual for the North American Genera of the Family Braconidae (Hy-
menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 ............
15.00
11.00
12.00
10.00
11.00
18.00
5.00
18.00
Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume
to non-members and $13.00 per volume to members of the Society.
Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10
per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with
the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
NOTES
DAVIDSON, R. L.—Note on the habitat of Prerostichus (Pseudomaseus), tenuis (Casey) (Co-
leoptera: Carabidae) with six new state records
HALSTEAD, J. A.—A gynandromorph of Hockeria rubra (Ashmead) (Hymenoptera: Chalci-
VOEGTLIN, D. and M. B. STOETZEL — Hyadaphis tataricae (Homoptera: Aphididae): 10 years
after its introduction into North America
BOOK REVIEW
AMRINE, J. W., JR. Bees and Their Keepers
NEW MEMBERS FOR 1987
SOCIETY MEETINGS AND REPORTS OF OFFICERS FOR 1987
JULY 1988 NO. 3
(ISSN 0013-8797)
mu a PROCEEDINGS
=a of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON
PUBLISHED
QUARTERLY
CHAN, W. P., G. T. BAKER, and M. M. ELLSBURY —Sensilla on the larvae of four Hypera
Species (Colcoplerd sO urcHll ONIGAE) Migr 8. hs. 0 eval aire era Cia icicle Goose eels, Aahela arse yee plate 269
DREA, J. J. andR. W. CARLSON — Establishment of Cybocephalus sp. (Coleoptera: Nitidulidae)
from Korea on Unaspis euonymi (Homoptera: Diaspididae) in the eastern United States 307
EMERSON, K. C. and R. D. PRICE—A new species of Haematomyzus (Mallophaga: Hae-
matomyzidae) off the bush pig, Potamochoerus porcus, from Ethiopia, with comments on
Li GER CUE ONIG PRE Ts tis Mopeds ALE ett hie Tce Sin oem alata oa ers AM a ered Hiciay aetls ae toes 338
HEYDON, S. L. and E. E. GRISSELL—A review of Nearctic Merismus Walker and Toxeuma
Walker (Hymenoptera: Chalcidoidea: Pteromalidae) ..................0.000025 eee eee 310
KURCZEWSKI, F. E., E. J. KURCZEWSKI, and M. G. SPOFFORD—Nesting behavior
of Aporinellus wheeleri Bequaert and A. taeniolatus (Dalla Torre) (Hymenoptera: Pompil-
MATHIS, W. N.—First record of the genus Procanace Hendel from North America, with the
description’ ofa. new species (Diptera? Ganacidae)\)..). 2.6 ce ce ee eee 329
MATHIS, W. N.—First record of the shore-fly genus Placopsidella Kertész from North America
CD Iptera ae PMy AIGA) yas Mae eseyt pichd Meee a sea ake oars Bc ic Suet han ial ane ranteg edn da¥v ager Les cde eae 334
REEVES, R. M.—Distribution and habitat comparisons for Carabodes collected from conifer
branches with descriptions of brevis Banks and higginsi n. sp. (Acari: Orabatida: Carabo-
PULA) ete ren a PRR ee ce AU SEE ede aN eget Bene MMI Air ar cl days Servis ine, Seale Gude ers 373
ROBBINS, R. K.— Male foretarsal variation in Lycaenidae and Riodinidae, and the systematic
Placement OL Styx. yennals (Lepidoptera): stews vast oehe sacs bam owe otek ease aes 356
SANTIAGO-BLAY, J. A. and J. MALDONADO-CAPRILES— Observations on the true bugs
Emesa tenerrima, a possible spider mimic, and Ghilianella borincana (Hemiptera: Redu-
RnGabsEmesinacyaLOnl POCELON RICO! Patt. Kiva mom h Renn AMR A ee eM UG ita fa cot 369
SPECKER, D. R. and W. T. JOHNSON —Biology and immature stages of the rhododendron
gall midge, Clinodiplosis rhododendri Felt (Diptera: Cecidomyiidae) ...................
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1988
F. EUGENE Woon, President WARREN E. STEINER, JR. Program Chairman
F. CHRISTIAN THompSON, President-Elect GEOFFREY B. WHITE, Membership Chairman
RICHARD G. Rossins, Recording Secretary ANNE M. WIEBER, Custodian
JOHN M. KINGSOLVER, Corresponding Secretary MANYA B. STOETZEL, Delegate, Wash. Acad. Sci.
NORMAN E. WoopDLEy, 7reasurer
HirRAM G. LAREw, Editor
Publications Committee
REBECCA F. SURDICK GEORGE C. STEYSKAL
Book Review Editor
B. V. PETERSON
Honorary President
Curtis W. SABROSKY
Honorary Members
LoulsE M. RUSSELL THEODORE L. BISSELL
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tution, Washington, D.C. 20560.
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STATEMENT OF OWNERSHIP
Title of Publication: Proceedings of the Entomological Society of Washington.
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Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Wash-
ington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash-
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PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 269-287
SENSILLA ON THE LARVAE OF FOUR HYPERA SPECIES
(COLEOPTERA: CURCULIONIDAE)
Wal P. CHAN, GERALD T. BAKER, AND MICHAEL M. ELLSBURY
(WPC, GTB) Department of Entomology, P.O. Drawer EM, Mississippi State Univer-
sity, Mississippi State, Mississippi 39762; (MME) USDA-ARS, Crop Science Research
Laboratory, Forage Research Unit, P.O. Box 5367, Mississippi State, Mississippi 39762.
Abstract.—The morphology, number and distribution of the antennal, mouthpart and
body sensilla of the fourth instar larva of Hypera meles (F.) are described and compared
to those of H. nigrirostris (F.), H. postica (Gyllenhal) and H. punctata (F.); comparisons
also are made among the first, second and third instar larvae of H. meles. The number
and distribution of the antennal and mouthpart sensilla are very similar for the four
species, and they are the same for the different larval instars of H. meles. Only H. punctata
exhibits some morphological differences. The antenna has two sensilla auricillica, three
sensilla basiconica and one sensillum campaniformium. Sensilla campaniformia and sen-
silla chaetica are found on the clypeus, labrum, mandible, stipes and prementum. Six
scolopophorous sensilla are found on the mandible. Only sensilla chaetica are found on
the postmentum. The epipharynx has six sensilla basiconica on the anterior edge, and 12
sensilla basiconica and two sensilla chaetica deeper in the buccal cavity. The adoral surface
of the mala has 11 sensilla basiconica. Ten sensilla basiconica and two sensilla campan-
iformia are found on the apex of the maxillary palpus whereas eight and two of the
respective sensilla are found on the apex of the labial palpus; both palpi have one sensillum
digitiformium on the side of the apical segment. Two types of setae are found on the
body. The first type is hair-like, resembling a sensillum chaeticum, and the second type
is capitate on the first instar larva of H. me/es but clavate on the other instars of H. meles
and the fourth instar larvae of the other three species. The second type can be found on
the prodorsum, postdorsum, and spiracular area of all segments except the prothorax.
Hypera punctata has thinner sensilla aurillica and no trifurcate sensillum basiconicum on
the antenna, a different arrangement of epipharyngeal sensilla basiconica, and eight scol-
opophorous sensilla in the mandible. Also, H. punctata has a short bifurcate sensillum
chaeticum on the stipes, a shorter and thinner sensillum digitiformium on the maxillary
palpus, emarginated dorsal sensilla of mala without scythe-like projections, and smooth-
walled clavate body setae.
Key Words: | Curculionidae, Hypera spp., larvae, morphology, sensilla, setae
Larvae of North American species of the
genus Hypera feed mostly on plants in the
legume (Fabaceae) and buckwheat (Polygo-
naceae) families (Titus 1911, Kissinger
1964). In the southeastern United States
there are four introduced species of Hypera
which feed on clover, Trifolium sp., alfalfa,
Medicago sativa L. and sweet clover, Mel-
ilotus sp.: (1) clover-head weevil, H. meles
(F.); (2) lesser clover-leaf weevil, H. nigri-
rostris (F.); (3) alfalfa weevil, H. postica
(Gyllenhal); and (4) clover-leaf weevil, H.
270
punctata (F.). Hypera meles and H. postica
are known to be serious pests on alfalfa and
clover. The life history, biology and feeding
preferences of these four Hypera species are
summarized by Chapin and Oliver (1981
and references cited within). The larvae have
distinct feeding preferences: larvae of H.
meles and H. nigrirostris feed on the flowers
and developing seeds, larval H. postica feed
inside leaf buds and on leaves, and larval
H. punctata feed on leaves. To understand
their feeding preferences, we need to know
the various types of cephalic sensilla that
are involved in the feeding behavior and
physiology. However, the morphology,
number and distribution of the antennal and
mouthpart sensilla which play an integral
role in feeding behavior are documented
only for the alfalfa weevil, H. postica (Bland
1983).
It is common for more than one of the
four above-mentioned species to infest the
same field together. The ability to differ-
entiate the four species and their different
life stages is critical for decision making in
pest management. A key to the fourth instar
larvae of 14 Hypera spp. which includes
these four introduced species is available
(Anderson 1948). However, the couplet that
separates H. meles from H. nigrirostris de-
pends on the color of the head capsule and
the relative length of the abdominal post-
dorsal setae (Anderson 1948: couplet 4).
These characters are hard to determine in
preserved specimens.
Dyar’s rule (1890) often is applied to re-
hiably differentiate the different larval in-
stars of these weevils (Detwiler 1923, Mail-
loux and Pilon 1975, Tower and Fenton
1920). The different larval instars of the four
above-mentioned species have been de-
scribed (Detwiler 1923, Titus 1911, Tower
and Fenton 1920). However, no other def-
inite characters are reported that would en-
able the differentiation of the different larval
instars of the same species, except in alfalfa
weevil (Gurrea-Sanz and Cano 1983).
This study provides information on (1)
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
the morphology, number, and distribution
of the antennal and mouthpart sensilla on
the fourth instar larvae of the lesser clover-
leaf weevil, alfalfa weevil, and clover-leaf
weevil and also those of the four larval in-
stars of the clover-head weevil; and (2) the
morphology of their body setae.
MATERIALS AND METHODS
Laboratory-reared fourth instar larvae of
H. meles, H. nigrirostris, H. postica, and H.
punctata were placed in boiling water for
30 s, transferred to Perfix® overnight, rinsed
five times with distilled water, sonicated in
a dilute detergent solution for 2 s, and rinsed
three times with distilled water; or, they were
fixed in Poly/LEM with 4% Triton® X-100,
sonicated in the fixative for 2 s, and rinsed
five times in distilled water. The mouth-
parts were dissected from the head and
placed in a flow-through cell that was made
from size 00 BEEM® capsules and embed-
ding bag material. Specimens were osmi-
cated 24-48 h in 4% aqueous OsO,, rinsed
in distilled water, dehydrated in a graded
ethanol series, and either critical point dried
in CO, or air dried from pentane. Specimens
were attached to aluminum stubs with dou-
ble stick tape or conducting silver paint and
sputter-coated with gold/palladium. They
were examined with a Hitachi HHS-2R or
JEOL JSM-35CF scanning electron micro-
scope at 20 kV.
The reduced silver staining method of
Schafer and Sanchez (1976) was used to
demonstrate porous sensilla. Stained larval
mouthparts were dehydrated in graded eth-
anol series, then cleared in clove oil for 3
days and rinsed for 5 min in xylene. The
specimens were mounted in Permount® for
examination with a compound microscope.
A modified Essig’s technique (Essig 1948)
was used to study body setae. Decapitated
fourth instar larvae were cut open either
along the horizontal or sagittal plane. They
were cleared in Essig’s fluid for 25 min at
50°C and then overnight at room temper-
ature. The clearing procedure was repeated
VOLUME 90, NUMBER 3
until a clear specimen was obtained. Larval
skins were rinsed ten times in distilled water,
dehydrated in a graded ethanol series,
cleared in clove oil for 3 days and rinsed
for 5 min in xylene, then mounted in Per-
mount® and examined with a compound
microscope. The nomenclature of the setae
is based on the terminology of Anderson
(1947) and that of the sensilla on Schneider
(1964) and Snodgrass (1935). Descriptions,
figures and measurements given are those
of the fourth instar larva of H. me/es unless
otherwise stated.
RESULTS
The antenna and mouthparts did not dif-
fer from the published description (Ander-
son 1948). The former is located on the side
of the epicranium between the anterior
stemma and the base of the mandible. Lar-
vae of the four Hypera spp. have hypogna-
thous mouthparts, but usually the labrum
is treated as the most dorsal mouthpart and
the labium the most ventral. The labrum
joins to the clypeus which is connected to
the frons and underneath is the epipharynx.
A pair of mandibles is situated behind the
labrum. The ventral mouthparts consist of
the labium and a pair of maxillae.
The average head capsule widths for the
fourth instar larvae used in this study were
0.47 mm (n = 18) for H. meles, 0.39 mm
(n = 3) for H. nigrirostris, 0.56 mm (n = 2)
for H. postica, and 1.2 mm (n = 28) for H.
punctata (Figs. 1-4). The average head cap-
sule widths for the various instar larvae of
H. meles are 0.2 mm (n= 5) for the first
instar, 0.25 mm (n = 5) for the second instar
and 0.33 mm (n = 5) for the third instar.
Antenna.—The antenna has seven sen-
silla on a short segment (Figs. 7, 8, 13, 16).
The anteriorly located sensillum basiconi-
cum is surrounded partially by three smaller
sensilla basiconica, two sensilla auricillica
and one sensillum campaniformium. The
larger sensillum basiconicum is ca. 17 wm
long, and 8 um wide at the base where some
plugged pores with diameters of ca. 400 to
271
600 nm are situated (Fig. 12). The outer
surface of this sensillum is pitted as seen in
the SEM micrographs (Fig. 9) and silver
staining preparations (Fig. 7 insert). Frac-
tured sensillum has a reticulated inner sur-
face and the wall is ca. 0.75 to 1 wm thick
(Fig. 11). Terminal pores are apparent on
two of the three smaller sensilla basiconica
which are ca. 2 to 3 um long and | to 1.5
um wide at the base (Fig. 10). The sensillum
basiconicum between the sensilla auricillica
is trifurcate and does not have a terminal
pore (Fig. 14). The sensillum auricillicum
is leaf-like, ca. 8 to 10 wm long, 2 wm wide
at the base and constricted to ca. 1 wm half-
way up the sensillum where a pore of ca.
250 nm in diameter is located (Fig. 15). It
widens again towards the tip to form a blade
and the widest part measures ca. 2.5 um.
On the other side of the blade there is a
midrib-like structure and in some speci-
mens the blade is emarginated. All of these
sensilla lack a socket. A sensillum campan-
iformium, an oval depression with a raised
rim of ca. 1.5 um in diameter, is located
near one of the sensillum auricillicum (Fig.
7). It often is not discernible when the cu-
ticle is distorted during preparation. In H.
punctata the sensillum auricillicum appears
to have a narrower blade and the sensillum
basiconicum between the sensilla auricillica
is not trifurcate (Fig. 16).
Labrum, clypeus and epipharynx. — There
are three sensilla chaetica, ca. 14 to 50 um
long and 2 to 3 um wide at the base, and
one sensillum campaniformium on each side
of the bilobed labrum (Figs. 1-6). Four sen-
silla chaetica, ca. 4 to 9 um long and | to 2
um wide at the base, are situated in the me-
dial notch. A median sensillum campani-
formium is located above the notch. Two
sensilla chaetica, ca. 10 to 15 um long and
2 to 3 um wide at the base, are located on
each side of the clypeus near the epistomal
suture. One sensillum campaniformium is
located between them. The sensilla chaetica
are smooth-walled, attenuated, straight or
slightly curved, and each has a socket. Some
272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6.
sensilla campaniformia possess a depres-
sion of ca. 200 nm in diameter in the center.
On each side of the outer edge of the epi-
pharynx there are three sensilla basiconica
(Fig. 17). They are ca. 10 to 21 um long and
4 to 5 um across the widest part. These sen-
silla are slightly flattened, sunken and con-
stricted at the base, and directed mesad.
Deeper in the buccal cavity are 12 sensilla
basiconica and two sensilla chaetica. Mi-
1-4, Head. 1, H. meles (frontal view). 2, H. nigrirostris (frontal view). 3, H. postica (frontal view).
4, H. punctata (frontal view). 5, Clypeus and labrum, H. postica. 6, Labrum, H. meles. Arrows = sensilla
campaniformia.
crotrichia are present and they are directed
toward the pharynx. The two larger sensilla
basiconica are ca. 10 to 12 wm long, 2 to 3
um wide at the base and sit in a cavity (Figs.
19, 21). Their tips may be blunt or sharp.
The other ten sensilla basiconica are ar-
ranged in two fields of five on each side of
the midline (Fig. 17). In each field, four of
the sensilla are grouped together and the
other one is further away from the midline.
VOLUME 90, NUMBER 3 273
\
| A
-_— . * y Ye
14 opm 4 015 10 fi
Saeed ¢. <y
Figs. 7-16. Antenna. 7, H. meles (insert shows pitted surface of the larger sensillum basiconicum, silver
staining, scale bar = 25 um). 8, H. nigrirostris. 9-11, H. meles. 9, Larger sensillum basiconicum with pitted
outer surface. 10, Sensillum basiconicum with terminal pore. 11, Fractured larger sensillum basiconicum shows
reticulate inner surface. 12, Plugged pores at the base of the larger sensillum basiconicum, H. punctata. 13, H.
postica. 14, Trifurcate sensillum basiconicum, H. meles. 15, Sensillum auricillicum shows a pore, H. meles. 16,
H. punctata.
274 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
S——=—— —/?
‘
;
Figs. 17-22. Epipharynx. 17-19, H. meles. 17, Low magnification. 18, High magnification of the small
sensilla basiconica. 19, Central region. 20, H. nigrirostris. 21, H. postica. 22, H. punctata.
The same are found in H. nigrirostris (Fig. like, ca. 300 to 400 nm wide, and sit inside
20) and H. postica (Fig. 21) but in H. punc- an oval depression of ca. 1 wm in diameter.
tata the five sensilla are grouped together A pore about 100 nm in diameter, which
(Fig. 22). These sensilla are short, nipple- appears plugged, is located at the tip of these
VOLUME 90, NUMBER 3
sensilla basiconica (Fig. 18). Two sensilla
chaetica ca. 7 um long and 1.5 um wide at
the base, are located between these sensilla
basiconica and the microtrichia (Figs. 17,
19). They are similar to the sensilla chaetica
on the labrum.
Mandible.—The anterior surface bears
two sensilla chaetica which are ca. 10 um
long and 1.5 um wide at the base (Figs. 23,
28). There are three sensilla campaniformia:
one is located on the center of the dorsal
surface, one on the anterior surface near the
dorsal articulation and one at the boundary
between the anterior and posterior surface
(Fig. 23). In silver-stained preparations, the
sensillum campaniformium appears as a ca-
nal in the heavily sclerotized cuticle (Fig.
30). Each mandible of H. meles, H. nigri-
rostris and H. postica has two apical teeth.
Inside each tooth there are three scolopoph-
orous sensilla (Fig. 24). Each mandible of
H. punctata has four apical teeth (Fig. 28).
Inside the fourth (ventral most) tooth are
five scolopophorous sensilla (Fig. 29)
whereas inside the second tooth there are
only three (Fig. 26). The scolopophorous
sensillum appears as a long canal extending
from the internal cavity of the mandible
into the teeth.
Maxilla.—Each maxilla consists of a fused
cardo and stipes with the mala on the adoral
side, and bears a two-segmented palpus on
the other side (Figs. 31, 32). No sensilla are
found on the cardo. The stipes has four sen-
silla chaetica, ca. 5 to 75 wm long and 3 to
5 um wide at the base, and four to five sen-
silla campaniformia. Two longer sensilla
chaetica, each accompanied by a sensillum
campaniformium, are found on the aboral
side of the stipes: one near the cardo and
one near the palpus. Two shorter sensilla
chaetica are found on the ventral surface
near the palpus and the shortest one, ac-
companied by a sensillum campanifor-
mium, is closer to the mala. In H. punctata
the shortest sensillum chaeticum is bifur-
cate. Another sensillum campaniformium
is situated on the aboral side of the articu-
275
lating membrane between the palpus and
the stipes. Itis a short cone or dome situated
in an oval depression of ca. 2 um in di-
ameter (Fig. 32 insert). The sensillum cam-
paniformium located on the dorsal surface
of the stipes where it joins the epicranium
is found only in H. meles and H. nigrirostris
but not in the other two species.
The basal segment of the palpus has one
short sensillum chaeticum, ca. 3 um long
and 1.5 um wide at the base, and two sen-
silla campaniformia on the ventral surface
(Fig. 32). The second segment has one sen-
sillum digitiformium on the aboral surface,
one sensillum campaniformium on the ven-
tral surface adjacent to the sensillum digi-
tiformium and a group of 12 sensilla on the
apex (Figs. 32-36). The sensillum digitifor-
mium is ca. 12 um long, 3 um wide, rod-
shaped, smooth-walled, socketed and lies in
a groove. In some preparations a pore can
be found on the distal tip. In H. punctata,
the sensillum digitiformium is much short-
er and thinner when compared to the size
of the second palpal segment (Fig. 31 insert)
whereas in the other three species it is al-
most as long as the second palpal segment
(Fig. 32). The 12 sensilla on the apex consist
of two sensilla campaniformia and ten sen-
silla basiconica. The sensillum campanifor-
mium isa short cone ca. 500 nm in diameter
and sits in an oval depression of ca. 700 nm
in diameter on an elevated base. The rim
of the depression is lower on the outer edge
thus exposing the cone. One sensillum cam-
paniformium is found facing the ventral
surface and the other facing the adoral sur-
face (Figs. 33-36). The sensilla basiconica
are ca. 1.5 to 2 wm high and 0.5 to 2 um
wide at the base, cone to peg-like, and with-
out a socket. In some of the sensilla basi-
conica, the terminal pore is surrounded by
a lip and in the others by finger-like projec-
tions (Fig. 37). The centrally located sen-
sillum basiconicum is larger than the rest
and only lightly stained by silver just below
the tip, whereas the other sensilla basiconica
were deeply stained (Fig. 38). It does not
276 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tl
Figs. 23-30. Mandible. 23, Anterior and dorsal surface, H. meles. Arrows = sensilla campaniformia. 24,
Scolopophorous sensilla, H. nigrirostris. 25, Posterior surface, H. postica. 26, Scolopophorous sensilla, H. punc-
tata. 27, Sensillum campaniformium, H. meles. 28, Anterior surface, H. punctata. 29, High magnification of
scolopophorous sensilla, H. punctata. Note only four of the five sensilla are visible. 30, Sensillum campanifor-
mium at the boundary between the anterior and posterior surface, silver staining, H. punctata.
VOLUME 90, NUMBER 3 aM fa)
Figs. 31-39. Maxilla. 31, Dorsal view, H. meles (insert shows the distal palpal segment, H. punctata, scale
bar = 50 um). 32, Anterior-ventral view of the palpus, H. me/es (insert shows sensillum campaniformium at
the junction of the palpus and stipes, scale bar = 2.5 wm). 33-39, Apex of the palpus. 33, H. meles. 34, H.
nigrirostris. 35, H. postica. 36, H. punctata. 37, High magnification of sensilla basiconica and sensillum cam-
paniformium on the ventral region, H. meles. 38, Sensilla basiconica stained at the tips, silver staining, H.
punctata. Note sensillum campaniformium is not stained (arrow). 39, Larger sensillum basiconicum with a
plugged pore near the base, H. postica.
278 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
VOLUME 90, NUMBER 3
appear to have a terminal pore. However,
a plugged pore which is ca. 300 nm in di-
ameter, is found at the base of this sensillum
(Fig. 39). It may be the ecdysial pore.
The adoral surface of the mala bears 11
sensilla and the ventral surface bears only
one sensillum campaniformium near the
apex (Fig. 47). Along the dorsal margin of
the adoral surface extending into the buccal
cavity are six sensilla basiconica, whereas
in the central region and along the ventral
margin of the distal adoral surface, there are
five sensilla basiconica (Figs. 40-47). These
11 sensilla project mesad. Anderson (1947)
called them the dorsal and ventral setae of
mala, respectively. These sensilla range from
ca. 6 to 37 wm long, and ca. 2 to 3 wm wide
at the base. In H. meles, H. nigrirostris and
H. postica, they have a smooth wall and
some of the dorsal sensilla of mala bear ir-
regular scythe-like projections at the tip (Fig.
42). In H. punctata, these sensilla measure
ca. 6 to 48 um long, and ca. 3 to 11 um wide
at the base and possess emarginated walls.
Their tips are rough and have no scythe-like
projections (Fig. 47). In optical sections of
the silver staining preparation these sensilla
appear to have a thick wall (Fig. 47 insert).
In all four species these malar sensilla have
sockets. A terminal pore is apparent on the
two shorter medial ventral sensilla of mala
(Fig. 45) and they were stained by silver at
the tips (Fig. 43).
Labium.— The labium consists of the pre-
mentum and the postmentum separated by
the premental sclerite. A pair of one-seg-
mented labial palpi is situated on each side
of the prementum. There are one pair of
sensilla chaetica, ca. 10 um long and 2.5 um
wide at the base, and one pair of sensilla
campaniformia, ca. 2.5 um in diameter, on
279
the prementum near the edge of the buccal
cavity (Figs. 48, 49). Another pair of similar
sensilla chaetica and sensilla campanifor-
mia are located between the palpi, and a
third pair of sensilla chaetica which are ap-
proximately three times as long are near the
premental sclerite. One sensillum campan-
iformium is found on the base of the aboral
side of the palpus (Fig. 49). Posterior to this,
on each side of the premental sclerite one
short sensillum chaeticum which is ca. 2.5
um long and 1.5 wm wide at the base, is
found adjacent to the postmentum and also
accompanied by a sensillum campanifor-
mium (Fig. 49). Both sensilla campanifor-
mia are ca. | to 1.5 wm in diameter. The
former is dome-shaped, whereas the latter
is crateriform. The palpus has one sensillum
digitiformium on the dorsal surface (Fig.
56), two sensilla campaniformia on the ven-
tral surface (Fig. 52), and a group of ten
sensilla on the apex (Fig. 48). Unlike the
sensillum digitiformium on the maxillary
palpus, this one is wider than long, measures
ca. 2 um long and 3 um wide, and does not
seem to have a socket (Fig. 57). Although
the number of sensilla is different from the
apex of the maxillary palpus, the spatial ar-
rangement and morphology of the sensilla
are very similar. Seven sensilla basiconica
and two sensilla campaniformia partially
surround a central larger sensillum basicon-
icum (Figs. 50, 51, 53, 54, 55).
There are three pairs of sensilla chaetica
on the postmentum. Two of these are sit-
uated laterally and one pair medially. The
anterior-most pair is the shortest, ca. 15 um
long and 2 um wide at the base. The pair
adjacent to them is the longest, ca. 40 um
long and 6 wm wide at the base. The medial
pair are ca. 24 wm long and 4 um wide at
—
Figs. 40-47.
Mala. 40-43, H. meles. 40, Mala (ventral view). 41, Dorsal sensilla of mala (arrows), silver
staining. 42, High magnification of dorsal sensilla of mala. 43, Medial ventral sensilla of mala, silver staining.
Arrow indicates stained tip. 44, H. nigrirostris. 45, Ventral sensilla of mala (arrows), H. meles. Note the position
of the two medial sensilla as compared with Fig. 43. 46, H. postica. 47, H. punctata (insert shows the thick-
walled dorsal sensilla of mala, silver staining, scale bar = 50 um).
280 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
i \asc
in
Figs. 48-55. 48, Prementum, H. meles. 49, Premental sclerite, H. meles. 50-55, Labial palpus. 50, Apex,
H. meles. 51, Apex, H. nigrirostris. 52, Sensilla campaniformia on the ventral surface (arrows), silver staining,
H. meles. 53, Apex, H. postica. 54, Apex, H. punctata. Arrows = sensilla campaniformia. 55, Larger sensillum
basiconicum on the apex with a plugged pore near the base, H. meles.
VOLUME 90, NUMBER 3
the base. The base of the postmentum is
covered with asperites similar to those found
on the body.
Body.—The number and distribution of
setae on the body are the same for the fourth
instar larvae of H. meles, H. postica, and
H. punctata. They are similar to the de-
scription on Pissodes strobi (Peck) (Ander-
son 1947) but have more setae, ca. 10 wm
long and 2 um wide at the base (Fig. 59), in
the following regions:
1. Two setae on the side between the
pronotum and the epicranium.
2. Three setae on the side between the pedal
area and epicranium adjacent to the
postmentum.
3. Two setae on the spiracular area and one
seta anterior to the pedal area of the
meso- and metathorax.
4. One seta on the spiracular area of the
first to the seventh abdominal segment.
Moreover, the anal lobe (tenth abdominal
segment) has no setae. The differences in
the relative sizes of the setae are similar to
those previously reported by Anderson
(1948) except that the postdorsal seta 2 of
H. postica on the fifth and sixth abdominal
segments is clavate instead of attenuate
(Anderson 1948: figs. 20, 21).
In H. meles two morphological types of
setae are present on the body. The first type
is smooth-walled and resembles a sensillum
chaeticum. It is straight or slightly curved
and varies from ca. 10 to 200 um long and
2 to 7 wm wide at the base (Fig. 59). The
second type 1s clavate and can be found on
the prodorsum, postdorsum and spiracular
area of all segments except the prothorax.
Both types of setae have sockets. A unique
variation of the second type is found in the
first instar larva. It is capitate, smooth-
walled, ca. 70 to 230 um long, 3 to 8 um
wide at the base and the distal bulb is ca. 7
to 10 um in diameter (Fig. 58). The longer
ones are found on the last three abdominal
segments. In later instars, clavate setae have
overlapping plates on the distal end and a
281
smooth middle piece (Figs. 60-62). On the
last three abdominal segments, some of the
clavate setae are replaced by long sensilla
chaetica-like setae which are ca. 160 um
long and 8 um wide at the base.
The first type of setae is found on all four
Hypera spp. The clavate seta has the same
appearance in H. nigrirostris and H. postica,
but in H. punctata it does not have over-
lapping plates and the whole seta appears
smooth (Fig. 63).
DISCUSSION
The number and distribution of the an-
tennal and mouthpart sensilla are very sim-
ilar for the four species, and they are the
same for the different larval instars of H.
meles. Only H. punctata exhibits some
morphological differences.
The morphology, number and distribu-
tion of the antennal and mouthpart sensilla
of H. postica reported here are different from
those previously reported by Bland (1983)
in the following aspects. Bland (1983) did
not mention that the sensillum basiconicum
(sensillum 4— Bland 1983) between the sen-
silla auricillica is trifurcate. In both the
maxillary and labial palpi he did not men-
tion the sensilla digitiformia and sensilla
campaniformia found on the side, and called
all sensilla on the apex of both palpi, sensilla
basiconica. But, I suggest that the two sen-
silla campaniformia be named as such be-
cause they resemble those in the wireworms
(Bellamy 1973, Doane and Klingler 1978),
adult Dendroctonus ponderosae Hopkins
(Whitehead 1981) and adult /ps typogra-
phus L. (Hallberg 1982). In the last two
species tubular bodies are present in the sen-
silla, which suggests that they are mechano-
receptors. On some of the ventral sensilla
of the mala (galea— Bland 1983) the shape
of the tip is variable rather than trifurcate
as reported by Bland (1983). The sensilla
found in these four Hypera spp. also resem-
ble those on the rice weevil larvae, Sitoph-
ilus oryzae (L.) in their distribution pattern.
However, the latter has three to four more
282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ws
Nea 60g
Figs. 56-63. 56, Sensillum digitiformium on the dorsal surface of the labial palpus, H. punctata. 57, High
magnification of the sensillum digitiformium similar to Fig. 56. 58, Capitate seta, first instar larva, H. meles.
59, Short sensilla chaetica-like setae (arrows) on the prothorax near the epicranium, H. meles. 60-63, Clavate
seta. 60, H. meles. 61, H. nigrirostris. 62, H. postica. Scale bar for Figs. 61-62 = 25 um. 63, H. punctata. Scale
bar = 50 um.
VOLUME 90, NUMBER 3
sensilla on the antenna, and sensilla auril-
lica are absent. It also has three less sensilla
basiconica on both the mala and apex of the
labial palp (Speirs et al. 1986).
Sensilla auricillica are common on the an-
tennae of adult Lepidoptera. They are re-
ported to be chemoreceptors (Flower and
Helson 1974, Faucheux 1984), temperature
and humidity receptors (Subchev 1980), or
receptors for selecting oviposition sites
(Flower and Helson 1976). These sensilla
are quite variable in shape (Subchev 1980).
They are perforated and different from those
found in this study. On the antenna of the
adult cave beetle, Aphaenops cryticola Lin-
der, the sensilla auricillica which are called
“sensilla basiconica inflata’ are considered
to be chemoreceptors (Juberthie and Maz-
zoud 1977). Bland (1983), however, sug-
gested that the sensillum auricillicum in H.
postica may be simply flattened mechano-
receptors.
The pitted surface and the reticulate inner
wall structure suggest that the larger sensil-
lum basiconicum of the antenna is a sin-
gle-walled, multiporous sensillum. This is
similar to the “lobe membraneux”’ of Speo-
phyes lucidulus Delar larva (Corbiere 1969)
and the antennal sensory appendix of Cre-
nicera destructor (Brown) larva (Scott and
Zacharuk 1971). Plugged pores on the base
of the antennal sensory cone of Aedes aegypti
(L.) larva, which serve as attachments of
dendritic sheath (Zacharuk et al. 1971), are
also found on these four Hypera spp.
The sensilla basiconica on the apex of the
maxillary and labial palpi are believed to
be multiporous because they are only stained
slightly on the tip by the silver stain and a
terminal pore is not apparent. The presence
of plugged (ecdysial) pores at the base may
indicate the possibility of absence of a ter-
minal pore (Zacharuk 1980). In adult Jps
typographus, single-walled sensilla that oc-
cur on the maxillary and labial palpi which
have a system of pores penetrating the apex,
are considered to be both gustatory and ol-
283
factory in function (Hallberg 1982). Ther-
moreceptor cells are known to bind to mul-
tiporous double-walled chemoreceptors in
Periplaneta americana (L.) (Altner et al.
1977). The function of these sensilla basi-
conica found in this study may be primarily
olfactory.
The sensilla basiconica on the antenna,
epipharynx, mala, maxillary and labial pal-
pi, which possess a terminal pore (unipo-
rous) may serve as gustatory receptors (Za-
charuk 1980). They can also be bimodal,
both mechanoreceptor and gustatory recep-
tor, like the lateral sensillum of the galea in
larval Entomoscelis americana Brown
(Mitchell 1978, Mitchell et al. 1979), uni-
porous pegs of the maxilla and labium in
adult Dendroctonus ponderosae (Whitehead
1981) and the terminal pore sensilla in adult
Ips typographus (Hallberg 1982). The lip
and finger-like projections around the pore
may regulate the pore opening to expose the
dendrites to or conceal them from stimu-
lants (Blaney and Chapman 1969). The nip-
ple-like sensilla basiconica on the epiphar-
ynx of Ctenicera destructor larva are also
found as two groups of five sensilla in which
Zacharuk (1962) termed the ‘oral plate or-
gan’ and suggested that they were gustatory
receptors.
The trifurcate sensillum basiconicum
found on the antenna, the six sensilla basi-
conica found on the anterior edge of the
epipharynx, the pair of sensilla basiconica
found in front of the nipple-like sensilla ba-
siconica on the epipharynx, and the dorsal
sensilla of mala, exept the medial two, do
not possess a terminal pore and their lu-
mens do not pick up the silver stain. In the
colorado potato beetle larvae, Leptinotarsa
decemlineata (Say) the longer sensilla on the
edge of the galea are believed to be mecha-
noreceptors protecting the two central con-
tact chemoreceptors (Mitchell and Schoon-
hoven 1974). Some non-porous sensilla with
inflexible sockets found on the antenna of
Locusta migratoria (L.) (Altner et al. 1981)
284
and Drosophila spp. (Altner et al. 1983), and
also on the maxillary palpus of Periplaneta
americana (Altner and Stetter 1982) are
known to be thermo- and hygroreceptive.
The information gathered for these sensilla
basiconica in this study is not enough to
ascribe a particular function.
Sensilla campaniformia and sensilla
chaetica are believed to be mechanorecep-
tors (McIver 1975). The form of sensilla
campaniformia vary from simple depres-
sions (Fig. 27) to elaborate dome-shaped
organs (Figs. 32 insert, 37). Pringle (1961)
proposed that specialization of arthropod
proprioceptors arose by slowly increasing
restriction of conditions, such as their ori-
entation and the thickness of their cuticle,
producing the strain. This is illustrated by
the sensilla campaniformia on the halteres
of the muscid flies (Pringle 1948) and Dro-
sophila melanogaster Meigen (Chevalier
1969). Zill and Moran (1981) found that the
responses of the tibial sensilla campanifor-
mia in Periplaneta americana are related to
their position and cap orientation. Thus, it
is possible that the simple-structured sen-
sillum campaniformium like those on the
prementum may respond to one form of
cuticular stress while the more elaborate ones
that are situated between the palpus and
stipes may respond to another form.
In larval Ctenicera destructor, the sensil-
lum digitiformium on the labial palpus is
found to be responsive to mechanical stim-
ulations (Zacharuk et al. 1977). In the max-
illary palpi of Agabas bipustulatus (L.) and
Hydrobius fuscipes (L.), the sensilla digiti-
formia are reported to be hygro- and ther-
moreceptor because they do not possess a
special socket and tubular body, and the
outer dendritic segment divides into several
branches in the shaft (Guse and Honomichl
1980). However, these two kinds of sensilla
only differ in morphology by the possession
of a socket. It is possible that the sensilla
digitiformia found in this study are mecha-
noreceptors.
The scolopophorous sensilla are similar
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
to those found in larval Speophyes sp. (Cor-
biere-Tichane 1971). Zacharuk and Albert
(1978) found that in elaterid larva, the scol-
opophorous sensillum responds electro-
physiologically to an outward deformation
or bending of the mandibular teeth, there-
fore they serve as proprioceptors monitor-
ing the stress and deformation in biting and
chewing. In Hypera spp. these scolopopho-
rous sensilla probably also function simi-
larly while the sensilla campaniformia and
sensilla chaetica on the surface serve as pro-
prioceptors and exteroceptors, respectively.
In these four Hypera spp. a general pat-
tern, in which a larger sensillum basiconi-
cum is surrounded by a group of other sen-
silla, is found on the antenna, and apexes
of the maxillary and labial palpi, but the
number of surrounding sensilla varies. The
number of sensilla on the apex of the max-
illary palpus 1s greater than the number on
the apex of the labial palpus. Both condi-
tions are true in larval Sitophilus oryzae
(Speirs et al. 1986) and larval Lyctus brun-
neus (Stephens) (Iwata and Nishimoto
1981). In Tribolium spp. larvae, the apex
of the maxillary palpus has one more sen-
sillum than that of the labial palpus (Ryan
and Behan 1973). The same conditions are
also found in the adult Dendroctonus pon-
derosae (Whitehead 1981) and Jps typogra-
phus (Hallberg 1982). The functional im-
plication of such a pattern, if there is any,
is unknown.
Zacharuk (1962) reported that the num-
ber, distribution and structure of the ce-
phalic sensilla among different genera and
species of larval elaterids are very similar,
and the minor differences may be related to
differences in habitat. In Tribolium spp. lar-
vae, Ryan and Behan (1973) found that there
is no difference in number and distribution
of sensilla on the last instar of two species.
In this study, a similar situation is found.
Hypera punctata is the only species among
the four to show some morphological dif-
ferences. The similarity in the number and
distribution of antennal and mouthpart sen-
VOLUME 90, NUMBER 3
silla among the four Hypera spp. indicates
that the difference in feeding preferences
among the fourth instar larvae is probably
due to physiological differences either at the
receptor level or at the central nervous sys-
tem integration level. The exact function of
the sensilla can only be determined by ex-
tensive electrophysiological and ultrastruc-
tural studies. The findings of this study pro-
vide the basis for such experiments.
The capitate sensillum is also found in
the first instar of H. postica (Bland 1983).
The body setae or sensilla, including the
capitate and the clavate sensilla, are be-
lieved to be mechanoreceptors. Studies on
the function of larval body sensilla are
scarce. Chapman (1982) in his review men-
tioned that Henig (1930) showed that sen-
silla on the thoracic tarsi of the caterpillar,
Agrochola lota (Clerck) is equipped with a
single neuron which suggests their role in
mechanoreception.
In exopterygota, the number of antennal
and mouthpart sensilla of different nymphal
instars varies considerably as illustrated by
Periplaneta americana (Schafer and San-
chez 1973) and Locusta migratoria (Chap-
man 1982 and references cited within). In
endopterygota, these characters are not
highly variable. In Coleoptera, the carnivo-
rus larvae of Macrodytes spp. have more
sensilla basiconica in the third instar than
the first (Hamon 1961). Different instar lar-
vae of the powder-post beetle, Lyctus brun-
neus, a xylophagous species, exhibit a dif-
ference in the shape of the apical sensilla of
the maxillary palpus and in the length of the
antennal segments, but there 1s no mention
of the number of sensilla found in each lar-
val instar (Iwata and Nishimoto 1981). Our
study shows that the number of sensilla on
the antenna and mouthparts of H. meles, a
phytophagous insect, remain the same
throughout larval development. This is in
agreement with the similar feeding prefer-
ence of the different larval instars. All larval
instars feed on flowers, ovaries and devel-
oping seeds of the host (Tippins 1957).
285
No additional diagnostic characters were
found that would enable differentiation of
the fourth instar larvae of H. meles, H. ni-
grirostris and H. postica. Hypera punctata
can be identified easily because of the larger
size and number of mandibular teeth (An-
derson 1948: couplet 2). Revision cannot
be made at this stage. The distribution of
body setae of the different larval instars may
provide an alternative to using measure-
ments of head capsule width, to differentiate
the instars as was proposed for alfalfa weevil
larva by Gurrea-Sanz and Cano (1983).
ACKNOWLEDGMENTS
This paper is assigned No. 6784 of the
Mississippi Agriculture and Forestry Ex-
periment Station. We thank B. Perrigin for
typing the manuscript. Mention of trade-
mark or proprietory products does not con-
stitute a guarantee or warranty of the prod-
ucts by the U.S. Department of Agriculture
and does not imply approval to the exclu-
sion of other products.
List of Abbreviations
(A) = Antenna. (C) = Clypeus. (Cr) = Cardo. (E) =
Epicranium. (f) = Finger-like projections. (F) = Frons.
(L) = Labium. (Lb) = Labrum. (Lp) = Labial palpus.
(M) = Maxilla. (Ma) = Mala. (Md) = Mandible. (Mp)
= Maxillary palpus. (p) = Pore. (Pm) = Prementum.
(Ps) = Premental sclerite. (SA) = Sensillum auricilli-
cum. (SB) = Sensillum basiconicum. (SC) = Sensillum
campaniformium. (SCh) = Sensillum chaeticum. (SD)
= Sensillum digitiformium. (Si) = Stipes. (SS) = Scol-
opophorous sensillum. (St) = Stemma.
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PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 288-293
A NEW SPECIES OF BRACON (HYMENOPTERA: BRACONIDAE)
PARASITIC ON EOREUMA LOFTINI (DYAR)
(LEPIDOPTERA: PYRALIDAE)
R. A. WHARTON AND D. L. J. QUICKE
(RAW) Department of Entomology, Texas A&M University, College Station, Texas
77843; (DLJQ) Department of Zoology, University of Shefheld, Western Bank, Sheffield
S10 2TN, United Kingdom.
Abstract. — A new species of Bracon(Hymenoptera, Braconidae) is described. This species
is parasitic on stem-boring Pyralidae (Lepidoptera) in grasses, and has been reared during
biological control programs directed against Eoreuma loftini (Dyar). Unusual features,
not previously associated with the genus Bracon, are described.
Key Words:
An undescribed species of Bracon was
reared during the course of a biological con-
trol program directed against the Mexican
rice borer, Eoreuma loftini (Dyar). It is de-
scribed here to provide a name for research-
ers working on stem borers, and to identify
several unusual features not previously re-
ported for Nearctic Bracon species. This
species has been reared only from E. /oftini
in grasses of the genera Setaria, Cynodon,
and Sorghum.
Terminology for the description follows
van Achterberg (1979) and Quicke (1987),
except that the total length of the ovipositor
has been measured rather than just the por-
tion extending beyond the metasoma. Mea-
surements presented only as ratios or ranges
are based on five individuals, with ratios
representing median values. Ranges fol-
lowed by a mean and standard deviation in
parentheses are based on 10 individuals.
Bracon rhyssaliformis
Quicke & Wharton, NEw SPECIES
Figs. 1-11
Males.—Length of body 2.5-5.0 mm and
of fore wing 2.1-4.0 mm.
Bracon, Braconidae, Biocontrol, stem borer, Pyralidae
Antennae with 38-45 flagellomeres. Ter-
minal flagellomere strongly acuminate (Fig.
3), 3.0 times longer than maximally wide.
Penultimate flagellomere |.8-2.4 times
longer than wide. Median flagellomeres | .7—
2.0 times longer than wide. First flagello-
mere 1.05-1.20 times and 1.10-1.25 times
longer than the 2nd and 3rd respectively,
the latter being 2.0-2.3 times longer than
wide. Hypoclypeal depression dorsally
strongly rounded, bordered dorsally by a
distinct lamelliform carina. Clypeus sepa-
rated from face dorsally by a narrow shallow
sulcus. Height of clypeus: inter-tentorial
distance : tentorio-ocular distance = 1:2.6:
1.7. Face moderately densely short setose
laterally, more or less glabrous medially;
smooth and shiny between the punctures at
the bases of the setae except for the malar
space which is broadly imbricate. Width of
head : shortest distance between eyes : height
of eye = 1:0.55:0.46. Eyes moderately
densely short-setose medially, glabrous lat-
erally (Fig. 1). Frons hardly impressed be-
hind the antennal sockets, with a well-de-
veloped mid-longitudinal sulcus. Vertex,
temples and occiput sparsely setose. Ocelli
VOLUME 90, NUMBER 3
Figs. 1-2.
small, shortest distance between posterior
ocelli : narrower diameter of elliptical pos-
terior ocellus :shortest distance between
posterior ocellus and eye = 1:0.7:2.1. Head
subparallel immediately behind eyes (Fig.
2).
Fore wing (Fig. 11). Veins CC +SC + R
and 1-SR forming an angle of approxi-
mately 65°. Vein 1-SR + M straight. Length
of veins r:3-SR:SR1 = 1:2.4:6.2; SR1 2.13-
3.08 (2.54 + 0.32) times longer than 3-SR;
3-SR 1.13-1.43 (1.27 + 0.10) times longer
than 2-SR. Vein r-m straight; vein 2-M dis-
tinctly curved. Vein 2-SR + M usually rath-
er long, 0.60-1.04 (0.82 + 0.12) times length
of m-cu. Veins 2-CU and 3-CU forming an
angle of approximately 75°. Vein cu-a often
(50% of specimens) interstitial, otherwise
antefurcal.
Hind wing (Fig. 11). Apex of vein C +
SC + R with | thickened bristle (hamule).
Vein Ir-m short, SC + R1 1.76-3.14 (2.29
+ 0.43) times longer than Ir-m. Vein 2-SC
+ R distinctly longitudinal. Base of wing
evenly densely setose.
Claws with acutely pointed basal lobes.
Length of fore femur : tibia: tarsus = 1:1.2:
1.5. Length of hind femur : tibia : basitarsus
Scanning electron micrographs of head, Bracon rhyssaliformis. 1, facial view. 2, dorsal view.
= 1:1.9:0.6. Hind tibia (Fig. 7) extraordi-
narily swollen for its whole length, 5.3-6.5
times longer than maximally deep, without
a distinct longitudinal, lateral furrow. Hind
basitarsus 5.1-6.6 times longer than deep.
Fore tibia with a longitudinal row of stout
setae interspersed with finer ones (Fig. 8).
Mesosoma (Figs. 4-6) smooth and pol-
ished. Pronotum in dorsal view long, bi-
sected by a deep, usually smooth, transverse
sulcus; this sulcus usually connected to an-
terior margin by a short, median longitu-
dinal groove. Mesonotum weakly decli-
vous, nearly bare, with patch of 5-10 setae
near base of notaulus and less than 15 setae
extending along each notaulus to the pos-
terior margin. Notauli unsculptured, deep
anteriorly, evanescent posteriorly. Prescu-
tellar pit narrow, containing 3-9 short, lon-
gitudinal ridges. Scutellum usually (70% of
specimens) with small median pit anterior-
ly. Metanotum (Fig. 4) often (40% of spec-
imens) with complete median longitudinal
carina. Propodeum usually (60% of speci-
mens) with rugae or carinae along midline,
forming an irregular longitudinal ridge, but
propodeum completely smooth in 30% of
specimens.
Figs. 3-6. Scanning electron micrographs, Bracon rhyssaliformis. 3, terminal flagellomere with acuminate
tip. 4, dorso-lateral, oblique view of metanotum showing median carina. 5 & 6, dorsal view of mesosoma
showing effect of angle of view on appearance of certain sculptural features.
VOLUME 90, NUMBER 3
Figs. 7-10. Scanning electron micrographs, Bracon rhyssaliformis. 7, hind femur and swollen hind tibia of
male. 8, spinose setae of fore tibia. 9 & 10, metasoma of male.
292
Fig. 11.
Fore and hind wing; scale = 2.0 mm.
First metasomal tergum approximately as
long as posteriorly wide, with distinct though
sub-lamelliform dorso-lateral carinae; pos-
terior 0.6—-0.7 of Ist tergum rugose and dis-
tinctly elevated medially. Second and 3rd
metasomal terga (Fig. 10) separated by a
broad, carinate suture. Second tergum with
narrow, relatively smooth, triangular plate
baso-medially, apex of triangle extending as
a carina to or nearly to the posterior margin;
median triangle centered within a larger,
raised, triangular area usually delimited lat-
erally by distinct, weakly converging, lateral
grooves; entire tergum rugoso-striate. Terga
3-5 (Fig. 9) completely rugose to rugoso-
striate, with metasomal sculpture weaker
posteriorly. Remaining terga often finely or
only incompletely sculptured.
Color: orange; gena, palps and fore wing
stigma often yellow; pedicel, flagellum, hind
tibia, and 5th tarsus of mid, hind, and some-
times fore leg dark brown to black; wings
usually uniformly infumate; 60% of speci-
mens with small dark spot on temple ad-
jacent eye, and varying amount of dark
markings around ocellar field.
Females.—Similar to males except as fol-
lows:
Length of fore wing 2.5—-4.3 mm. Anten-
nae with 36-48 flagellomeres. Median flag-
ellomeres 1.4—1.6 times longer than wide.
Clypeus short, inter-tentorial distance ap-
proximately 3.4—4.0 times height of clype-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
us. Ocellar triangle even smaller than in
male, with shortest distance between pos-
terior ocellus and eye approximately 2.5
times shortest distance between posterior
ocelli (ocellar measurements highly vari-
able). Length of fore wing veins r:3-SR:SR1
= 1:2.2:5.6; SR1 2.30-2.88 (2.53 + 0.17)
times longer than 3-SR; 3-SR 1.20-1.29
(1.23 + 0.03) times longer than 2-SR, these
ratios thus less variable than in males.
Length of hind femur: tibia: basitarsus =
1:1.7:0.5. Hind tibia not swollen, 10.5-12.1
times longer than deep. Hind basitarsus 4.2—
5.3 times longer than deep. Scutellum usu-
ally (70%) without small median pit. Meta-
notum with median carina nearly always
well developed. Propodeum with at least
some sculpture along midline in 90% of
specimens. Metasomal sculpture variable,
often (40% of specimens) considerably re-
duced posteriorad 3rd segment. Second
metasomal segment with median triangle
often weakly defined due to absence of dis-
tinct longitudinal depression on either side
of median triangle; larger median triangular
area usually set off laterally by a pair of
strongly converging grooves. Ovipositor
(total length) 1.7-1.9 times longer than me-
sosoma, with small, sharp subapical node
dorsally and well-defined serrations ven-
trally at tip; ovipositor sheath 1.1—1.2 times
longer than mesosoma. Color as in male,
but only 2 of the specimens examined with
a black spot on the temple; ovipositor sheath
black, ovipositor red.
Holotype ¢.—U.S.A.: Texas, Hidalgo
County, TAES Annex, 2 miles north of
Mercedes, ex. Setaria (Poaceae), 19.i1x.1984,
H. W. Browning, in U.S. National Museum
of Natural History (= USNM).
Paratypes.— MEXICO: Nuevo Leon,
Monterrey (Marin), 2-3.vii.1982, J. W.
Smith, Jr. & F. Bennett, Texas A&M Uni-
versity (= TAMU) Quarantine Number
82015, TAMU Voucher Number 208 (6 4,
4 2) reared from E. /oftini ex. Sorghum hal-
apense. U.S.A.: Texas, same data as holo-
type (6 6, 8 2); Hoblitzelle Farm, 5 miles
VOLUME 90, NUMBER 3
north of Mercedes, reared from Eoreuma
loftini (Dyar) on Bermuda grass (Cynodon),
15.vii.1984, H. W. Browning (2 6, 4 9); 4
miles north of Mercedes, ex. Eoreuma lof-
tini in Bermuda grass, 29.v.1985, R. Pfan-
nenstiel (2 2); 5 miles northwest of Weslaco,
1.vi.1983, C. W. Melton (1 2). Paratypes
deposited in TAMU Collection, USNM,
Canadian National Collection, and British
Museum (Natural History).
Diagnosis.— The enlarged hind tibia of the
male readily separates rhyssaliformis from
all previously described species of Bracon.
This species runs to oenotherae Muesebeck
in the latest revision of the Nearctic species
of Bracon (Muesebeck 1925). It is also
somewhat similar to mellitor (Say). As in
oenotherae and mellitor, the metasoma is
extensively sculptured, the propodeum is
unsculptured except along the midline, and
the fore wing stigma tends to be yellow.
However, females of both oenotherae and
mellitor have more extensive dark markings
on the hind legs and the petiolar sculpture
in both sexes is less rugose than in rhyssal-
iformis. Bracon mellitor also has a longer
Ovipositor and oenotherae a relatively
shorter second tergum.
Discussion.—The enlarged hind tibia of the
male, fore tibial spines, and petiolar mor-
phology place rhyssaliformis in a somewhat
isolated position amongst the species of
Bracon known from the Nearctic Region.
Since this region has not been well-studied,
and several species are as yet undescribed,
further discussion regarding the relation-
293
ships of rhyssaliformis would be premature.
Despite the presence of some doryctine
characteristics (notably the fore tibial
spines), rhyssaliformis is clearly a bracon-
ine, and readily fits the characterization of
Bracon given by Quicke (1987). Diagnostic
features for the genus include the 5-seg-
mented maxillary palps; upper margin of
mandible abutting lower margin of face
without formation of a groove; scape longer
dorsally than ventrally; fore and hind wing
as in Fig. 11; and absence of an occipital
carina, a prepectal carina, and a sculptured
sternaulus.
ACKNOWLEDGMENTS
We wish to thank F. Bennett, J. W. Smith,
Jr., and H. W. Browning for making this
material available for study, and for pro-
viding the rearing records. Approved as
TA#23276 by the Texas Agricultural Ex-
periment Station. This work was supported
in part by the Rio Grande Valley Sugar
Growers, Inc.
LITERATURE CITED
Muesebeck, C. F. W. 1925. A revision of the parasitic
wasps of the genus Microbracon occurring in
America North of Mexico. Proc. U.S. Natl. Mus.
67: 1-85.
Quicke, D. L. J. 1987. The Old World genera of
braconine wasps (Hymenoptera: Braconidae). J.
Nat. Hist. 21: 43-157.
Van Achterberg, C. 1979. A revision of the subfamily
Zelinae auct. (Hymenoptera, Braconidae). Tidschr.
Ent. 122: 241-479.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 294-306
NESTING BEHAVIOR OF APORINELLUS WHEELERI BEQUAERT
AND A. TAENIOLATUS (DALLA TORRE)
(HYMENOPTERA: POMPILIDAE)
FRANK E. KurRCZEWSKI, EDMUND J. KURCZEWSKI, AND MARGERY G. SPOFFORD
(FEK, MGS) Environmental and Forest Biology, State University of New York College
of Environmental Science and Forestry, Syracuse, New York 13210; (EJK) 2921 Reed
Street, Erie, Pennsylvania 16504.
Abstract. —The ecology and nesting behavior of Aporinellus wheeleri were studied from
late May through early October 1979-1987 in a field containing small hills of gravel and
shale and an abandoned gravel pit, both SE of Erie, Erie County, PA, and in a patch of
gravelly soil at the edge of an active quarry E of Auburn, Cayuga County, NY. Nests in
sand or gravel comprised short cylindrical burrows with terminal cells, whereas those in
gravel or crushed shale were simple conical depressions. Wasps provisioned their nests
with various Salticidae, predominantly immature Pellenes borealis, and, rarely, Thomisi-
dae (Xysticus spp.).
Additional observations were made on 4A. taeniolatus nesting in bare sand behind a
beach at Presque Isle State Park, Erie County, PA during July-September 1969-1978.
This species constructed relatively long, cylindrical burrows with terminal cells and pro-
visioned with P. borealis. Behavioral similarities between 4. wheeleri and A. taeniolatus
included searching for prey, paralysis, cachement, transport, position of prey in the cell,
and oviposition site. Manner of burrow excavation and closure was dictated by the kind
of soil in which the wasps nested. The two species often differed in these behavioral
activities.
A comparison of the geographic distribution, altitude, habitat (soil type), flight period,
adult external morphology, and nesting behavior of 4. wheeleri and A. taeniolatus suggests
that the two taxa have separate gene pools in the eastern United States.
Key Words: | Nesting behavior, Pompilidae, spider wasps, Aporinellus, Salticidae, prey,
sibling species, biosystematics
The genus Aporinellus contains eight
species in North America. The six species
in the Fasciatus and Apicatus groups are
easily recognized in the male sex, using gen-
italic characters. Females belonging to these
groups are very similar and in some cases
cannot be separated reliably. Slight differ-
ences in head shapes appear to separate the
females of some species in the Fasciatus
group, but this character is difficult to use
without prior experience. In the Apicatus
group the females of some species may be
separated by small differences in the lengths
of antennal segments (Evans 1951, 1966).
In the Taeniolatus group the reverse is
true. Males of A. taeniolatus (Dalla Torre),
the correct name for A. taeniatus (Kohl) (A.
S. Menke 1980, pers. comm., see Kurczew-
ski and Kurczewski 1987), and A. wheeleri
Bequaert are indistinguishable by the cri-
teria of external morphology and termi-
nalia. However, females of the two taxa are
VOLUME 90, NUMBER 3
separated readily in the eastern United States
by body coloration, pubescence, and fore-
tarsal comb length (Evans 1951, Kurczew-
ski and Kurczewski 1987). In the extreme
southern and western U.S. and Mexico the
situation is complicated by much intraspe-
cific variation in 4. taeniolatus, specifically
coloration of body and pubescence, width
of temples, and number of comb-spines on
the forebasitarsus (Evans 1951, 1966). Two
and three distinct color forms of A. taeni-
olatus females have been collected at a sin-
gle locality in Mexico (Evans 1966). In the
eastern United States only 4. wheeleri ex-
hibits a black and red body with mostly
rufous legs, but in the western U.S. several
forms of A. taeniolatus demonstrate varying
amounts of rufous coloration on the body
and legs (Evans 1951, Wasbauer and Kim-
sey 1985).
Species of Aporinellus occur in sandy or
gravelly areas, feed as adults on honeydew
and flowers, and store paralyzed Salticidae,
less often Oxyopidae or Thomisidae, as food
for their larvae (Krombein 1979). Studies
on the biology of the Nearctic species are
limited. Prey records exist for A. basalis
Banks, A. completus Banks, A. medianus
Banks, A. taeniolatus (Dalla Torre), and A.
yucatanensis (Cameron) (Evans 1951, 1959,
Evans and Yoshimoto 1962, Krombein
1959, 1961, 1964, Kurezewski and Kurc-
zewski 1968a, b, 1973, Peckham and Peck-
ham 1898). Information on nesting is scant
and available for only 4. basalis (Evans
1959), A. completus (Krombein 1961), A.
medianus (Peckham and Peckham 1898,
Krombein 1959, Kurczewski and Kurczew-
ski 1968a), 4. taeniolatus (Kurczewski and
Kurczewski 1973) and A. yucatanensis
(Krombein 1959).
Nothing is known about the nesting be-
havior of A. wheeleri (Krombein 1979). De-
spite the relative abundance of 4. taeniola-
tus (Evans 1951, 1966), little is known about
its nesting behavior. Evans (1951) collected
a female with the salticid Pellenes (Habro-
nattus) calcaratus (Banks) near Jacksonville
295
FL. Krombein (1964) noted cache and prey
(Pellenes sp.) of a female near Lake Annie
FL. Kurczewski and Kurczewski (1973) re-
corded cache, tumulus size, entrance and
burrow dimensions of one nest, and prey
type and prey weight ratios of two females
at Weymouth, NJ. They reported a 2 Pel-
lenes (H.) agilis (Banks) and an imm. P. (#.)
viridipes (Hentz) as prey from this locality.
The following observations on the ecology,
nesting behavior, and prey preference of A.
wheeleri and A. taeniolatus expand consid-
erably the amount of biological information
about this genus and emphasize the ecolog-
ical and behavioral differences between the
two species (see Kurczewski and Kurczew-
ski 1987). Related species of Pompilidae are
often readily separated by habitat and prey
type rather than specific sequencing of be-
havioral components (Evans and Yoshi-
moto 1962).
Aporinellus wheeleri Bequaert
Observations on A. wheeleri were made
during nine field seasons: 21 July-2 Oct.
1979, 9 July—27 Sept. 1980, 15 June-14 Sept.
1981, 1 June-14 Sept. 1982, 6 July—6 Sept.
1983, 11 June-19 Sept. 1984, 19 June-20
Sept. 1985, 26 May-27 July 1986, and 28-
31 July 1987, ina field containing small hills
of gravel and shale in Wintergreen Gorge
Cemetery, 1.6 km SE of Erie, Erie County,
PA (site I), and in a large, abandoned, over-
grown gravel pit containing fine and coarse
gravel mixed with sand, 2.4 km SE of Erie
(site II) (Figs. 1, 2), and during 4-19 Sept.
1985 in a gravelly area at the edge of an
active quarry 2.3 km E of Auburn, Cayuga
County, NY (site III) (Fig. 3). Approxi-
mately 50 wasps nested annually at site II
during 1979-1982 in flat areas of gravel and
in gravelly hills, both nearly devoid of vege-
tation (Fig. 2), but their numbers decreased
to just a few individuals during 1984-1987.
Twelve males of 4. wheeleri were col-
lected in Erie County, PA during 21 July-
14 Aug. 1979, 7 Sept. 1980, 21 July—3 Sept.
1981, 18-28 June 1982, 11 June-28 Aug.
Fig. 1. Habitat of Aporinellus wheeleri and various other species of ground-nesting Pompilidae and Sphecidae,
2.4 km SE of Erie, Erie County, PA.
Fig. 2. Nesting site of Aporinellus wheeleri in Erie County, PA. Females excavated and provisioned nests in
the bare gravel in foreground.
VOLUME 90, NUMBER 3
Fig. 3.
the gravel in foreground but avoided the peripheral sandy areas.
1984, 24 May-24 July 1985, and 29 May
1986 while attempting to mate with nesting
females of A. wheeleri. The males trailed the
females on the ground and in flight as the
latter searched for prey and prospective
nesting sites.
Females of 4. wheeleri searched for spi-
ders from 0950 to 1805 h (EDT) among the
sparse vegetation within and at the periph-
eries of the nesting areas. Wasps flushed spi-
ders from upright and decumbent vegeta-
tion and pursued them rapidly on the
ground, either by running or in flight. The
wasps caught and stung spiders once or a
few times in rapid succession in the under-
side of the cephalothorax after which they
cleaned and malaxated the prey. Some large
prey were stung, examined, malaxated, and
then stung again. Using their mandibles, the
wasps then grasped the spiders by the face
or pedipalps, dragged them backwards to a
position on an upright plant, grass blade,
297
Nesting site of Aporinellus wheeleri in Cayuga County, NY. Females dug and provisioned nests in
dead twig, raised stone or soil ridge, and
released them dorsal side upward. Most prey
were placed on the leaves of low, upright
plants up to 8.3 cm above ground level. One
paralyzed spider was cached vertically on
the side of a stone. Wasps often cleaned
their antennae with the strigilis following
cachement. The entire stinging-cleaning-
malaxation-cachement-cleaning sequence
often took less than | min.
One wasp was observed searching for
jumping spiders in their silken retreats be-
neath stones. She flushed a relatively large
female salticid from underneath a stone,
pursued it on the pebbles, caught it, stung
it in the underside of the cephalothorax, ex-
amined it with her antennae for 15s, malax-
ated it, and then stung it again. She pulled
it to a small Melilotus alba Desr. plant,
placed it 2 cm above the ground, and began
a burrow 67 cm away. She abandoned this
burrow, rebuffed the advances of a male,
298
and then a few minutes later, stung a second
prey and cached it similarly. She began and
abandoned a second burrow. Unlike the first
spider, this prey recovered from the effects
of the sting in less than 15 min. The wasp
repeated the process of capturing and sting-
ing a third, fourth, and fifth salticid, each
time caching the prey 2-4 cm above ground
ona white sweet clover plant, and beginning
and abandoning a burrow. These prey re-
covered rapidly from the effects of the sting,
the last one taking only 2 min. This suggests
that the wasp’s venom became less effective
with subsequent captures. All five spiders
were adult females of Pellenes (Habronat-
tus) borealis. They weighed (wet) 38, 54, 42,
39, and 40 mg (wasp wt. 7 mg). Why the
wasp did not use these prey for provisioning
nests is unknown. Perhaps she was unmat-
ed, which seems unlikely in view of the
above-mentioned rebuff of a male, or the
spiders were too heavy to transport toa nest.
She had little difficulty in transporting them
the short distances to their caches. The spi-
ders were not used for adult feeding.
Selection of a nesting site often required
15 min and some females searched for over
30 min. In Erie County PA females sampled
many gravelly and sandy areas, usually dug
in gravel, but frequently abandoned such
burrows. The wasps often selected tiny
patches of compacted sand between gravel
particles, and, although these sites were often
abandoned, most females eventually dug
burrows in such areas. Wasps near Auburn,
NY invariably nested in gravel even though
loose sand was only 50-75 cm away (Fig.
3):
Females excavating burrows used their
mandibles in unison to loosen the soil and
forelegs alternately to throw the loosened
sand and gravel underneath and behind the
abdomen. One female weighing 9 mg, when
beginning to dig, pulled backward with her
mandibles pebbles weighing | 1 7-182 mg to
a distance of 25 mm from the opening. Be-
tween periods of digging some wasps ex-
amined their prey by walking or flying to
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
the cache 11 cm—5 m away, and a few fe-
males moved the spider closer to the ex-
cavation at this time. Such periodic exam-
inations may have been related to the rather
light paralysis and rapid recovery of some
of the prey. Nest excavation usually took 4—
28 (x = 13.8, n = 24) min except for one
wasp that spent 43 min constructing a nest
in gravel. Much of this time was spent in
dragging various-sized pebbles backwards
with the mandibles onto a crescent-shaped
tumulus.
Upon finishing their burrows females ex-
ited headfirst, turned 1-3 circles in front of
the opening—possibly a form of orienta-
tion—interspersed with antennal cleaning,
or entered and exited 1-3 times, and then
walked in a mostly straight line toward the
cached prey. Such females occasionally
paused and cleaned their antennae en route
to the prey. One of 31 wasps apparently
became disoriented while en route and al-
ternated short, quick, circular flights with
walking.
Provisioning females paused at the cache,
examined the spider with their antennae,
grasped it dorsal side upward with the man-
dibles by the face, or pedipalps in the case
of one male spider, and flew to the ground.
Transport to the nest then involved walking
rapidly backwards in a rather straight line,
interspersed with occasional turns and flights
forward, in either case retaining the grasp
of the prey. Some wasps released their prey
one to several times on the gravel, dorsal
side upward, walked to the nest, flew or
walked back to the spider, and resumed
transporting it toward the nest. Eventually
the wasps released the spiders directly in
front of the entrances, then went inside,
turned around, came out headfirst, reposi-
tioned the spiders, and, using the mandi-
bles, pulled in the prey by its spinnerets. If
prey recovered from the effects of the sting
while cached or during transport, they were
stung again in the cephalothorax and some-
times malaxated.
One to 7 min later, females reappeared
VOLUME 90, NUMBER 3
Fig. 4. Closed nest of Aporinellus wheeleri showing circular mound of pebbles (center) covering entrance.
in the openings, throwing sand and gravel
backwards with the forelegs which moved
alternately, and tamping the soil in the bur-
rows with the end of the abdomen. Some
wasps periodically turned 180° and checked
the amount of fill. After 4-15 min of closing,
a female began retrieving small pebbles with
her mandibles and placed these in the bur-
row and on the filled entrance. She inter-
mittently cleaned her antennae, threw small
pebbles or sand onto the area with the fore-
legs, and spent much time rearranging the
surface by placing pebbles atop bare sand
near the entrance. After 5.5-22 (x = 11.8,
n = 28) min of such closing activity females
paused, cleaned their antennae, and flew
away. The resulting closure comprised a cir-
cular mound of fine gravel and pebbles, 15-
30 mm in diameter (Fig. 4). Such a closure
prevented the lightly paralyzed spider from
escaping from the cell.
In Ene County, PA the burrow structure
reflected the substrate in which the nest was
built. Nests in mixed sand and gravel had
cylindrical burrows, 20-28 mm (x = 25, n
= 23) long, that terminated in cells 14-23
mm (X = 19, n = 23) beneath the ground
surface, including cell height (Fig. 5). The
entrances and burrows were 2.5-4.0 mm
and 2.5-3.5 mm in diameter, respectively.
Nests in gravel comprised conical depres-
sions, 9-22 mm (X = 17, n = 40) deep (Fig.
5). Near Auburn, NY, on the other hand,
nests in gravel contained cylindrical bur-
rows, 18-26 mm (X = 22, n = 6) long, which
ended in cells 15-23 mm (¥ = 17, n = 6)
beneath the surface. Three cells were 6-7
mm wide and 10-12 mm long. In either
substrate at all localities the paralyzed spi-
der was placed in the crude cell dorsal side
upward, either sideways or head toward the
entrance. The eggs of A. wheeleri were 1.25-
2.00 mm long and placed on the undersides
of the spiders’ abdomens near the base, per-
pendicular to the longitudinal axis of the
body (Fig. 6), but two eggs were attached
300
=e OG
Vs
6
Fig. 5. Longitudinal sections of nests of Aporinellus
wheeleri in sand (top) and gravel (bottom), as viewed
from the side.
Fig.6. Egg (arrow) of Aporinelllus wheeleri attached
to base of abdomen of Pellenes borealis, ventral view.
about midway from the abdominal base,
obliquely and lateroventrally.
Prey taken from provisioning females of
A. wheeleri or their nests in Erie County, PA
comprised the following species of Saltici-
dae: Pellenes (Habronattus) borealis, 8 ad.
46, 43 imm. 66, 14 ad. 92, 81 imm. 99, 12
imm.,; P. (H.) decorus (Blackwall), 4 ad. 42,
5 imm. 64, 3 ad. 22, 10 imm. 92, 3 imm.,;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Eris sp., 21mm. °°; Tutelina elegans (Hentz),
1 ad. 9; Metaphidippus protervus (Walck-
enaer), | ad. 4, 5 ad. 22; and Metaphidippus
sp., |imm. The capture of | 6 Yysticus ferox
(Hentz) (Thomisidae) is noteworthy be-
cause it is the first record of a crab spider
taken as prey by this species of pompilid.
Spiders taken from A. wheeleri cells near
Auburn, NY included P. (H.) borealis, 3 99:
P. (H.) decorus, 1 2, 1 imm. &; and, X. sp.,
1 imm. Prey were larger in late summer and
late spring (overwintering?) and comprised
relatively more adults, and were smaller in
midsummer and included a high proportion
of immatures.
Many of the spiders recovered from the
paralysis of the wasp’s sting shortly after
being removed from the cells. Wasps weigh-
ing (wet) 8-11 mg (X = 8.8, n = 5) were
particularly ineffective in paralyzing larger
salticid spiders weighing (wet) 22-34 mg (x
= 27.0, n = 5), whereas one small wasp (7
mg) completely and permanently paralyzed
an immature crab-spider weighing only 12
mg. Larger prey had to be restung periodi-
cally by the wasps to effect partial paralysis
and acquiescence. The spider weighing 34
mg recovered quickly from repeated wasp
stings, was placed in a vial, spun a silken
retreat, and fed on a small fly introduced
thereto. In several nests excavated 48 h after
closure, adult spiders bearing wasp eggs were
found enclosed in silken retreats and were
extremely lively. Smaller, immature spiders
with wasp eggs had no such surrounding
retreats, were lethargic, and appeared to be
partly paralyzed. The construction of silken
retreats by the larger, adult spiders may be
related to the ineffectiveness of the wasps’
stings (relative amount of venom?), which
enabled the spiders to behave normally as
if not stung at all.
After feeding on the spider in the cell the
wasp larva spun a cocoon of saliva, silk and
sand and gravel particles (Fig. 7). Some
wasps emerged by chewing through their
own cocoon and then the surrounding spi-
der retreat (Fig. 8). Two nests from Auburn,
VOLUME 90, NUMBER 3
iMETRIC 1
Fig. 7.
are In mm.
Fig. 8.
spider retreat. Units are in mm.
NY provisioned on 17 Sept. 1985 and ex-
cavated on 19 Sept. 1985 gave rise to two
males on 26 May 1986, leaving ample time
for the production of another, mid- or late
summer generation of 4. wheeleri at this
latitude.
A few provisioning females of 4. wheeleri
were trailed by females of the cleptopara-
sites Sphenometopa tergata (Coquillett) and
Hilarella hilarella (Zetterstedt) (Sarcophag-
idae: Miltogramminae). The flies also pur-
sued wasps searching for places to dig and
paused nearby on raised stones or other
perches during burrow construction. De-
spite this activity, none of the cells or prey
examined contained maggots. Several spi-
ders were stolen from the wasps’ caches by
workers of the ant Formica subsericea Say.
The completed nest of one female was ex-
cavated partly by a female of Evagetes par-
vus (Cresson) (Pompilidae), but this clep-
toparasite could not penetrate fully the
301
METRIC 1
Tt
.
Completed cocoon of Aporinellus wheeleri showing construction of silk, saliva and soil particles. Units
Emergence hole in cocoon of Aporinellus wheeleri showing inner wasp cocoon and surrounding silken
stone-filled closure and abandoned the ex-
cavation.
Aporinellus taeniolatus (Dalla Torre)
Five females of A. taeniolatus were ob-
served nesting at Presque Isle State Park,
Erie County, PA on 7 July 1969, 7, 12 Sept.
1976, and 23 Aug. 1978, in bare sand be-
hind Beach 10 (Fig. 9). These wasps searched
for spiders on the sparse beach vegetation,
pursued them on the sand, and captured and
stung them in a manner similar to 4. whee-
leri. The paralyzed spiders were dragged
backwards on the sand and cached above
ground on vegetation.
The wasps sampled many areas of sand
before beginning to dig. Females used the
mandibles to loosen the sand crust and fore-
legs alternately to remove loosened sand
from the burrow, backing out periodically
to deposit the load in a crescent-shaped tu-
mulus partly surrounding the entrance. Two
*
yy. ae
Wash
ae
Fig. 9.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Nesting site of Aporinellus taeniolatus, Beach 10, Presque Isle State Park, Erie County, PA. Females
excavated and provisioned nests in the bare sand in foreground.
such tumuli were 25-45 mm long and 40-
55 mm wide. One burrow took 27 min to
dig. Exiting females made orientation turns
on the sand near their entrances. Removal
of the spider from the cache and transport
of it to the nest occurred as in A. wheeleri.
Nests of A. taeniolatus comprised cylin-
drical burrows, 55-70 mm (X = 59, n = 5)
long which terminated in roughly ovoidal
cells, 44-57 mm (* = 48, n = 5) beneath
the sand surface. The paralyzed spider was
placed in the cell dorsal side upward and
head toward the entrance. Wasp eggs were
ca. 1.8 mm long and attached to the un-
dersides of the spiders’ abdomens perpen-
dicularly near the base. Prey taken from 4.
taeniolatus nests comprised the salticid Pel-
lenes (H.) borealis (2 9°, 2 imm. 22, | imm.).
One spider weighed (wet) 24 mg.
DISCUSSION
In A. wheeleri a black-red-black body pat-
tern coupled with red and black legs is ob-
viously disruptive yet highly cryptic against
a gravelly substrate and reminiscent of the
body coloration of Tachysphex pechumani
Krombein which ts also associated with a
gravelly habitat (see Kurczewski and Elliott
1978). Aside from theories expounding
cryptic or aposematic coloration, one ex-
planation for the relatively large amount of
erythrization in A. wheeleri is that the de-
veloping adult may be melanized less in
gravelly than in sandy soil. Water percolates
readily through the larger gravel particles
and is not intimately bound to the wasp’s
cell, thus approximating soil conditions in
some arid regions of the western U.S. and
possibly enhancing erythrization of the adult
exoskeleton. In sandy soil water adsorbs
tightly to the numerous small particles and
may keep the cell confines evenly humid,
thus resulting in more melanization of the
adult exoskeleton.
The dark pubescence exhibited by 4.
wheeleri might enhance the absorption of
VOLUME 90, NUMBER 3
ONTARIO
303
s
eS
oe
Fig. 10. Distribution of Aporinellus taeniolatus (@) and A. wheeleri (@) in the northeastern United States and
Canada.
radiant energy at higher altitudes and cooler
temperatures; however, males of A. wheeleri
have silvery pubescence which, according
to G. C. Eickwort (personal communica-
tion) is genetically unusual. Some females
of A. taeniolatus from the extreme southern
U.S. and Mexico also have dark pubes-
cence. Aporinellus completus Banks females
show geographic variation in the amount of
grey pubescence on the body with some
specimens appearing essentially entirely grey
and others mostly black. Females of 4. com-
pletus from Presque Isle State Park and Erie
County, PA were, however, identical in body
coloration.
A. taeniolatus has a broad geographic
range, occurring throughout southern Can-
ada and the United States and extending
into Mexico and Central America (Evans
1951, 1966, Krombein 1979). In the north-
eastern U.S. A. taeniolatus has a coastal dis-
tribution, whereas A. wheeleri exhibits an
upland distribution (Fig. 10). In the North-
east, 4. taeniolatus has been collected at al-
titudes ranging from near sea level (Well-
fleet, MA, Bayville, NY, Manumuskin, NJ)
to 580’ (Presque Isle St. Pk., PA, Long Point
Prov. Pk., ONT). Altitudes for A. wheeleri
range from +900’ (nr. Ithaca, NY) to 1985’
(Asheville, NC), except for specimens from
Bridgeport, NY, Princeton, NJ (det. K. W.
Cooper), and Stonybrook Res., Middlesex
Co., MA (det. J. Bequaert). In Erie County,
PA, where the two taxa are virtually sym-
patric, A. taeniolatus has been collected at
elevations of 575-580’ (Presque Isle St. Pk.)
and A. wheeleri, only 9-11 km away, at el-
evations of 1050-1200’ (Wintergreen Gorge
304
Table 1.
Characteristic
Flight season (PA)
Habitat (NE U.S.)
Soil type (NE U.S.)
Prey search
Sting
Cache
Burrow excavation
Excavation time
Orientation form
Prey transport type
Closure type
Burrow length
Cell depth
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
A. wheeleri
May-—October
Upland gravel pits, roadways, paths
Gravel, loam, shale
On plants, under stones
Underside of cephalothorax
Upright plant, stone
Mandibles and forelegs
4-43 min (% = 15.0)
Circles near entrance
Mandibles; ground/flight
Pile of pebbles on entrance (mandi-
bles)
18-28 mm (mixed sand and gravel)
14-23 mm (mixed sand and gravel);
Major ecological and behavioral characteristics of females of Aporinellus wheeleri and A. taeniolatus.
A. taeniolatus
June-September
Lowland soils near bodies of water
Sand
On plants
Underside of cephalothorax
Upright plant
Mostly forelegs
27 min
Circles near entrance
Mandibles; ground/flight
Sand fill (forelegs)
55-70 mm (sand)
44-57 mm (sand)
9-22 mm (gravel)
Prey placement
sum up
Oviposition site
Egg length
Prey type (PA)
Prey sex or stage
Prey weight (wet)
1.25-2.00 mm
6, 2, immature
12*-34 mg
* Thomisidae (Xysticus sp., immature).
Cemetery, Four Mile Creek nr. route I-90).
Although there is some overlap in altitude
when one compares their ranges in the
Northeast, 4. wheeleri occurs mostly at the
higher and A. faeniolatus, mostly at the low-
er elevations.
Habitat (soil type) is a better indicator of
distribution of the two taxa in the eastern
U.S. than altitude (Table 1). A. taeniolatus
is restricted to mostly sandy soils near bod-
ies of water (Fig. 9) whereas 4. wheeleri oc-
curs on heavier, often gravelly, soils away
from water (Figs. 1-3). Bequaert’s (1919)
type specimen (2) of A. wheeleri was col-
lected on a “stony woodroad” in Massa-
chusetts. Krombein (1958) reported 4.
wheeleri in association with gravelly soil in
West Virginia. Near Bridgeport, Madison
County, NY, an area characterized by sandy-
loamy soils, one female of 4. wheeleri was
collected on a pile of gravel near the base
of a utility pole connecting a power line
crossing a road. This gravel was not the typ-
Cephalothorax toward entrance, dor-
Underside of abdomen, near base
Mostly Salticidae (Pellenes borealis)
Cephalothorax toward entrance,
dorsum up
Underside of abdomen, near base
Ca. 1.8 mm
Salticidae (Pellenes borealis)
2, immature
24 mg
ical soil of the area but had been trucked in
from elsewhere and dumped to support the
pole. In the western U.S. and Mexico, A.
taeniolatus inhabits gravelly as well as sandy
soils (Evans 1951, 1966), thus filling the
niches of both species. One can envision the
evolution of 4. wheeleri from A. taeniolatus
in the eastern U.S. by progressively farther
movements inland away from the sandy
beaches near water (Great Lakes, Atlantic
Ocean) to higher altitudes and gravelly soil.
Such movements would obviously have
been influenced by the most recent period
of glaciation.
The inclusive flight periods for both taxa
in Erie County, PA (4. wheeleri, 24 May-2
October; A. taeniolatus, 28 June—22 Sep-
tember) suggest more than one generation
per year (Table 1). In New York we reared
males of 4. wheeleri in May from the pre-
vious September’s nests, extending the op-
portunity for a second generation in mid-
or late summer. Both A. taeniolatus and A.
VOLUME 90, NUMBER 3
wheeleri exhibit peak abundance in Erie
County, PA and Cayuga County, NY in late
summer (Aug.—Sept.), and occur only spar-
ingly throughout June and July. Evans
(1951) collected A. taeniolatus at East Hart-
ford, CT from 13 June to 19 Sept. but could
not ascertain the number of generations per
year.
In comparing the nesting behaviors of 4.
taeniolatus and A. wheeleri we found essen-
tially identical components to be the man-
ner of prey searching, method of paralysis,
cachement, transport, position of prey in
the cell, oviposition site, and kind of prey,
predominantly Pellenes borealis (Salticidae)
(Table 1).
Two major behavioral activities, burrow
excavation and closure, and the result there-
of, nest structure, are linked to the habitat
in which the wasp nests (Table 1). Thus, 4.
taeniolatus females dig and close nests in a
fashion typical of sand inhabiting pompi-
lids, i.e. they back out of the burrow while
throwing sand backward with the forelegs
which move alternately, and close the bur-
row while backing in, using the forelegs al-
ternately along with the abdominal apex for
tamping the sand into place. Females of A.
wheeleri which nest in patches of compacted
sand in gravelly soil dig their burrows sim-
ilarly. However, females of 4. wheeleri nest-
ing in broken shale or gravel use the man-
dibles considerably and forelegs less
frequently during burrow construction and
closure. As is typical of females in the family
Pompilidae, species inhabiting sand have
more elaborate foretarsal digging rakes, 1.e.
longer spines and more of them, than species
inhabiting heavier soils such as gravel, loam
and clay (Evans 1950). Thus, females of A.
taeniolatus collected on sand in the North-
east have a more extensive foretarsal rake
with longer spines than females of 4. whee-
leri; however, females of A. taeniolatus col-
lected in the extreme southern U.S. and
Mexico on gravel have a short foretarsal
rake similar to that of A. wheeleri.
In both taxa the architecture of the bur-
305
row reflects the soil in which the nest is
excavated (Table 1). Nests of A. taeniolatus
in loose sand comprise elongate (55-70 mm)
cylindrical burrows, those of 4. wheeleri in
compacted sand and gravel, short (20-28
mm) cylindrical burrows, and those of A.
wheeleri in gravel, short cylindrical burrows
or shallow conical depressions (Fig. 5).
Closed burrows of 4. taeniolatus are indis-
tinguishable from the surrounding sand. For
A. wheeleri the burrow closure is clearly vis-
ible as a characteristic circular mound of
small pebbles covering the filled nest en-
trance (Fig. 4).
In conclusion, although Evans (1951)
considered 4. wheeleri to be nothing more
than a subspecies of 4. taeniatus (= taenio-
latus) based upon its localized geographic
distribution in the Alleghenian Fauna and
distinctive body coloration, the available
morphological, distributional, ecological
and behavioral data suggest that the two
taxa have separate gene pools in the eastern
United States (Kurczewski and Kurczewski
1987). Aporinellus wheeleri and A. taeniola-
tus are distinguished by a combination of
geographic, altitudinal, soil, behavioral and
morphological (2) criteria; the two taxa are
similar in their inclusive flight periods, 4
morphology (including genitalia), prey pref-
erences and many behavioral features. In
the western U.S. and Mexico, A. taeniolatus
inhabits heavy, somewhat gravelly, as well
as sandy soil (Evans 1951, 1966), thus oc-
curring in areas similar to those inhabited
by 4. wheeleri in the Northeast. The dis-
tinctive ecology and morphology exhibited
by these two taxa in the northeastern U.S.
may reflect character divergence now com-
pounded by human disturbance.
ACKNOWLEDGMENTS
We thank R. A. Norton, SUNY-CESF,
for identifying the species of Salticidae used
as prey and reviewing the manuscript, and
M. S. Wasbauer, Calif. Dept. of Food and
Agric., for reviewing the manuscript and ad-
dressing the biosystematics problem of the
306
Taeniolatus group. We appreciate the input
of G. C. Eickwort, Cornell University, on
the specific statuses of the two species of
Aporinellus. We also thank R. J. Gagné and
D. R. Smith, both of BBH, ARS, USDA,
for determining Sarcophagidae and For-
micidae, respectively. E. Tucker, Jr., Beh-
rend College of The Pennsylvania State
University, provided information on the el-
evations in Erie County, PA. A. F. Newton,
Jr., Museum of Comparative Zoology, Har-
vard University, L. L. Pechuman, Cornell
University, Marjorie Favreau, American
Museum of Natural History, and A. S.
Menke, BBII, ARS, USDA, furnished dis-
tributional data for A. wheeleri and A. tae-
niolatus.
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. 1959. Prey records for some midwestern and
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PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 307-309
ESTABLISHMENT OF CYBOCEPHALUS SP. (COLEOPTERA: NITIDULIDAE)
FROM KOREA ON UNASPIS EUONYMI (HOMOPTERA: DIASPIDIDAE)
IN THE EASTERN UNITED STATES
J. J. DREA AND R. W. CARLSON
(JJD) Beneficial Insects Laboratory, PSI, Agricultural Research Service, USDA, BARC-
East, Beltsville, Maryland 20705; (RWC) PSI, Systematic Entomology Laboratory, Ag-
ricultural Research Service, USDA, BARC-West, Beltsville, Maryland 20705 (formerly
at the Asian Parasite Laboratory, % American Embassy, Seoul, Republic of Korea).
Abstract.—In 1983, a species of Cybocephalus, tentatively identified as C. prob. nip-
ponicus Endrody-Younga, was imported from the Republic of Korea and released on
euonymus trees and shrubs infested with the euonymus scale, Unaspis euonymi (Com-
stock). The predaceous nitidulid is well established at three release sites in the metropolitan
Washington, D.C. area and has reduced the scale populations on the host plants. Distri-
bution is in progress to establish this beneficial beetle at other locations in the eastern
United States.
Key Words:
In 1980 the Agricultural Research Ser-
vice, United States Department of Agricul-
ture, initiated a Small Farm Research proj-
ect to study the biological control of several
species of armored scale pests. Among the
targeted insects was the euonymous scale,
Unaspis euonymi (Comstock), an exotic pest
of Asian origin that is devastating to many
species of ornamental plants of the genus
Euonymus (Gill et al. 1982).
Because U. euonymiis common in Korea,
the USDA Asian Parasite Laboratory, Seoul,
Republic of Korea (ROK), was assigned the
task of obtaining natural enemies of the scale
as a part of its research effort. Among the
many biotic agents found was a species of
Cybocephalus that preyed on all stages of
the euonymus scale. We have tentatively
identified this nitidulid as C. prob. nippon-
icus Endrody-Younga through comparison
with paratypes of C. nipponicus in the col-
lection of the United States National Mu-
seum of Natural History, Washington, D.C.
Biologicai control; armored scales; predator; Diaspididae; Nitidulidae.
Species of Cybocephalus are important
predators of diaspine scales. They may rank
second to the Coccinellidae as predators of
armored scales but nitidulids and their im-
pact on scale populations have received rel-
atively little study. However, several species
have been introduced into various parts of
the world for scale control (Drea 1988).
Limited introductions of Cybocephalus spp.
have been made into North America but
with little success. A species of Japanese
origin, referred to as C. probably gibbulus
Erichson, was introduced and established in
California in 1932-1933 for the control of
the California red scale, Aonidiella aurantii
(Maskell), (Rosen and DeBach 1978). Al-
though the predator expanded its host range
to include two other species of scale, it has
not proven to be effective (Bumgardner
1945). In 1956-1957, a Cybocephalus sp.
from India was introduced into California
but did not become established (Rosen and
DeBach 1978).
308
Our introductions of C. prob. nipponicus
began in 1984. Three sites were selected in
the metropolitan Washington area for re-
leases of the beetle. On April 5, 1984, a total
of 29 adults (24 2, 5 4), from Namhansan-
seong, Kyeonggi Province and Seoul, ROK,
were released at the Chadwick Overlook at
the National Arboretum on Euonymus for-
tunei (Turcz.) Hand.-Mazz., a shrub heavily
infested with U. euonymi. A colony was es-
tablished and living individuals were reg-
ularly collected at this site through 1986.
A second site in the National Arboretum
was a grove of euonymus trees near the Ad-
ministrative Center. In August, 1984, 15
adults (11 2, 4 4) from Sacheon, Kyeong-
sangnam Province, ROK, were released on
E. europaeus L. heavily infested with the
euonymus scale. Within several weeks the
colony expanded and the beetles spread to
neighboring euonymus trees, E. hamilton-
ianus var. nikoensis (Nakai) Blakelock and
E. kiautshovicus Loes., also heavily infested
with the scale. The scale population on the
release tree had been reduced to an insig-
nificant level by late summer, 1986, and the
tree has shown increased growth.
The third release site was an unidentified
euonymus tree at the USDA Beltsville Ag-
ricultural Research Center in Maryland. In
May, 1984, a total of 32 adults (25 9, 7 4)
were released on the tree and quickly in-
creased in number. Individuals were re-
covered from the tree in 1985 and 1986.
Because a laboratory colony of the beetle
was not established, it was necessary to uti-
lize field collected insects for the releases.
However, before any beetles were released,
samples from the Korean material were dis-
sected and examined for parasites, but none
were found. Furthermore, Williams et al.
(1984) report that there are no known par-
asites of adult Cybocephalus, although lar-
vae of the genus are recorded as hosts of
Zeteticontus (Encyrtidae), Aphanogmus
(Ceraphronidae), and Zatropis (Pteromali-
dae).
Once a field colony was established, no
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
additional releases were made with Korean
specimens. Consequently, a total of 76 adults
(60 2, 16 4) were liberated in the three sites
of the metropolitan Washington area. Using
the trees in the National Arboretum as a
natural insectary, more than 4000 Cybo-
cephalus adults were collected and shipped
during 1985 and 1986 for release by co-
operators in Pennsylvania, New Jersey,
Delaware, North Carolina, and Ohio.
Blumberg and Swirski (1982) present a
comprehensive study of the biologies of C.
micans Reitter and C. nigriceps nigriceps
(Sahlberg) that prey on diaspine scales in
Israel. Our observations suggest that C. prob.
nipponicus has a life cycle similar to these
two nitidulid species.
At the National Arboretum the beetles
had 3, and possibly 4, generations per year.
The greyish-white eggs, 0.5 x 0.2 mm, were
laid in or under empty scale coverings and
in other protected areas. According to
Blumberg and Swirski (1982), eggs of the
species they studied hatched in about one
week, and the larvae completed 3 instars in
one or more weeks. Pupation reportedly oc-
curs on the plant, in debris near or on the
plant, or in the soil (Clausen 1956, Kartman
1946). Pupae of C. prob. nipponicus were
found on leaves of the plant.
Adults of C. prob. nipponicus are small,
about 1.0 mm in length, hemispherical in
appearance. The female is entirely black, the
male has a yellow head and pronotum. The
adults can live 1 month or more depending
upon the season. Overwintering beetles at
the Arboretum survived at least 5 months,
from November to April. During the winter
beetles were active on the trees when the
temperature was about 10°C.
There appears to be a degree of host spec-
ificity in species of Cybocephalus. Blumberg
and Swirski (1974) noted a difference in the
effect of the species of scale on the devel-
opment of the species of Cybocephalus. They
observed that C. micans preyed mainly on
the nymphal stages of A. aurantii and other
diaspine scales and that younger instars of
VOLUME 90, NUMBER 3
the predator were not able to feed on mature
female scales. Our laboratory and field ob-
servations with C. prob. nipponicus indi-
cated that the small male scales are the chief
prey of the adults. More beetles were found
on leaves infested with the males of U. eu-
onymi than on branches where the female
scales were more abundant. Microscopic ex-
amination indicated that more male than
female scales were damaged by the beetles.
The sex ratio, was 1:1 determined from
756 individuals of 7 field samples, collected
from May through September, 1986.
At present, C. prob. nipponicus is well
established at 3 sites in the metropolitan
Washington area. The beetle has had an im-
pact on the populations of euonymus scales
at these sites. The predator has increased to
a level that has permitted the collection and
shipment of the beetle to other states in the
eastern United States. However, the pop-
ulation of the host at the original release
sites has been reduced to a very low level,
because of the predation by C. prob. nip-
ponicus and by a coccinellid, Chilocorus ku-
wanae (Silvestri), also introduced from Ko-
rea and established at the same study sites
(Drea and Carlson 1987).
ACKNOWLEDGMENTS
We wish to thank Ho-Yeon Han and Jang-
Hoon Lee, personnel of the Asian Parasite
Laboratory, who assisted in the original col-
lections in the ROK; S. G. March, U.S. Na-
tional Arboretum, Washington, D.C. for as-
sistance in locating scale populations; and
L. R. Ertle and R. M. Hendrickson, Jr., Ben-
309
eficial Insects Research Laboratory, Agri-
cultural Research Service, USDA, Newark,
Delaware for assistance in quarantine clear-
ance and release of the beetle in northern
areas.
LITERATURE CITED
Blumberg, D. and E. Swirski. 1974. The development
and reproduction of cybocephalid beetles on var-
ious foods. Entomophaga 19: 437-443.
. 1982. Comparative biological studies on two
species of predator beetles of the genus Cyboceph-
alus (Col.: Cybocephalidae). Entomophaga 27: 67-
76.
Bumgardner, R. J. 1945. Cybocephalus established
in California. J. Econ. Entomol. 38: 128.
Clausen, C. P. 1956. Biological control of insect pests
in the continental United States. U.S. Dep. Agric.
Tech. Bull. 1139. 151 pp.
Drea, J. J. 1988. Other Coleoptera. /n Rosen, D., ed.,
Armored Scale Insects, Their Biology, Natural
Enemies, and Control: Natural Enemies. Elsevier
Science Publications. (In press.)
Drea, J. J. and R. W. Carlson. 1987. The establish-
ment of Chilocorus kuwanae (Coleoptera: Cocci-
nellidae) in eastern United States. Proc. Entomol.
Soc. Wash. 84: 821-824.
Gill, S. A., D. R. Miller, and J. A. Davidson. 1982.
Bionomics and taxonomy of the Euonymus scale,
Unaspis euonymi (Comstock), and detailed bio-
logical information on the scale in Maryland (Ho-
moptera: Diaspididae). Univ. Md. Agric. Exp. Stn.
Misc. Pub. 969. 36 pp.
Kartman, L. 1946. A new host for Cyhocephalus sp.,
a predator of diaspine Coccidae. J. Econ. Entomol.
39: 814.
Rosen, D. and P. DeBach. 1978. Diaspididae, pp.
78-128. Jn Clausen, C. P., ed., Introduced Para-
sites and Predators of Arthropod Pests and Weeds:
A World Review. U.S. Dep. Agric., Agric. Handb.
No. 480. 551 pp.
Williams, R. N., M. J. Weiss, M. Kehat, and D. Blum-
berg. 1984. The hymenopterous parasites of Ni-
tidulidae. Phytoparasitica 12: 53-64.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 310-322
A REVIEW OF NEARCTIC MERISMUS WALKER AND TOXEUMA WALKER
(HYMENOPTERA: CHALCIDOIDEA: PTEROMALIDAE)
S. L. HEYDON AND E. E. GRISSELL
(SLH) Section of Faunistic Surveys and Insect Identification, Illinois Natural History
Survey, Champaign, Illinois 61820; (EEG) Systematic Entomology Laboratory, Agricul-
tural Research Service, USDA, % U.S. National Museum, NHB 168, Washington, D.C.
20560.
Abstract.—The genus Toxeuma Walker, herein reported for the first time from the
Nearctic region, is represented by four new species: aciculare Heydon, aquilonium Heydon,
gerra Heydon, and inopinum Heydon. The specimen from which Toxeuma gerra is de-
scribed was reported as Merismus rufipes Walker in the Nearctic, but this is a misiden-
tification and the record should be eliminated. However, the Palearctic species Merismus
megapterus Walker and M. lasthenes (Walker) are reported for the first time from the
northern Nearctic region. A key to females of Nearctic Toxeuma is given.
Key Words:
mineae
The Palearctic species Merismus rufipes
Walker was first reported in the Nearctic
region by Burks (1979). The specimen upon
which this record was based is in the col-
lection of the U.S. National Museum of
Natural History and represents an unde-
scribed species of Toxeuma, a genus not
previously known from North America. In
this paper we report Toxeuma from the
Nearctic for the first time and describe four
new species. Additionally, we report Mer-
ismus megapterus Walker and M. lasthenes
(Walker) for the first time from North
America. Both species were previously
thought to be restricted to the Palearctic,
but the former is widespread in the Nearctic
and the latter has been found in Alaska.
This paper is part of a series by the senior
author reviewing the Nearctic fauna of the
miscogasterine Pteromalidae (Heydon
1988a, b, Heydon and LaBerge 1988). Tox-
euma and Merismus are phenetically sim-
ilar genera and belong with Cryptoprymna
Chalcidoidea, Pteromalidae, Merismus, Toxeuma, Agromyzidae, Gra-
Foerster in a group of miscogasterine genera
characterized by the following synapomor-
phies: 1. The genal concavities extending
around one-third of the malar distance. 2.
A tendency for the female club to have a
large patch of micropilosity. 3. A propo-
deum which is usually rather elongate (ex-
cept in Merimus megapterus), arched front
to back, and with plicae that tend to curve
regularly till they converge posteriorly. The
plicae in most other closely related genera
are initially parallel, and then there is a sharp
angle at the point where they begin to con-
verge. Cryptoprymna is distinct from Mer-
ismus and Toxeuma by its loss of metallic
coloration, its smooth frenum, and elongate
hypopygium which extends to the tip of the
gaster. Merismus and Toxeuma retain me-
tallic coloration, a reticulate frenum, and a
hypopygium extending less than three-
fourths the gastral length. Nearctic Cryp-
toprymna is treated by Heydon (1988a).
The following abbreviations are used for
VOLUME 90, NUMBER 3
institutions in the text: BMNH = British
Museum (Natural History), London, En-
gland; USNM = U.S. National Museum of
Natural History, Washington, D.C., U.S.A.;
CNC = Canadian National Collection, Ot-
tawa, Canada; SEC = Snow Entomological
Collection, University of Kansas, Law-
rence, Kansas, U.S.A.; INHS = Illinois Nat-
ural History Survey, Champaign, Illinois,
U.S.A. Abbreviations used in the descrip-
tions are: LOD = lateral ocellar diameter;
OOL = ocelocular line; POL = posterior
ocellar line; LOL = lateral ocellar line; F =
funicular segment; T = tergal segment (ap-
parent gastral terga, excluding petiole). Body
sizes are given as thoracic lengths measured
from anterior of pronotal neck to posterior
of propodeal nucha; the long petiole of the
gaster allows so much flexion that total body
length often cannot be measured accurately.
Measurements given are units from a Wild
120 unit reticle at 50 x and can be converted
to millimeters by multiplying by 0.02.
Toxeuma Walker
Toxeuma Walker, 1833: 371, 378. Type
species: Toxeuma fusicornis Walker. Des-
ig. by Westwood, 1839: 68.
This genus is known from the Palearctic,
Neotropical, and Australian regions and was
revised in the Palearctic by Graham (1959,
1969) and Dzhanokmen (1978). The dis-
tribution of world species by regions is as
follows (the generic placement of these
names was not confirmed and we cite them
as currently recognized without attempt at
correction): Palearctic: acilius (Walker), dis-
cretum Graham, fuscicorne Walker, mu-
cronatum Graham, paludum Graham, sub-
truncatum Graham, Neotropical: aphareus
(Walker), faceta Girault, orobia (Walker);
Oriental: affinis Ashmead, ferrugineipes
Ashmead, hawaiiensis Ashmead, nigrocy-
anea Ashmead, nubilipennis Ashmead, tar-
sata Ashmead; Australian: pax Girault. To
this list we add the four new Nearctic species
described below.
The only biological record is for T. fus-
311
cicorne, which was reared from seeds of A ve-
na (Graham 1969); the actual insect host
within the seeds is unknown.
The genus 7oxeuma may be recognized
as follows: body metallic green or blue; clyp-
eus with anterior margin straight, lacking
denticles (Figs. 10, 12); gena with distinct
hollow above mandible (Fig. 10) except i
T. gerra (Fig. 12); antennal formula 1:1:2
6:3, female club with (Fig. 11) or without
large ventral patch of micropilosity and api-
cal spine sometimes present (Figs. 4, 11);
pronotum with collar with sharp anterior
transverse carina; notauli complete, furrow-
like; prepectus reticulate, without carinae;
scutellum with 2 or 3 pairs of lateral setae,
frenum set off by punctate sulcus; propo-
deum with median carina and plicae, usu-
ally also with weak rugae; gastral petiole
reticulate, transverse to elongate, with basal
flange laterally and ventrally, lateral longi-
tudinal carinae present (Figs. 6, 7) except in
T. mucronatum, gaster fusiform or lanceo-
late; Tl often nearly covering dorsum of
gaster, hind margin straight mesally and lat-
erally. Characters to differentiate Merismus
from Toxeuma are listed under the former
genus.
Key TO FEMALE NEARCTIC TOXEUMA
The following key will separate females
of the Nearctic species of Toxeuma. Males
of 7. gerraare unknown. Because males have
a simple apical club segment (no spine or
micropilosity) and vary in color and wing
setation patterns, it is not possible to pro-
vide reliable characters to identify unasso-
ciated males at this time.
1. Antennal club with terminal spine (Figs. 4, 9,
LUI 2 rcs x site eenvish amen tea eros fr oaiateten steers 2
— Antennal club rounded apically, lacking ter-
minal spine (Fig. 1) :
ne _ aquilonium Heydon,: new species
Club with needle- like spine at apex (Figs. 4, 9);
basal vein with 1-7 setae (rarely bare) (Figs. 5,
8); gena with distinct hollow extending “4 to 1
malar length (Fig. 10) . 3
— Club with conical spine at apex (Fig. 11); basal
vein bare; genal hollow obscure, extending
malar length (Fig. 12) . gerra Heydon, new species
to
312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-12. 1-2, Toxeuma aquilonium n. sp., female. 1, Club. 2, Forewing. 3-6, Toxeuma inopinum n. sp.,
female. 3, Antenna. 4, Club. 5, Forewing. 6, Petiole. 7-10, Toxeuma aciculare n. sp., female. 7, Petiole. 8,
Forewing, 9, Antenna. 10, Clypeus and lower face. 11-12, Toxeuma gerra n. sp., female. 11, Club. 12, Clypeus
and lower face. Scale bar = 0.1 mm.
3. Funicular segments I-3 elongate (Fig. 3); gas- Toxeuma aciculare Heydon,
tral petiole as long as wide (Fig. 6) ......... New SPECIES
shore crave rewire inopinum Heydon, new species Figs. 7-10
— Funicular segments 1-3 quadrate (Fig. 9); gas- :
tral petiole 1.5-2.0 as long as wide (Fig. 7) Holotype female.—Thoracic length 0.90
Conoco aciculare Heydon, new species mm. Body and coxae dark green except: oc-
VOLUME 90, NUMBER 3
ciput, neck of pronotum, and gastral terga
after Tl darker, almost black; mandibles
dark reddish brown with teeth reddish; legs
beyond coxae yellow-brown, mid- and hind
tarsi cream, pretarsi dark; antenna brown
with basal 73 of scape yellow-brown; wing
veins translucent brown. Head subreticulate
with clypeus microalutaceous; thorax retic-
ulate except pronotum with smooth band
along hind margin (length equal to median
length of collar), and frenum subreticulate;
propodeum with region between plicae re-
ticulate with some rugae, nucha subreticu-
late; petiole rugulose between irregular lon-
gitudinal carinae; gastral terga smooth. Head
ovate in anterior view, |.2 x as wide as high
(29.5:25), 2.0x as wide as long (29.5:15);
occiput moderately concave; clypeus with
anterior margin produced, anterior tentorial
pits obscure; anterior margins of face lat-
erad of clypeus produced (Fig. 10); gena with
hollow extending almost 3 malar length (1.5:
5.5); toruli | diameter above lower eye mar-
gin (2:2); intermalar distance 3.1 x malar
length (17:5.5); eye height 2.7 x malar length
(15:5.5); ratio of LOD:OOL:POL:LOL as 3:
4:7:4: antenna with pedicel plus flagellum
1.0x head width (30:29.5), ratio of lengths
of scape: pedicel: annelli 1 + 2: funicular
segments 1-6: club as 13:4:1:2.5:2.5:2.5:3:
2.5:2.5:8.5, funicular segments | and 6 both
0.91 x as long as wide (2.5:2.75) (Fig. 9),
apical segment of club with subterminal
needle-like spine and patch of micropilosity
ventrally, ratio of ventral lengths of basal :
second club segments 1.25 (2.5:2). Thorax
with propodeum shorter than scutellum (11:
14), nucha set-off by low carina anteriorly.
Forewing with ratio of submarginal : mar-
ginal : postmarginal : stigmal veins as 38:15:
12:7; basal vein with 2 setae on right wing
and | on left; basal cell bare; costal cell with
1 complete ventral row of setae (Fig. 8).
Petiole 1.8 x as long as wide (9:5) (Fig. 7),
median and sublateral carinae weakly de-
veloped. Gaster ovate, 1.6 x as long as wide
(32:20), basal tergum 0.75 x median length
(24:32), a row of 3 setae distal to each pos-
313
terolateral corner of basal fovea; T3-6 re-
tracted beneath T2.
Allotype male.—Thoracic length 0.90
mm. Similar to female except generally light
green; thorax with strong golden reflections;
mid- and hind femora darkly pigmented
over most of length, mid-tibia with dark
band on basal third; antenna with pedicel
plus flagellum 1.2 x head width (35:28), ra-
tio scape : pedicel : annelli | + 2: funicular
segments 1-6: clubas 1 1:4:1:3.5:3.5:3.5:3.5:
3:3:10, funicular segments decreasing from
longer than wide (FI 1.4 as long as wide,
3.5:2.5) to as wide as long (F6 3:3); ratio of
gaster length: width as 26:16, T1 0.80
median length (20:25).
Type material.—Holotype 2, Illinois,
Champaign Co., Champaign (from railroad
siding at the end of Gerty Street on the South
Farms of the University of Illinois), 26-V-
1985, S. L. Heydon. Allotype 3, 2 2 and 17
é paratypes with same data. Additional 24
paratypes as follows. —CANADA: New
Brunswick: Fredricton (Acadia Field Sta-
tion), 13-VII-1970, 1 2: Kouchibouguac Na-
tional Park, 5-VIII-1977, 2 2; 16-VII-1977,
2 6. Ontario: Innisville, 16-VII-1963, 2 9,
18-VII-1962, 1 2. Quebec: Cap Rouge, 4-VII-
1956, 1 2 7-VII-1953, 1 ¢. UNITED
STATES: //linois: same data as holotype ex-
cept 19-V-1985, 2 2, 1 4; 8-VI-1985, 42, 1
6; 19-VI-1985, 3 2; 21-VI-1985, 1 4; 22-VI-
1985, 5 9; 24-VI-1985, 1 9; 21-VIII-1981,
1 2. Michigan, Iron Co., 27-VIHI-1952, 1 °.
Holotype female, allotype male, and para-
types in USNM. Additional paratypes in
BMNH, CNC, and INHS.
Etymology.—The species name is de-
rived from the Latin acicularis—like a
needle—referring to the needle-like process
on the tip of the female antenna.
Variation.—In females, the body varies
from green to dark green to blue-green. The
topotypic females nearly all have dark green
bodies, only a couple being more blue-green.
However, about half the females from Can-
ada are blue-green. The scape varies from
completely pale to brownish with a trace of
314
metallic coloration over the apical half.
Femora are always pale. The thoracic length
varies between 0.62 and 0.90 mm. The bas-
al funicular segments are rarely slightly
longer than wide (2.5:2) but F3 is always
quadrate. The petiole varies from 1.4 to
2.0x as long as wide (X = 1.56, n = 10).
One specimen does not have gastral terga
3-6 retracted beneath T1 and T2. In males,
which were all collected from the type-lo-
cality, the dark bands on the femora and
tibiae may be strongly or only faintly de-
veloped, the scape may be entirely dark or
pale, the gold reflections of the thorax may
not be visible and there may be some setae
in the basal cell. The thoracic length varies
between 0.70 and 0.86 mm. In most spec-
imens, all funicular segments are longer than
wide. In both sexes, the number of setae on
the basal vein varies from 0-5 but 84% of
female wings examined had between | and
3 setae (n = 20).
Discussion. — Toxeuma aciculare closely
resembles inopinum in having the nucha set
off anteriorly by a fine carina and having a
small area of micropilosity ventrally and
needle-like subterminal spine on the apical
club segment of the female antenna. These
species are readily separated by characters
given in the key and also because aciculare
almost never has setae in the basal cell while
inopinum almost always does. The eye
height is generally more than 2.5 x the ma-
lar length in aciculare while the eye height
is less than this in inopinum. The scape of
T. aquilonium is always strongly metallic,
and though the scape of T. aciculare is rarely
brown for half its length, even then, it has
only a trace of metallic coloration. The Pale-
arctic species, 7. mucronatum Graham has
a terminal spine on the antennal club, but
differs from all such Nearctic Toxeuma
species in having no rugae on the propo-
deum, and the petiole uniformly cylindrical
and reticulate. In Nearctic Toxeuwma species,
the petiole always has a pair of distinct lat-
eral carinae and it usually has rugae dor-
sally.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Toxeuma aquilonium Heydon,
New SPECIES
Figs. 1-2
Holotype female.—Thoracic length 1.0
mm. Body and coxae blue-green except:
frons and discs of mesoscutum and scutel-
lum which are green, occiput and neck of
pronotum dark; mandible yellowish brown;
legs yellow with fore and hind tibiae with
brown bands basally, pretarsi brown; an-
tenna brown with scape metallic blue-green;
wing veins translucent brown; setae of body
reddish brown. Sculpture similar to 7. aci-
culare except face and frenum alutaceous.
Head 1.2 as wide as high (28:22.5), 2.0 x
as wide as long (28:14); clypeus with ante-
rior margin truncate, tentorial pits obscure;
anterior margin of face laterad of clypeus
not produced; genal hollow extending ma-
lar length (1:5); torulia little over 1 diameter
above lower eye margin (2:1.5); intermalar
distance 3.2 x malar length (17:5); eye height
2.7x malar length (14.5:5); ratio of LOD:
OOL:POL:LOL as 2.5:3.75:6.25:3; antenna
with pedicel plus flagellum 1.1 x head width
(30.5:28), ratio of lengths of scape : pedicel :
annelli | + 2: funicular segments 1-6: club
asl 325:4:12295:3:3:3:°3:32 5:8 wieleaas
long as wide (2.5:2.25), F6 0.71 as long
as wide (2.5:3.5), apical segment of club
without spine but with patch of micropi-
losity on ventral surface, ratio of ventral
lengths of basal: second club segments 1.7
(5:3) (Fig. 1). Thorax having propodeum al-
most as long as scutellum (11:12), costulae
distinct, nucha acarinate anteriorly. Fore-
wing with ratio of submarginal : marginal :
postmarginal : stigmal veins as 27:16:15:8,
basal vein with 3 setae on left wing and 5
on right, basal cell with 2 setae, costal cell
with | complete row of setae and partial
second row apically (Fig. 2). Petiole 1.5 x
as long as wide (6:4), with complete median
carina, a pair of lateral carinae basally which
converge with the median carina basally,
and a pair of short sublateral carinae api-
cally. Gaster fusiliform, 1.7 x as long as wide
VOLUME 90, NUMBER 3
(32.5:19), T1 extending 0.6 x median length,
2 setae present distal to each posterolateral
corner of basal fovea, all terga visible, T2-
6 with | transverse row of setae.
Allotype male.—Thoracic length 0.94
mm. Body blue. Sculpture similar to female.
Antenna with pedicel plus flagellum 1.2 x
head width (35:28), ratio of scape : pedicel :
annelli | + 2: funicular segments 1-6: club
as 11:4:1:4:3.5:3.5:3.75:3.5:3.25:9, all fu-
nicular segments elongate (Fl 1.6 as long
as wide (4:2.5), F6 1.3 as long as wide
(3.25:2.5); basal vein with 2 complete but
irregular rows of setae.
Type material.—Holotype female, Alas-
ka, Matanuska Susitna Borough, Matanus-
ka, 12-15-VII-1945, J. C. Chamberlin (Soil
emergence cage 118-119, series II, Lot No.
45-19986). Allotype and | paratype male
same locality, 20-VI-1945, A. Linn (with
thrips swept from grass; Lot No. 45-19986).
Additional 23 paratypes as follows. —CAN-
ADA. Alberta: Bilby, 1-VI-1924, 1 2. British
Columbia: Hixon, 11-29-VII-1965, 7 9°:
Kaslo, 1943, 1 9. Ontario: Ottawa, 27-VI-
1982, 1 9, 1 6; One Sided Lake, 16-VII-
1960, 1 2. Quebec: Laniel, 14-VI-1942, 1 ¢;
8-VII-1944, | 2. Saskatchewan: White Fox,
7-VU-1944, 1 9. UNITED STATES Alaska:
Kenai-Cook Inlet, Kenai Peninsula, 1 mi.
S Jct. Hwy. #4 and Homer Road, 30-VI-
1957, F. W. Preston, 5 4. Colorado: Doo-
little Ranch (Near Mt. Evans), 23-VIII-1961,
1°. New Mexico: Lincoln National Forest,
26-30-VII-1977, 3 2, 2 8. Holotype, allo-
type, and paratypes in USNM. Additional
paratypes in CNC, INHS, and SEC.
Etymology.—The species name is de-
rived from the Latin aquilonium, meaning
northerly.
Variation.—The females from Kaslo,
British Columbia; Colorado; and Saskatch-
ewan lack the dark bands on the legs, but
still have the scape nearly totally metallic.
The number of setae in the basal cell varies
between 0 and 6, with 17 of 23 wings having
at least | and 10 having at least 2. The pet-
iole varies from as long as wide to 1.4 times
315
as long as wide (x = 1.24, n = 10). The
amount of telescoping of the gastral terga
depends on the method of drying. Most ter-
ga are retracted beneath T1 in air-dried
specimens. The allotype male has a quad-
rate petiole while the other male paratypes
have elongate petioles. The other male from
Matanuska has broken antennae and there-
fore was not selected as the allotype.
Discussion.—Females of aquilonium are
distinct from those of the other three Nearc-
tic species of Toxeuma because they lack
the apical spine on the clava. Males and
females resemble inopinum in the pattern
of wing setae and closely resemble those in-
opinum from western Canada in color pat-
tern. However, in aquilonium the petiole
averages longer than wide and the basal fla-
gellar segments are usually transverse or
quadrate, whereas the petiole in inopinum
is as long as wide and the funicular segments
are always elongate. The scape is nearly
wholly metallic throughout the range of
aquilonium, while outside western Canada
and montane western U.S.A., the scape of
inopinum 1s light over at least the basal third.
Toxeuma inopinum also has setae in the
basal cell, but in this species the setae are
almost invariably inserted adjacent to those
on the basal vein while in aquilonium the
setae are often inserted at some distance
from the basal vein. Toxeuma aquilonium
would key out to subtruncatum in Graham
(1969) but differs from this species in having
the sides of the pronotal collar distinctly
convergent anteriorly in dorsal view.
Toxeuma gerra Heydon,
NEw SPECIES
Figs. 11-12
Merismus rufipes, Burks, 1979: 789; nec
Walker, 1833: 378. (Misidentification)
Holotype female.—Thoracic length 0.79
mm. Body and coxae blue-green except: oc-
ciput and neck of pronotum dark blue,
mesoscutum and scutellum green; mandible
brown; legs yellow, pretarsi brown; wing
316
veins translucent brown. Sculpture like T.
aciculare except face and frenum alutaceous
and gastral tergum 7 microalutaceous. Head
1.1 x as wide as high (21:19), 1.8 as wide
as long (19:12); clypeus with anterior mar-
gin produced medially, anterior tentorial pits
distinct (Fig. 12); genal hollow narrow, ex-
tending 0.14 malar length (1:7) (Fig. 12);
face with subantennal tubercle ventrally
curving abruptly to meet dorsal edge of
clypeus; toruli | diameter above lower eye
margin (2:2); intermalar distance 3.0 x ma-
lar length (15:5); eye 2.2 malar length
(11.5); ratio of LOD:OOL:POL:LOL as 1.5:
4:6:3; antenna with pedicel plus flagellum
1.1 head width (24:21), ratio of lengths of
scape : pedicel : annelli | + 2 : funicular seg-
ments 1-6:club as 10:3.5:1:2:2:2:2:2:2:8,
Fl 1.1 as long as wide (2:1.75) and F6
0.8 x as long as wide (2:2.5), ratio of flagellar
length to head width 1.1 (24:21); apical seg-
ment of club with conical terminal spine
and ventral patch of micropilosity extend-
ing more than halfway to club base, sutures
between the club segments strongly oblique
(ratio of ventral lengths of second : basal club
segments 0.5 (1:2) although dorsal lengths
subequal (4:4) (Fig. 11)). Thorax with pro-
podeum shorter than scutellum (7:10), with
plicae and medina carina complete and dis-
tinct and with irregular rugae posteriorly,
basal fovae smooth and bordered on all sides
by carinae, spiracular sulci shallow, nucha
collar-like, transversely rugulose with traces
of carina anteriorly. Forewing with ratio of
submarginal : marginal : postmarginal :
stigmal veins as 24:13:8:5, basal cell and
vein bare, costal cell with single complete
row of setae ventrally. Petiole 1.3 x as long
as wide (6:4.5), median and sublateral ca-
rinae present anteriorly. Gaster ovate; 2.1 x
as long as wide (29:14); T1 extending 0.75 x
median length, 2 setae present distal to each
posterolateral corner of basal fovea; only
posterior margin of T2 and all of T7 visible.
Male. — Unknown.
Type material.—Holotype female, Vir-
ginia, Montgomery Co., 21-VIII-1973, J. M.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Beisler (on sedge). One additional paratype
female from Nova Scotia, Bridgetown, 17-
VIII-1912. Holotype in USNM, paratype in
CNC.
Etymology.—The name is from the Latin
gerra—trifle—referring to the smallness and
rareness of this species.
Discussion. — Females of gerra have a ter-
minal spine and patch of micropilosity on
the club like those of aciculare and inopin-
um but the terminal spine is conical not
lanceolate and the patch of micropilosity
extends more than halfway to the base of
the club (Fig. 11). In females of aciculare
and inopinum the patch extends less than
halfway to base of the club (Figs. 4, 9).
Unique characteristics of females, and pos-
sibly males, of this species are the lack of
setae on the basal cell and basal vein, the
distinct anterior tentorial pits, and narrow
genal hollows which extend only ¥ genal
length.
Toxeuma inopinum Heydon,
New SPECIES
Figs. 3-6
Holotype female.—Thoracic length 0.88
mm. Head with face blue-green, frons green,
vertex from lateral ocellus and occiput dark
blue; thorax with pronotum black except
smooth posterior hind margin dark yellow-
green, remainder, including coxae, blue-
green, propodeum with yellowish reflec-
tions; gastral petiole dark blue-green, gaster
blue-green with yellowish reflections; man-
dibles yellow-brown with teeth brownish;
legs beyond coxae yellow except pretarsi
black and mid- and hind tarsi cream; an-
tenna with scape yellow, remainder dark
brown; wing veins pale brown. Sculpture
similar to aciculare; T2—7 coriarious except
for smooth band along hind margin. Head
ovate in anterior view, 1.2 x as wide as high
(29:23.5), 1.9 as wide as long (29:15); oc-
ciput concave; clypeus with anterior margin
produced, anterior tentorial pits obscure;
anterior margins of face just laterad of clyp-
VOLUME 90, NUMBER 3
eus not produced; gena with hollow extend-
ing “3 malar length (2:6); toruli 1 diameter
above lower eye margin (2:2); intermalar
distance 3.0 x malar length (18:6); eye height
2.3x malar length (14:6); ratio of LOD:
OOL:POL:LOL as 2:5:7.5:3.5; antenna with
pedicel plus flagellum 1.1 head width
(31.5:29), ratio of lengths of scape : pedicel :
annelli | + 2: funicular segments 1-6: club
as 114:421.5:3:3:3:2.5:2.5:2.5:9.5, Fl 1):5.x as
long as wide (3:2), F6 0.83 x as long as wide
(2.5:3) (Fig. 3), apical segment of club with
subterminal needle-like spine and patch of
micropilosity ventrally (Fig. 4), club sutures
perpendicular, ratio of ventral lengths of
basal: second club segments 1.8 (3.5:2).
Thorax with smooth strip along hind mar-
gin of pronotal collar extending just over
halfway to anterior transverse carina; pro-
podeum as long as scutellum (13:13), nucha
bordered anteriorly by fine but sharp carina.
Forewing (Fig. 5) with ratio of submarginal :
marginal : postmarginal : stigmal veins as 29:
17:13:8; basal vein with 3 setae; basal cell
with setae near basal vein—2 on right wing,
1 on left; costal cell with 1 complete ventral
row of setae. Gastral petiole 1.0 as long
as wide (5.5:5.5) (Fig. 6), median and sub-
lateral carinae weakly developed: gaster
ovate, 1.5 as long as wide (34:22); T1 ex-
tending 0.5 = median length of gaster (17:
34), slightly produced medially, 2 setae
present distal to each posterolateral corner
of basal fovae; all terga visible dorsally
(specimen prepared in critical point dryer
so less telescoping of gastral terga relative
to that in air-dried specimens).
Allotype male.—Thoracic length 0.80
mm. Similar to female except face and
mesoscutum green, pedicel and flagellum
pale brown; antenna with pedicel plus fla-
gellum 1.4 x head width (35:25), ratio scape:
pedicel: annelli | + 2: funicular segments
1-6; club as 12:3.5:1:4:4:3.5:3.5:3.5:3.5:10,
Fl 2.0 as long as wide (4:2), F6 1.8 as
long (3.5:2); gaster 1.7 x as long as wide (29:
17.5), T10.55 x median length of gaster (16:
29).
317
Type material.—Holotype female, Ore-
gon, Benton Co., Mary’s Peak (near Cor-
vallis), 15-VIII-1984, M. E. Schauff, E. E.
Grissell, roadside meadow. Allotype male
and 32 female and | male paratypes with
same data. Fifteen additional paratypes as
follows.—CANADA: Alberta: Lethbridge,
26-VI-1956, 1 9; McMurray 8-VIII-1943, 1
2; 30-VII-1953, 3 2. British Columbia, Hatz-
ic Lake, 22-20-VII-1953, 4 2; Mission City,
26-VH-1953, 1 2. New York: Whiteface
Mountain, 19-VII-1962, 1 2, 1 6. New Mex-
ico: Lincoln National Forest, 28-VII-1977,
1 ¢. Utah: Cache Co., Logan, 7-XI-1954 (al-
falfa), 1 9; Washington: San Juan Island Co.,
Carter’s Point on San Juan Island, 23-VII-
1944, 1 2. Holotype, allotype, and paratypes
in USNM. Additional paratypes in BMNH,
CNC, and INHS.
Etymology.—The species name is de-
rived from the Latin inopinus — unexpected,
unlooked for—referring to the discovery of
this species after several drafts of this paper
had been completed.
Variation. —In females, body color varies
from a basic dark blue to dull green with
yellowish tints; the scape is sometimes dark-
er in the apical fifth. Females from Alberta,
British Columbia, Washington, and Utah
have the scape metallic over most of its
length, and dark bands on the femora like
female 7. aquilonium. The thoracic length
varies between 0.82-0.96 mm. The basal
vein has 2-7 setae with 15 of 20 wings ex-
amined having 4-6. The basal cell has 0-5
setae with 11 of 20 wings examined having
2 or 3. Generally, in those individuals hav-
ing 4 or more setae along the basal vein,
there 1s a partial row of setae apically in the
cubital cell. The male paratype is a little
paler than the allotype and it has strong
yellow reflections on the propodeum and
gaster.
Discussion.—Females of this species
closely resemble those of acicu/are in
structure of club and both sexes in structure
of the nucha but can be distinguished by the
characters given in the discussion section of
318
the latter species. Specimens of inopinum
from western Canada and montane west-
ern United States resemble aquilonium
closely in color. Additional characters to
separate these two species are given in the
discussion section for the latter species.
Toxeuma inopinum is unique in generally
having the smooth strip along the hind mar-
gin of the collar narrow, extending medially
only a little more than halfway to the an-
terior carina.
Merismus Walker
Merismus Walker, 1833: 371, 375. Type
species: Merismus rufipes Walker. Desig.
by Westwood, 1839: 68.
Kentema Delucchi, 1953: 218. Type species:
“Lamprotatus ovatum Walker” (= Mis-
cogaster ovata Walker). Orig. Desig.
Stylomerismus Graham, 1969: 171. Type
species: Merismus (Stylomerismus) ru-
fipes Walker. Orig. Desig. New Synony-
my.
Kentema Delucchi was synonymized with
Merismus by Graham, 1956. The subgenus
Stylomerismus was created by Graham
(1969) to characterize members of the group
of species to which rufipes belongs, but un-
fortunately rufipes is the type of the genus
(and thus subgenus) Merismus. Therefore
the name Stylomerismus is invalid. Boucek
(in litt.) pointed out to us that Kentema
would be the valid subgeneric name if one
were used, but we believe it is sufficient to
recognize the two species-groups proposed
by Graham.
With the findings of this paper, the genus
Merismus is now represented by the follow-
ing world species (the generic placement of
these names is not confirmed and we cite
them as they are currently recognized with-
out attempt at correction): Holarctic: /as-
thenes (Walker), megapterus Walker; Pale-
arctic: nitidus (Walker), rufipes Walker,
splendens Graham; Australian: scutellaris
Dodd and Girault, squamosus Girault. Re-
visions of Palearctic species have been writ-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ten by Graham (1969), Hedqvist (1974) and
Dzhanokmen (1978).
Host records are known only for two Pale-
arctic species, both reared from Agromy-
zidae: M. splendens from Agromyza albi-
pennis Meigen (Graham 1969) and M.
megapterus from Cerodontha (Poemyza)
pygmaea (Meigen) (Graham 1969), C. (P.)
incisa (Meigen) (Graham 1969), C. (P.) pyg-
maea on Deschampsia caespitosa (Hansson,
1987), and Cerodontha (Dizigomyza) ireos
Robineau-Devoidy on /ris pseudacorus
(Hansson 1987). The U.S. National Mu-
seum of Natural History houses specimens
of rufipes reared from “‘wheat stubble” in
France.
Merismus differs from Toxeuma in hav-
ing 3 asymmetrically arranged anterior den-
ticles on the clypeus (Fig. 14); female club
with a linear patch of micropilosity (Figs.
19-20); notauli sometimes shallow poste-
riorly; prepectus smooth, with oblique ca-
rina delimiting posterior triangular area;
propodeum with median carina and plicae
less well developed, usually more rugose;
gastral petiole sometimes also with dorsal
basal flange (Fig. 20), well developed lateral
longitudinal carina absent (Figs. 20, 22); T1
never nearly covering dorsum of gaster.
Merismus lasthenes (Walker)
Figs. 13-20
Sphegigaster Lasthenes Walker, 1848: 108,
165-166, 2. Lectotype, BMNH, exam-
ined.
This species was described apparently
from one female collected in England and
now housed in the British Museum. The
only other known specimen of this species
was reported by Graham (1969) as a female
collected in Scotland. We have discovered
a series of 4 female and 7 male specimens
collected 30 miles north of Fairbanks, Alas-
ka, 30-VII-13-VIII-1984, by S. and J. Peck
from a mixed birchwood forest. These are
in the collection of the CNC (except | 9, 2
6, USNM).
Although it seems unlikely that the Alas-
VOLUME 90, NUMBER 3
\
V3
Ae
Figs. 13-18.
319
Merismus lasthenes. 13, Head, dorsal view, female. 14, Head, anterior view, female. 15, Man-
dibles, dorsal view, female. 16, Antenna, lateral view, female. 17, Antenna, lateral view, male. 18, Forewing,
dorsal view, female. Scale bar = 0.1 mm.
kan specimens would be the same species
as the apparently rare British /asthenes, we
have been unable to find any morphological
structures to distinguish the specimens.
Measurements as well as superposition of
drawings and specimens have given no in-
dication that more than one species 1s in-
volved. Because the male of this species has
never been described and we now can con-
tribute some information on variation in
females, we take this opportunity to rede-
scribe the species as follows (the main de-
scriptive sections are based upon the Alas-
kan specimens; measurements, ratios, and
differences for the lectotype female are giv-
en in brackets):
Female.—Thoracic length 0.9-1.0 mm
[0.9]. Body and coxae blackish metallic
green; smoky yellow [to orange] are: basal
¥4 of scape, tibiae, and tarsi 1-4; mandibles
mahogany; wing veins dark brown. Retic-
ulate sculpture fairly uniform on head, tho-
rax, and coxae; gastral petiole faintly retic-
ulate (compared to thorax) and with irregular
wrinkles; smooth are: clypeus, posterior
margin of pronotum, prepectus, upper mes-
epimeron, metapleuron, lateral sides of pro-
podeum, dorsellum, and dorsum of nucha;
propodeum with irregular median and lat-
eral carinae [no distinct median carina] in
addition to reticulation: gastral terga alu-
taceous. Head in anterior view with inter-
ocular distance 1.3-1.4= eye height [1.3],
in dorsal view 2.1-2.2 x [2.2] as long as
broad, temples converging only slightly and
about '3 length of eye (Fig. 13), POL 1.3-
1.7 length of OOL [1.3], genal hollow ca.
4 distance to eye, clypeal apex as in Fig. 14,
both mandibles with 4 teeth (Fig. 15), scape
barely reaching ventral margin of midocel-
lus, antennal proportions as in Fig. 16, club
with ventral area of micropilosity and rows
of sensillae on last 2 segments [unknown for
lectotype because antenna glued flat to card],
apex of club with spine (Fig. 19, arrow).
Notauli weakly delimited over posterior
third, frenum evenly sculptured much like
rest of scutellum, propodeum 7% length of
scutellum. Forewing (Fig. 18) with marginal
vein 0.9-1.1* postmarginal [0.9], apical
margin of costal cell with complete setal row
on ventral surface but dorsal surface with
setal row only in distal half, ventrally also
are a few scattered setae distally, basal vein
320
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 19-22.
Scale bar
setose, basal cell with 0 to | seta, cubital
vein with basal 73 bare and 3 or 4 setae
distally, speculum open below, wing surface
evenly setose from speculum to apex. Gas-
tral petiole 1.0-1.2 x as long as wide [1.0],
flanged dorsally, slightly shorter than pro-
podeum; gaster ovate in outline, 1.4—1.7 x
longer than wide [1.5], T1 ca. 4 length of
gaster and 0.5-0.7 x as long as wide [0.7],
T2-7 subequal in length.
Male.—Generally similar to female ex-
cept as follows: thoracic length 0.8-0.9 mm;
legs except for coxae entirely straw-yellow;
interocular distance 1.2-1.5x eye height;
POL 1.5-1.8 x OOL; antennal proportions
as in Fig. 17, no area of micropilosity on
venter of club, club without spine at tip but
last segment conical and sharply pointed;
19-20, Merismus lasthenes, scanning electron micrographs. 19, Club of antenna, ventral view,
female (arrow points to apical spine). 20, Gastral petiole, dorsal view, male. 21-22, Merismus megapterus,
scanning electron micrographs. 21, Club of antenna, ventral view, female. 20, Gastral petiole, dorsal view, male.
= 0.05 mm.
marginal vein 0.8-1.0 x postmarginal vein,
0 to 4 setae in basal cell; propodeum varies
from nearly completely reticulate with faint
rugulosity, to irregularly rugulose with in-
terrupted median carina as in Alaska fe-
males, to irregularly rugulose with no me-
dian carina as in lectotype female; gastral
petiole 1.1-1.5 x as long as broad (Fig. 20),
carinate dorsally on distal half; gaster 1.7-
3.0 x longer than wide, tending to telescope
greatly, Tl ca. 3 length of gaster and 0.7-
0.8 as long as wide, T2-7 subequal in
length.
Discussion. — Based upon Graham’s 1969
key to European Merismus and an exami-
nation of rufipes Walker (lectotype 8°,
BMNH), /asthenes (Walker) (lectotype 2,
BMNH), and nitidus (Walker) (paralecto-
VOLUME 90, NUMBER 3
type ¢, 2 additional @ and 2 4, England,
BMNH), /asthenes may be separated from
other species in the rufipes species-group by
the 4-toothed mandibles (left 3, right 4 in
other species) and the short gastral petiole
(1.0 to 1.5< as long as wide and shorter
than the propodeum, versus 1.5 to 2.0 as
long as wide and equal to the propodeum
in other species).
In the Nearctic region, /asthenes may be
distinguished: from megapterus by the
species-group characters, namely Jasthenes
with gastral petiole flanged (Fig. 20; absent
in megapterus, Fig. 22), notauli superficial
posteriorly (complete in megapterus), scu-
tellar frenum evenly reticulate (longitudi-
nally wrinkled with smooth interspaces in
megapterus), and female with spine on api-
cal tip of club (Fig. 19; absent in megapterus,
Fig. 21).
Merismus megapterus Walker
Figs. 21-22
Merismus megapterus Walker, 1833: 377,
6, 2. Lectotype 4, paralectotypes, BMNH,
examined.
Merismus clavicornis Walker, 1833: 377, @.
? Miscogaster tenuicornis Walker, 1833: 462, 2.
Miscogaster ovata Walker, 1833: 462, 2.
Sphegigaster Agriope Walker, 1848: 108,
165, 6.
Merismus megalopterus Schulz, 1906: 143
(emendation).
?Kentema viride Delucchi, 1955: 94, 96, é, 9.
The above synonymy is taken from Gra-
ham (1969) and has not been changed since
his work. Merismus megapterus has been
reported throughout western and central
Europe (Boucek 1977). One of us (EEG) first
discovered specimens of megapterus in ma-
terial reared and submitted for identifica-
tion to the Systematic Entomology Labo-
ratory by Dr. Chris Maier (The Connecticut
Agricultural Experiment Station, New Ha-
ven). Four females and six males emerged
in April 1980 from “grass plants” collected
in March. These were identified initially us-
ing Graham’s key (1969) to European Mer-
32)
ismus, then compared to specimens iden-
tified by Dr. Z. Boucek (Commonwealth
Institute of Entomology, London), and fi-
nally compared with the type material in
1982. Subsequently a large number of spec-
imens were collected and accumulated by
the senior author as part of a study on the
miscogasterine pteromalids.
We have seen a total of 38 females and
48 males of this species from eastern Can-
ada and the northern United States (USNM.,
CNC, INHS, Cornell University, Ithaca,
NY). Records from Canada include Que-
bec, New Brunswick, and Nova Scotia.
United States records include (from west to
east) California, Washington, North Da-
kota, Colorado, Texas, Nebraska, Minne-
sota, Wisconsin, Michigan, Missouri, IlIli-
nois, Indiana, New York, Massachusetts,
Connecticut, Washington, D.C., Virginia,
and West Virginia.
ACKNOWLEDGMENTS
We thank the following individuals for
the loan of specimens: Z. Boucek and J.
Noyes, British Museum (Natural History),
London, England: Chris Maier, The Con-
necticut Agricultural Experiment Station,
New Haven, U.S.A.; G. A. P. Gibson, Bio-
systematics Research Centre, Ottawa, Can-
ada. For reviewing this paper and offering
substantial improvements we thank G. Gib-
son, Tom Henry (Systematic Entomology
Laboratory, Washington, D.C.), and W. E.
LaBerge and G. L. Godfrey (Illinois Natural
History Survey, Champaign, IL).
LITERATURE CITED
Boucek, Z. 1977. A faunistic review of the Yugosla-
vian Chalcidoidea (Parasitic Hymenoptera). Acta
Entomol. Jugosl. 13(Suppl.): 1-145.
Burks, B. D. 1979. Pteromalidae, pp. 768-853. Jn
K. V. Krombein et al., eds., Catalog of Hyme-
noptera in America North of Mexico. Vol. I. Sym-
phyta and Apocrita (Parasitica). Smithsonian In-
stitution Press. Washington, D.C. 1198 pp.
Delucchi, V. 1953. Neue Chalcidier aus der subfam-
ilie der Lamprotatinae. Mitt. Schweiz. Entomol.
Gesell. 26: 201-218.
1955. Contribution a l’etude des Lampro-
322
tatinae. Acta Univ. Lund, (n.s.) Avd. 2, 50(20):
1-97.
Dzhanokmen, K. A. 1978. Pteromalidae, pp. 57-228.
In [Key to the Insects of European Part of the
USSR. Vol. III. Hymenoptera.] Opredelitel: po
faune SSSR, No. 120 Leningrad, “Nauka” Len-
ingradskoe Otedel. 756 pp.
Graham, M. W. R. de V. 1956. A revision of the
Walker types of Pteromalidae (Hym., Chalcidoi-
dea). Part I (Including descriptions of new genera
and species). Entomol. Mon. Mag. 92: 76-98.
1959. Notes on Pteromalidae (Hym., Chal-
cidoidea) with descriptions of new genera and
species. Trans. Soc. Brit. Entomol. 13: 97-112.
. 1969. The Pteromalidae of Northwestern Eu-
rope (Hymenoptera: Chalcidoidea). Bull. Brit.
Mus. (Nat. Hist.). Entomol. Suppl. 16: 1-908.
Hansson, C. 1987. New records of Swedish Eulo-
phidae and Pteromalidae (Hymenoptera: Chalci-
doidea), with data on host species. Entomol. Tidskr.
108: 167-173.
Hedqvist, K.-J. 1974. The genus Merismus Walk. in
Sweden and description of a new genus and species
(Hym. Pteromalidae, Miscogasterinae). Notes on
Chalcidoidea (Hym.). VI. Entomol. Scand. 5: 143-
147.
Heydon, S.L. 1988a. A review of the Nearctic species
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of the genus Cryptoprymna Foerster, with the de-
scription ofa new genus, Po/lstonia (Hymenoptera:
Pteromalidae). Proc. Entomol. Soc. Wash. 90: I-
11.
. 1988b. The Sphegigasterini: A cladistic anal-
ysis and generic classification with reviews of se-
lected genera (Hymenoptera: Pteromalidae). Ph.D.
Thesis, Univ. of Illinois at Urbana-Champaign,
Urbana. 167 pp.
Heydon, S. L. and W. E. LaBerge. 1988. A review of
North American Sphegigaster north of Mexico with
a review of the biology (Hymenoptera: Pteromali-
dae). J. Kans. Entomol. Soc. (In press.)
Schulz, W. A. 1906. Spolia Hymenopterologica. Al-
bert Pape, Paderborn. 355 pp.
Walker, F. 1833. Monographia Chalcidum. Entomol.
Mag. |: 367-384, 455-466.
. 1848. List of the specimens of Hymenopter-
ous insects in the collection of the British Museum.
II. Chalcidites. Additional species. British Mu-
seum, London. 237 pp.
Westwood. 1839. Synopsis of the genera of British
insects, pp. 1-158. Jn An Introduction to the Mod-
ern Classification of Insects. Part II. [Dated 1840
but issued in 1839], Longman et al., London. 587
pp.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 323-328
NOMENCLATURAL NOTES ON POLISTINAE
(HYMENOPTERA: VESPIDAE)
JAMES M. CARPENTER AND MICHAEL C. DAY
(JMC) Museum of Comparative Zoology, Harvard University, Cambridge, Massachu-
setts 02138, U.S.A.; (MCD) Department of Entomology, British Museum (Natural His-
tory), Cromwell Road, London SW7 SBD, United Kingdom.
Abstract. —Correct nomenclature is supplied for certain genus-group names of paper
wasps. The infrasubgeneric names of Saussure (1852-58) were made available in 1985
from the original dates of publication. A/pha Saussure, 1854, is thus a senior synonym of
Hypopolybia Richards, 1978; Phi Saussure, 1854, is a senior synonym of Monocyttarus
Richards, 1978. Trichinothorax is newly proposed as a replacement name for Trichothorax
Richards, 1978, non Montrouzier, 1860. Agelaia Lepeletier, 1836, is a senior synonym
of Stelopolybia Ducke, 1910.
Key Words:
During the course of various studies on
the paper wasps, certain nomenclatural
problems became apparent to both authors.
The purpose of this note is to correct the
nomenclature of the genus-group names.
GREEK LETTERS
The difficulties stem in large part from
the infrasubgeneric names proposed by
Saussure. In Volume 2 of his great work,
Etudes sur la Famille des Vespides (1852-
58), Saussure recognized 13 genera in the
tribe ““Vespiens”’ (a taxon equivalent to the
Vespidae of Richards 1962). He divided the
genus Polybia Lepeletier into two subgen-
era, Clypearia and Polybia ‘“‘proprement
dites,” and subdivided the latter into seven
“divisions.”” These divisions were Alpha,
Iota, Phi, My, Kappa, Omega and Para-
polybia. Subsequently, Saussure (1863) rec-
ognized Pseudopolybia as a division of the
genus Polybia.
Volume 2 of “Etudes” was published be-
tween 1853 and 1858; the divisions were
proposed in 1854 (see Griffin 1939). Shortly
Nomenclature, Polistinae, social wasps
thereafter, both Alpha and Phi were used
again by Saussure in Volume 3 of “Etudes”
(1854-1856; the part mentioning these
names appeared in 1855). The names were
applied to divisions of the eumenine genus
Eumenes (p. 137 and 145, respectively), and
Alpha was further used for a division of
Montezumia (p. 160). Other names derived
from Greek letters were used for divisions
of genera in this work. Delta, Epsilon, Omi-
cron and Zeta are currently applied to eu-
menine genera, and Beta was also used in
both Eumenes and Montezumia (see Car-
penter 1986). Saussure applied these names
to divisions first delineated in Volume | of
“Etudes” (1852-1853), where they had been
referred to by Roman numerals. In Volume
1, Saussure also delineated sections of some
of these divisions. These were referred to
typically by capital letters or arabic numer-
als, but in Zethus he also used the letters
“a and “8.” This practice was continued
throughout the “Etudes.” From thisitis plain
that Saussure was merely hierarchically par-
titioning his genera according to the prin-
324
ciples of Aristotelean division, and did not
consider these names to be validly pro-
posed. As explained in a footnote on p. 167
of Volume 2, he began applying “arbitrary”
names instead of numerals in order to fa-
cilitate interpolation of new taxa into his
classification, without the necessity of re-
numbering. Nevertheless, the criteria of
availability now in the International Code
of Zoological Nomenclature make it clear
that these names, even if proposed unin-
tentionally, are available.
Subsequent to Saussure, Dalla Torre
(1894: 161) treated Pseudopolybia as a ge-
nus, and Bingham (1897: 382) did the same
for Parapolybia. Dalla Torre (1904: 76) list-
ed the other polistine secondary names as
divisions of his subgenus Eupolybia, but
otherwise these names were generally ig-
nored. Then Bequaert (1933: 112) trans-
ferred Kappa and Omega to the synonymy
of Mischocyttarus Saussure, and designated
a type species for the former. Richards
(1941) subsequently raised them to subgen-
era. Later, Bequaert (1944a: 99) fixed the
identity of Alpha by designating a type
species, and Bequaert (1944b: 292) did the
same for My (as Mu) and Phi by designating
type species for them. Finally, Richards
(1978: 33) selected a type species for Jota.
However, it was Richards’ monograph
which crystallized the present problem.
The International Code of Zoological No-
menclature in force prior to 1985 did not
treat infrasubgeneric names as available. As
mentioned above, Saussure described nu-
merous “divisions” of subgenera in his
Etudes, especially in the eumenine genus
Odynerus. When the present trend of split-
ting this genus was begun 50 years ago by
Bliithgen (1938), these divisions gradually
came to be treated as genera (see Carpenter
and Cumming 1985, and Carpenter 1986).
In order to fix their status, van der Vecht
(1967) sought an Opinion ruling these names
as available from their original dates of pub-
lication, with original authorship. Opinion
893 (ICZN 1970) so ruled.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Richards (1978) did not follow this course
in Polistinae. While stating correctly that
infrasubgeneric names had no nomencla-
tural status under the Code in force at that
time, he misunderstood van der Vecht’s
treatment of the analogous case in Eumen-
inae, which Richards erroneously stated as
pertaining to primary divisions of genera.
Of course, van der Vecht’s proposal would
have been unnecessary if this were true! Fol-
lowing the Code, Richards treated the names
as first made available when they were given
nomenclatural standing by subsequent au-
thors, either as subgenera or by designation
of type species, with these later workers as
authors of the names. Thus he attributed
Parapolybia to Bingham (1897), Pseudo-
polybia to Dalla Torre (1894), and Kappa,
Omega and Alpha to Bequaert (1933) [4/-
pha should have been cited as dating from
Bequaert 1944a]. Under this procedure
Richards treated Omega as a junior objec-
tive synonym of Monacanthocnemis Ducke,
1905 in his subgeneric arrangement of Mis-
chocyttarus. Richards’ classification was
therefore inconsistent with the precedent es-
tablished in Vespidae by van der Vecht.
It is unfortunate that Richards miscon-
strued the situation, although to have fol-
lowed van der Vecht’s precedent would have
required submission of another case to the
Commission. However, no argument on the
point is now required. Under Art. 10(e) of
the revised International Code of Zoological
Nomenclature, secondary divisions are val-
id as genus-group names from their original
date of publication, with original author-
ship. Richards’ classification must therefore
be amended. Richards himself, in a list of
errata and addenda to his monograph pub-
lished in 1983, reversed his stance 1n part,
accepting Saussure’s authorship of Pseu-
dopolybia and Omega. In addition, Rich-
ards (1978), although citing Bequaert
(1944b), overlooked that that author had
designated type species for Phi and My, and
as a result Richards’ subgenus Monocytta-
rus is a junior subjective synonym. Thus,
VOLUME 90, NUMBER 3
Alpha and Phi become valid names for cur-
rent subgenera of Polistinae and preoccupy
the use of these names in the Eumeninae
(Carpenter 1986). The corrected synony-
mies of the relevant taxa are listed below.
But first two unrelated matters must be ad-
dressed.
HOMONYMY
Richards proposed a subgeneric classifi-
cation of Polybia in his monograph. The
naturalness of his arrangement remains to
be investigated, but that aside, one of his
subgenera is a junior homonym. As pointed
out by Day (1979), Trichothorax Richards,
1978 is preoccupied in Coleoptera (by Mon-
trouzier 1860). A replacement name is here-
with proposed. This name is the replace-
ment intended by Richards, as shown by an
unpublished manuscript at the British Mu-
seum.
THE IDENTITY OF AGELAIA LEPELETIER
The final matter concerns Agelaia fusci-
cornis Lepeletier, 1836. Since its proposal
this taxon has been unrecognized. Saussure
(1854: 210) placed it in Polybia, while
pointing out the similarity in color of its
description to that of Apoica pallida and
Polybia testacea (now in Stelopolybia). Fol-
lowing him Dalla Torre (1894: 161) placed
Agelaia in the synonymy of Polybia. How-
ever, Bequaert (1944b: 254) suggested that
it may have been a Polistes. After success
in identifying Aphanilopterus Meunier from
its description (see Richards 1978: 437), J.
van der Vecht inquired whether one of us
(MCD) might also be able to identify 4ge-
laia. It was determined that the description
almost certainly applies to Stelopolybia tes-
tacea (F.). The evidence for this is outlined
next.
Lepeletier (1836: 535) described Agelaia
as a member of his fourth family, ““Les Po-
listides.”” He mentioned (p. 536) that noth-
ing was known of its habits, nevertheless
the name was derived from the Greek for
“Vivant en société.”” The placement of Age-
325
laia, in between Polybia and Apoica, and
the similarities in the details of the descrip-
tions of these taxa, indicate that Agelaia
must be a social wasp. Lepeletier stated that
the collecting locality was unknown, but the
color pattern he described, largely ferrugi-
nous with the posterior part of the meta-
soma blackish, is that ofa well known South
American mimicry complex and does not
occur elsewhere. Three South American so-
cial wasps with this color pattern and of the
right size (10 lines, or approximately 21 mm,
cf. Mayr 1969) are known to us: Stelopo-
lybia testacea, Mischocyttarus flavicans and
Polistes testaceicolor. Two solitary vespids
of similar color pattern and size (Pachy-
menes orellanae and Montezumia analis)
may be dismissed because the clypeus is not
angular below. Only the Ste/opolybia
matches the description of the metasomal
petiole. The petiole is described as almost
conical and tuberculate laterally. The meta-
soma of Polistes testaceicolor is as in other
species of this genus described by Lepele-
tier: “*. . . sans pédicule distinct; son premier
segment se dilatant en cloche dés sa base.”
The petiole in Mischocyttarus flavicans is
elongate, as in the species of Apoica de-
scribed by Lepeletier. The second submar-
ginal cell (““cubitale’’) in .4gelaia is described
as scarcely narrowed towards the marginal
(“radiale’’) cell and little dilated towards the
discal cell. Saussure (1854: 210) considered
this feature the main obstacle in identifying
Agelaia fuscicornis as Stelopolybia testacea.
Saussure characterized the latter species as
having this cell strongly (‘‘entiérement’’)
narrowed, as in other paper wasps. Lepe-
letier was possibly in error, however the cell
in S. testacea is narrower posteriorly rela-
tive to the other two species, and so it ap-
pears less narrowed anteriorly. If this inter-
pretation is correct, the only discrepancies
concern the color pattern. The antennae are
described as blackish above, and the second
metasomal segment as blackish on the pos-
terior third. Most specimens of Stelopolybia
testacea which we have seen have the joints
326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
between the flagellomeres slightly darkened,
but the antennae are otherwise ferruginous,
and the second metasomal tergum has a fer-
ruginous band apically. However the anten-
nae do appear blackish if viewed with the
naked eye, and the extent of black and fer-
ruginous on the second tergum varies great-
ly. We regard these discrepancies as minor
compared to the general correspondence to
the description. In the absence of the orig-
inal material there can be no absolute cer-
tainty, but the description appears adequate
to recognize this taxon. Agelaia fuscicornis
Lepeletier, 1836 is thus considered here a
junior synonym of Stelopolybia testacea (F.,
1804).
This resolution of the identity of Agelaia
creates another problem. Age/aia Lepele-
tier, 1836 is a senior synonym of Ste/opo-
lybia Ducke, 1910. We regard replacement
of the name Stelopolybia by Agelaia with
equanimity, since we believe that stability
is best served through strict application of
the Principle of Priority. We do not think
this replacement will cause more than tem-
porary confusion, and the situation will
henceforth be stable. However, other work-
ers may disagree, in which case an appeal
to the Commission for suppression of Age-
/aia under the plenary powers should be
made. Were adoption of Age/aia the only
change required in an othewise stable no-
menclature, more force might accrue to an
argument for suspension of the rules. In par-
ticular, we note with interest that the ma-
jority of changes necessitated here arise from
the Commission’s recent adoption of re-
vised criteria of availability with respect to
infrasubgeneric names. It would have been
so easy to make these criteria dependent on
previous accepted usage for extant names!
CoRRECT SYNONYMY
Polybia Lepeletier, 1836: 533. Type species
Polistes liliacea F., 1804. By subsequent
designation of Ashmead, 1902: 166.
subgenus Polybia
Tota Saussure, 1854: 174, explanations
to plates 22 and 24 (as division of
subgenus Polybia). Type species ‘‘Po-
lybia liliacea (F.).” By subsequent
designation (Richards 1978: 33) un-
der Art. 67(f).
Jota Dalla Torre, 1894: 161. Unjusti-
fied emendation of Jota.* New syn-
onymy.
Eupolybia Dalla Torre, 1904: 76. New
name for Polybia. Type species ‘*Po-
lybia liliacea (F.) (= Polistes liliacea
Fabricius, 1804). By subsequent
designation of Richards, 1978: 33.
Iota Richards, 1978: 33, 46; non Iota
Saussure, 1854. Unjustified emen-
dation of Jota. Unavailable under
Art. Ile.
subgenus A/pha Saussure, 1854: 167, ex-
planations to plates 21 and 22 (as di-
vision of subgenus Polybia). Type
species Polybia bifasciata Saussure,
1854. By subsequent designation of Be-
quaert, 1944a: 99. New status.
Hypopolybia Richards, 1978: v, 35, 52
(subgenus of Polybia). Type species
Polybia_ bifasciata Saussure, 1854.
Original designation. New synony-
my.
subgenus Myrapetra White, 1841: 320.
Type species Myrapetra scutellaris
White, 1841. By indication (monotyp-
ic).
Myraptera Saussure, 1854: 192, 194,
211, 249. Incorrect subsequent spell-
ing of Myrapetra.
My Saussure, 1854: 191, explanation
to plate 23 (as division of subgenus
Polybia). Type species Myrapetra
scutellaris White, 1841. By subse-
quent designation (Bequaert 1944b:
292) under Art. 67(f).
Mi Dalla Torre, 1904: 76. Unjustified
emendation of My. New synonymy.
Mu Bequaert, 1944b: 292. Unjustified
emendation of My. New synonymy.
*Note that unjustified emendations are available
names (Art. 33b(iii)).
VOLUME 90, NUMBER 3
subgenus Trichinothorax new name for
Trichothorax Richards, 1978, q. v.
Pseudopolybia Thering, 1896: 452, ju-
nior homonym of Pseudopolybia
Saussure, 1863. Type species Polistes
ignobilis Haliday, 1836. By subse-
quent designation of Bequaert, 1933:
1s
Trichothorax Richards, 1978: v, 35, 101
(subgenus of Polybia); non Montrou-
zier, 1860: 235. Type species Vespa
chrysothorax Lichtenstein, 1796.
Original designation.
Trichthorax Snelling, 1981: 416. In-
correct subsequent spelling of 7vi-
chothorax.
Mischocyttarus Saussure, 1853: 19. Type
species Zethus labiatus Fabricius, 1804.
By subsequent designation of Ashmead,
1902: 166.
Mischocytharus Saussure, 1853: viii,
footnote in Introduction to Vol. 1. In-
correct original spelling of Mischocyt-
tarus.
Mischocittarus Saussure, 1857: xi. Incor-
rect original spelling of Mischocyttarus.
Mischocythorus Krombein, 1979: 1516.
Incorrect subsequent spelling of Mis-
chocytharus.
subgenus Kappa Saurssure, 1854: 200 (as
division of subgenus Po/lybia). Type
species Polybia injucunda Saussure,
1854. By subsequent designation of Be-
quaert, 1933: 112.
subgenus Phi Saussure, 1854: 183, expla-
nations to plates 23 and 24 (as division
of subgenus Polybia). Type species Ves-
pa phthisica Fabricius, 1793. By sub-
sequent designation of Bequaert, 1944b:
292. New status.
Monocyttarus Richards, 1978: vi, 18,
273, 307. Type species Polybia flav-
itarsis Saussure, 1854. Original des-
ignation. New synonymy.
subgenus Omega Saussure, 1854: 206 (as
division of subgenus Po/ybia). Type
species Polybia filiformis Saussure,
1854. By indication (monotypic).
327
Monacanthocnemis Ducke, 1905: 6, 8,
21. Type species Polybia filiformis
Saussure, 1854. By indication
(monotypic).
Parapolybia Saussure, 1854: 207 (as divi-
sion of subgenus Polybia). Type species
Polybia indica Saussure, 1854. By sub-
sequent designation of Bingham, 1897:
382.
Pseudopolybia Saussure, 1863: 237 (as di-
vision of genus Polybia). Type species Po-
lybia vespiceps Saussure, 1863. By indi-
cation (monotypic).
Agelaia Lepeletier, 1836: 535. Type species
Agelaia fuscicornis Lepeletier, 1836 (=
Polistes testacea F., 1804). By indication
(monotypic).
Aglaia Dalla Torre, 1904: 75. Incorrect sub-
sequent spelling of Agelaia.
Stelopolybia Ducke, 1910: 452, 464, 517.
Type species Polistes angulata F., 1804.
By subsequent designation of Lucas,
1912: 210. New synonymy.
Gymnopolybia Ducke, 1914: 316, 317,
327. Type species Polybia vulgaris
Ducke, 1904 (= Vespa fulvofasciata
DeGeer, 1773). By subsequent desig-
nation of Richards, 1943: 45. New syn-
onymy.
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(Nat. Hist.), London. 4 pp.
Saussure, H. F. de. 1852-1858. Etudes sur la Famille
des Vespides, Vols. 1-3. V. Masson, Paris, and J.
Cherbuliez, Geneva. [For publication dates of the
various parts see Griffin 1939].
. 1863. Mélanges Hyménoptérologiques. Mém.
Soc. Phys. Hist. Nat. Genéve 17: 171-255.
Snelling, R. R. 1981. Systematics of social Hyme-
noptera, pp. 369-453. Jn Hermann, H. R., ed.,
Social Insects I]. Academic Press, New York.
Van der Vecht, J. 1967. The status of certain genus-
group names in the Eumenidae (Hymanoptera,
Vespoidea). Z.N.(S.) 1689. Bull. Zool. Nomencl.
24: 27-33.
White, A. 1841. Description of a South American
wasp which collects honey. Ann. Mag. Nat. Hist.
7: 315-322.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 329-333
FIRST RECORD OF THE GENUS PROCANACE HENDEL FROM
NORTH AMERICA, WITH THE DESCRIPTION OF A
NEW SPECIES (DIPTERA: CANACIDAE)
WAYNE N. MATHIS
Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C.
20560.
Abstract. —The genus Procanace, previously known only from the Indopacific, Hendel
is reported from North America for the first time, and a new species, P. dianneae, is
described and illustrated from specimens collected in Virginia. A revised key to the genera
occurring in North America is presented, as well as one to the species groups of Procanace.
Key Words:
Beach flies of the genus Procanace Hendel
were previously known from littoral habi-
tats, both marine and freshwater, within the
basins of the Pacific and Indian oceans (Af-
rotropical, Oriental, eastern Palaearctic, and
Oceanian regions). No species were known
from the Western Hemisphere, and it was
an unexpected surprise to discover a new
species of Procanace on the tidal shores of
the Potomac River in peninsular Virginia,
thousands of kilometers from its nearest
congener and within a drainage system of
the Atlantic Ocean.
A further anomaly of this discovery con-
cerns the climate of the area. Although a
few species of Procanace occur in temperate
regions (Japan), greater species diversity oc-
curs in the tropics and subtropics. Thus, the
discovery of a new species in temperate Vir-
ginia was again unanticipated.
The purpose of this paper is to describe
the new species, provide a diagnosis to the
genus and an annotated key to its species
groups, and to present a revised key to the
genera of the family Canacidae that occur
in North America. The key to genera will
essentially be an updating of Wirth’s key
(1987) that was recently published in the
Diptera, Canacidae, Procanace, species groups, new species
Manual of Nearctic Diptera, Volume 2. In
addition to Procanace, the generic key also
includes Paracanace Mathis and Wirth even
though that genus has not yet been recorded
from the United States. Paracanace has been
found in Cuba and Mexico, however, and I
would expect it to occur along the Gulf Coast
in the southeastern United States. The
chapter on the family Canacidae in the
Manual should be consulted for additional
details on the family, its biology, and char-
acterization.
It is timely that this species be described
now so that its record, including that of the
genus, can be incorporated in a checklist of
Nearctic Diptera that is being prepared.
Methods.—The descriptive format used
in this paper essentially adheres to that which
I have published elsewhere in the family
Canacidae (Mathis 1988). The terminology
used for anatomical structures follows
McAlpine (1981) with the exceptions that
have been noted previously (Mathis 1986).
For the convenience of the user, the defi-
nition of M vein ratio of the wing is: the
straight line distance along M_ between
crossveins rm and dm-cu/distance apical of
crossvein dm-cu.
330
Key TO GENERA OF CANACIDAE IN
NortTH AMERICA
1. Lateroclinate fronto-orbital setae 4. Presutural
acrostichal setae present; acrostichal setulae
numerous, in four irregular rows. Female cer-
cus with | large apical seta that is usually acute-
ly pointed (subfamily Canacinae)
Dey ee ee ren eet Canacea Cresson
~ Lateroclinate fronto-orbital setae 3. Presutural
acrostichal setae absent; acrostichal setulae
sparse or absent. Female cercus with 2 large
setae, | apical and | subapical, each usually
rather broadly rounded (subfamily Noctican-
acinae) . Se ae et, Settee te amt meee sees 2
Interfrontal setae absent, although anterior 1/3
of frons occasionally with scattered setulae . .
hg eco diets mn ee Seen e Procanace Hendel
— Interfrontal setae present, | or more pairs in
additional to any setulae ....
3. Two interfrontal setae present; postocellar se-
tae well developed, proclinate and slightly di-
vergent Paracanace Mathis and Wirth
— One interfrontal seta present; postocellar setae
either much reduced or absent . = 4
4. Disc of scutellum with setae; 3 large anaclinate
genal setae .. Canaceoides Cresson
— Disc of scutellum lacking setae; 2 anaclinate
genal setae . Nocticanace Malloch
to
Genus Procanace Hendel
Procanace Hendel 1913: 93. Type species:
Procanace grisescens Hendel, by original
designation.
Diagnosis.—General coloration whitish
gray, olivaceous, to blackish brown.
Head. Interfrontal setae absent, but with
a few setulae inserted anteriorly; fronto-or-
bital setae 3; ocelli arranged to form equi-
lateral or isosceles triangle, if isosceles, the
greater distance is between posterior ocelli.
Arista pubescent over entire length. Two
large anaclinate genal setae; anteroclinate
genal seta moderately well developed. Pal-
pus not bearing long setae. Epistomal mar-
gin, in lateral view, more or less horizontal.
Thorax: Acrostichal setae, especially a
prescutellar pair of large setae, usually lack-
ing (setulae present in species of the wil-
liamsi group); scutellar disc lacking setae (1-
2 pairs of scutellar disc setulae occur in P.
nakazatoi Miyagi of the wi/liamsi group); 2
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pairs of marginal scutellar setae, apical pair
not anaclinate; anterior and posterior no-
topleural setae present, length of both sub-
equal; anepisternum with scattered setulae.
Katepisternal setal usually present (lacking
in species of the grisescens group). Hind tib-
ia lacking spine-like setae apically.
Abdomen: Male genitalia as follows:
Epandrium in posterior view wider than
high; cerci reduced, poorly sclerotized; sur-
stylus with an anterior and posterior lobe,
the latter larger, sometimes markedly so and
shape unique to species.
Discussion. —Procanace is probably a
monophyletic taxon, although the evidence
is weak, i.e. the lack of interfrontal setae
and the more or less horizontal epistomal
margin. The possibility remains that the ge-
nus is paraphyletic, which isa common con-
dition for groups that include disjunct
species.
ANNOTATED Key TO SPECIES
Groups OF PROCANACE HENDEL
1. Katepisternal seta absent .... the grisescens group
4 species; Oriental, eastern Palaearctic, Oceanian,
Malagasy, Seychelles
= “Katepisternal'seta‘present 57. .-2....2.2.02% 2
2. Clypeus high, width about twice the height;
palpus blackish brown; proepisternal seta ab-
SOME 4H. cc:Aaec asaya agerteee the nigroviridis group
7 species; Hawaiian Islands
— Clypeus low, width at least 4 times the height;
palpus yellowish; proepisternal seta(e) present
3: Acrostichal setulae present, in 2 irregular rows
PrN PRE rial Cyaan the williamsi group
4 species; Hawaiian and Ryukyu Islands
— Acrostichal setulae absent ............. 4
4. Postocellar setae either absent or anche re-
Guced’s. es SNA PS: the fulva group
9 species; Oriental and eastern Palaearctic
— Postocellar setae present, subequal to length of
Ocellar'SCtay says sae cea eusetees the cressoni group
3 species; Oriental, Nearctic
Procanace dianneae Mathis,
New SPECIES
Fig. 1-3
Diagnosis. —Extenally this species is very
similar to those of the cressoni group, and
VOLUME 90, NUMBER 3 331
5th
0.40mm
4th
3
Figs. 1-3. Procanace dianneae. 1, Epandrium and surstylus, lateral view. 2, Surstyli, posterior view. 3, Sterna
4-5 of male, ventral view.
332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
I am tentatively placing it in that group. It
differs from the two species of that group,
P. cressoni Wirth and P. taiwanensis Del-
finado, as well as other congeners by the
following combination of characters: Pos-
tocellar setae well developed, subequal in
length to ocellar setae; clypeus low, height
Y, width; palpus yellowish. Scutum mostly
bluish black, sparsely microtomentose, scu-
tum densely microtomentose, brown; pro-
episternal seta present, pale; katepisternal
seta present; acrostichal setae absent. Shape
of the male genitalia unique (see figs. and
description below).
Description.— Moderately small to me-
dium-sized beach flies, length 2.0 to 3.1 mm;
general coloration whitish gray, olivaceous
to brown, scutum darker.
Head: Frons with ocellar triangle and
fronto-orbits mostly grayish but usually with
some golden coloration, especially on an-
terior half of fronto-orbits, mesofrons most-
ly golden brown but with some rust to red-
dish coloration toward base of ocellar
triangle; anterior half of mesofrons bearing
about 10 setulae; postocellar setae well de-
veloped, length subequal to that of ocellar
setae; middle fronto-orbital seta inserted
slightly closer to posterior fronto-orbital seta
than to anterior seta. First and 2nd antennal
segments and arista dark colored, grayish
black; Ist flatellomere reddish brown to
brown: palpus yellow; face mostly white but
with faint bluish to olivaceous coloration;
gena mostly concolorous with face, poste-
rior portion slightly more olivaceous. Gena
bearing | anteroclinate and 2 anaclinate,
well-developed setae and 2 setulae between
them. Clypeus low, height about 4 width.
Thorax: Scutum mostly subshining, bluish
black, sparsely microtomentose but micro-
tomentum becoming denser laterally, whit-
ish or olivaceous gray to brown; scutellum
densely microtomentose, brown; pleural
areas densely microtomentose, mostly gray
but with some olivaceous and light brown
coloration. Acrostichal setulae absent; 3rd
dorsocentral seta inserted at level of or an-
terior to supra-alar seta; anterior notopleu-
ral seta present, well developed, size sub-
equal to posterior seta; proepisternal setae
present but pale colored; anepisternum with
few scattered setulae, mostly in more or less
vertical arrangement in middle and along
posterior margin; katepisternal seta present,
well developed, katepisternum with several
setulae anterior to large seta. Femora, tibiae,
and most tarsomeres of male yellowish with
light dusting of whitish gray microtomen-
tum on dorsal surface, females with micro-
tomentum on legs more extensive and dark-
er; tarsomeres becoming darker apically,
apical 1-2 tarsomeres blackish; fore femur
bearing 5-6 moderately long and evenly
spaced setae along posteroventral margin,
basal 1-2 pale; mid femur bearing row of
setae, these more evident and closely set on
apical 3. Wing with length of apical section
of vein CuA, moderately long, subequal to
length of crossvein dm-cu; M vein ratio 0.6.
Abdomen: Unicolorous, olivaceous gray
with some faint brownish coloration. Male
abdomen as follows: 4th sternum (Fig. 3)
narrowly rectangular, over 2 as long as
wide; 5th sternum (Fig. 3) wider than long,
width of anterior margin subequal to that
of 4th sternum, becoming wider posteriorly,
lateral margins irregular, widest at posterior
margin, bearing a short process postero-
laterally; epandrium wider than high in pos-
terior view, bearing numerous setae, in lat-
eral view (Fig. 1) posterodorsal margin
broadly rounded, ventral margin nearly flat,
anterior margin nearly straight except for
anteroventral prong and irregular dorsal 1;
surstylus (Figs. 1, 2) as 2 processes, anterior
one much smaller, digitiform, bearing sev-
eral setulae preapically and apically, pos-
terior process much larger, length nearly
equal to that of epandrium and equally as
wide, in lateral view with posterior margin
irregularly arched, anteroventral process
very angulate in lateral view and spatulate
in posterior view.
Type material.—The holotype male is la-
beled “USA. VIRGINIA[:] Westmoreland
VOLUME 90, NUMBER 3
Co. & Park (bank Potomac River)[,] 9 Oct
1987[,] W. N. & D. Mathis.” Allotype fe-
male and 87 paratypes (65 6, 22 2; USNM)
bear the same label data as the holotype.
The holotype is double mounted (minute
nadel in a plastic elastomer block), is in ex-
cellent condition, and is deposited in the
National Museum of Natural History,
Smithsonian Institution.
Etymology.—It is a pleasure to name this
species after my wife, Dianne, in recogni-
tion of her many contributions to my stud-
ies of Diptera. Dianne was also a collector
of the type series.
Natural history.—All specimens of the
type series were collected from the shoreline
of the tidal portion of the Potomac River
at Westmoreland State Park. At the park,
the river is over a mile wide, due largely to
the tidal influence, and the water 1s slightly
brackish. The shore is either almost entirely
sand, the bathing area of the beach, or a
combination of sand, considerable gravel,
and some cobble and large rocks. In the
latter habitat, the shore is quite narrow, at
most two to three meters, and immediately
adjacent to the shore is a cliff. In the sandy
area, specimens occurred along the protect-
ed sides of narrow, wooden jetties that were
installed perpendicular to the shoreline to
break up the action of waves and prevent
erosion of the beach. In the sand/cobble/
rock habitat, specimens were found only on
the rocks and were easily collected by
sweeping immediately over and between the
rocks. Most of the rocks and jetties were
333
covered in part with algae, and I suspect
that the larvae of this species were feeding
on them.
Remarks. — The Chesapeake Bay is one of
the busiest commercial waterways in the
world, and I do not dismiss the possibility
that this species, albeit previously un-
known, was introduced in conjunction with
the large volume of traffic on these waters.
ACKNOWLEDGMENTS
The illustrations were skillfully prepared
by George Venable on a MacIntosh II com-
puter, and a draft of this paper was critically
reviewed by Oliver S. Flint, Jr. and Norman
E. Woodley. I thank these individuals for
their contributions to this paper.
LITERATURE CITED
Hendel, F. 1913. H. Sauter’s Formosa-Ausbeute:
Acalyptrate Musciden (Dipt.), II. Suppl. Entomol.
2: 77-112.
Mathis, W. N. 1986. Studies of Psilopinae (Diptera:
Ephydridae), I: A revision of the shore fly genus
Placopsidella Kertész. Smithson. Cont. Zool. 430:
1-30.
1988. Beach flies of the Republic of Sey-
chelles (Diptera: Canacidae). Bull. Biol. Soc. Wash.
8: 22-29.
McAlpine, J. F. 1981. Morphology and terminolo-
gy—adults [chapter], pp. 9-63. Jn McAlpine, J. F.
et al., eds., Manual of Nearctic Diptera. Vol. 1.
Mono. 27, Res. Branch, Agri. Canada. Hull, Que-
bec.
Wirth, W. W. 1987. Canacidae [chapter 102], pp.
1079-1083. In McAlpine, J. F., ed., Manual of
Nearctic Diptera. Vol. 2. Mono. 28, Res. Branch,
Agri. Canada. Hull, Quebec.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 334-337
FIRST RECORD OF THE SHORE-FLY GENUS PLACOPSIDELLA
KERTESZ FROM NORTH AMERICA (DIPTERA: EPHYDRIDAE)
WAYNE N. MATHIS
Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C.
20560.
Abstract.—The shore-fly genus Placopsidella Kertész is reported from North America
for the first time. The species, P. grandis (Cresson), was previously known from littoral
habitats within the Pacific and Indian Ocean basins, with probable introductions into the
eastern Mediterranean (Israel). This introduction apparently follows a similar pattern for
other recent ephydrid introductions into North America, and these are apparently abetted
by the large volume of commerce on the lower Chesapeake Bay and the natural history
of the flies.
Key Words:
Peninsular Virginia and the Eastern Shore
of Maryland and Virginia, especially their
seashores and tidal rivers, appear to be focal
points for the introduction of new taxa into
North America. During the last two years
alone, we have reported the introduction of
two other shore and beach flies (Mathis and
Steiner 1986, Mathis 1988) to these areas,
and the purpose of this paper is to report
yet another. The introduced shore fly, Pla-
copsidella grandis (Cresson), is the first oc-
currence of that genus and species in North
America, although previously it was re-
ported from Panama, the first record from
the Western Hemisphere (Mathis 1986).
I have adopted a taxonomic format to
present the published history of the intro-
duced species, chiefly to save space, and have
attempted to provide all references to the
species. Inasmuch as Placopsidella was re-
cently revised (Mathis 1986), I have not re-
peated the charcterizations of the genus and
species. A brief diagnosis for the genus and
species, however, plus figures of distinguish-
ing characters are provided to facilitate
identification. Consult Mathis (1986) for
Diptera, Ephydridae, Placopsidella grandis (Cresson), introduction
further information on the genus and its in-
cluded species.
Placopsidella grandis (Cresson)
Figs. 1-5
Gymnopa grandis Cresson 1925: 232; 1945:
54 [review].
Placopsidella grandis. —Cogan and Wirth
1977: 323 [comb.; Oriental cat.].— Math-
is 1986: 22-25 [revision].
Placopsidella opaca Miyagi 1977: 30-31.—
Cogan 1984: 129 [Palearctic cat.].—
Mathis 1986: 23 [syn. with P. grandis].
Mosillus grandis. —Tenorio 1980: 268
[comb.; review of Hawaiian species].
Diagnosis. — Placopsidella belongs to the
tribe Gymnopini and is closely related in
the North American fauna only to Mosillus
Latreille. Like the latter, the ocellar and
pseudopostocellar setae are either much re-
duced or absent, there is a single notopleural
seta that is inserted in the posterior angle
of the notopleuron, and the arista is bare or
nearly so. Placopsidella is distinguished from
Mosillus and other genera of Gymnopini by
VOLUME 90, NUMBER 3
335
Figs. 1-S.
its grayish brown to brown microtomentose
vestiture, the absence of an outer vertical
seta (the inner seta is present, although only
moderately well developed), and the pres-
ence of four to six large setae between the
postalar seta and the base of the scutellum.
Other genera of Gymnopini are mostly
shining black, nearly bare of microtomen-
tum, and there are no large setae between
the postalar seta and the base of the scutel-
lum. In Wirth et al.’s key (1987) to the gen-
era of North American Ephydridae, Pla-
copsidella keys to the first half of couplet 39
(Mosillus) but is distinguished by the char-
Placopsidella grandis. 1, Head, lateral view. 2, Head, anterior view. 3, Thorax, dorsal view. 4,
Male genitalia, lateral view. 5, Male genitalia, posterior view.
acters just outlined. Placopsidella is strictly
coastal in distribution, mostly intertidal,
whereas Mosil/lus occurs inland.
Within Placopsidella, this species belongs
to the /iparoides species group and is dis-
tinguished from congeners by its smaller size
(length 2.7 to 3.7 mm), silvery white facial
microtomentum, pattern of shining areas on
face, conically prominent face, the ante-
riormost aspect of the facial prominence
shining, darker antenna, number of scutel-
lar bristles (two), and the unique confor-
mation of the male genitalia (see figs.).
Canzoneri (1986) recently described P.
336
rossii from a specimen collected in West
Africa (Sierra Leone). That species is ap-
parently very similar to P. grandis and is
reported to differ from the latter only by
charcters of the male genitalia. I have not
examined the holotype male of this species,
but would not be surprised to learn that it
is conspecific with P. grandis.
Materials examined.—VIRGINIA.
Northampton Co., Kiptopeke, 2-5 Oct
1987, on flowers of Solidago sempervirens,
W.E. Steiner, J. M. Swearingen, J. M. Hill,
J. J. Marshall (1 3, 1 9; USNM). This site is
at the tip of the Eastern Shore of the Ches-
apeake Bay and is located directly across
the Bay from Norfolk and Virginia Beach.
Distribution.— Widespread throughout
the Pacific basin (Hawaiian Islands, Japan,
Panama, Taiwan); from there disjunct, pre-
sumably by introduction to the Mediterra-
nean (Israel), and Virginia.
Natural history.—In Hawaii, specimens
were collected from salt bush (Atriplex
semibaccata), a plant introduced from Aus-
tralia in the early part of this century. The
specimens collected in Virginia were swept
from goldenrod (Solidago sempervirens).
Other species of the genus are predators,
perhaps scavengers, on various molluscs,
especially small mussels, and barnacles.
Remarks.— Mathis (1988) has suggested
elsewhere that the lower Chesapeake Bay
and ajoining land are the recipients of many
introductions because of the large volume
of commerce on these waterways. The lower
Chesapeake Bay, the principal waterway as-
sociated with this portion of Virginia, is one
of the busiest in the world, serving the large
metropolitan areas of Baltimore and Wash-
ington, and housing a principal base for the
U.S. Navy on the East Coast. With a high
level of traffic, the possibility of an intro-
duction, as “‘extra baggage,” is much great-
er.
Other factors contributing to the intro-
duction of species concern their natural his-
tory (Lewin 1987). All species reported re-
cently, including this one, frequent the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
littoral zone where they are scavengers on
molluscs or feed on algae and/or seaweed
that has accumulated on the shore. These
food sources are plentiful in the littoral zones
of Virginia. Furthermore, these habitats are
not particularly rich in insect species, which
may contribute to the success of an insect
invader. The widespread distribution of P.
grandis has undoubtedly further abetted the
possibilities of invasion but has probably
had little or no influence on the stability of
the invasion afterwards.
Clearly much remains to be studied about
the immigration of insects and of this species
in particular, whose natural history and
ecology are largely unknown. Also, basic
survey work and general collecting are valu-
able tools in and prerequisites for the de-
tection of exotic organisms.
ACKNOWLEDGMENTS
The collecting efforts of Warren E. Stei-
ner, particularly in littoral habitats, are ap-
preciated. I also thank Allen L. Norrbom
for critically reviewing a draft of this paper.
The illustrations were rendered by George
Venable. I thank them all for their contri-
butions to this paper.
LITERATURE CITED
Canzoneri, S. 1986. Nuovi dati sugli Ephydridae
(Diptera) della Sierra Leone. Part I]. Accad. Naz.
Lincei 383(260): 67-75.
Cogan, B. H. 1984. Family Ephydridae, pp. 126-176.
In Soos, A., ed., Catalogue of the Diptera of the
Palaearctic Region. Hung. Acad. Sci., Budapest.
Cogan, B. H. and W. W. Wirth. 1977. Family Ephy-
dridae, pp. 321-339. Jn Delfinado, M. D. and
Hardy, D. E., eds., A Catalogue of the Diptera of
the Oriental Region, Vol. Il: Suborder Cyclor-
rhapha (excluding Division Aschiza). Univ. Press
Hawaii, Honolulu.
Cresson, E. T., Jr. 1925. Studies on the Dipterous
family Ephydridae, excluding the North and South
American faunas. Trans. Amer. Entomol. Soc. 51:
227-258.
1945. A systematic annotated arrangement
of the genera and species of the Indoaustralian
Ephydridae (Diptera), I: The subfamily Psilopi-
nae. Trans. Amer. Entomol. Soc. 71: 47-75.
VOLUME 90, NUMBER 3
Lewin, R. 1987. Ecological invasions offer opportu-
nities. Sci. 238: 752-753.
Mathis, W. N. 1986. Studies of Psilopinae (Diptera:
Ephydridae), I: A revision of the shore fly genus
Placopsidella Kertész. Smithson. Cont. Zool. 430:
1-30.
. 1988. The first report of the genus Procanace
Hendel from North America, with the description
of a new species (Diptera: Canacidae). Proc. Ento-
mol. Soc. Wash. 90(3): 329-333.
Mathis, W. N.and W.E. Steiner. 1986. Anadventive
species of Brachydeutera Loew in North America
337
(Diptera: Ephydridae). J. New York Entomol. Soc.
94(1): 56-61.
Miyagi, I. 1977. Ephydridae (Insecta: Diptera). Jn
Fauna Japonica. | 13 pp. Keigaju Pub. Co., Tokyo.
Tenorio, J. A. 1980. Family Ephydridae, pp. 251-
351. In Hardy, D. E. and M. D. Delfinado, eds.,
Diptera: Cyclorrhapha III, Insects of Hawaii, Vol.
13: 251-351. Univ. Press Hawai. Honolulu.
Wirth, W. W. et al. 1987. Ephydridae [chapter 98],
pp. 1027-1047. In McAlpine, J. F., ed., Manual
of Nearctic Diptera, Vol. 2. Res. Branch, Agric.
Canada. Monograph 28.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 338-342
A NEW SPECIES OF HAEMATOMYZUS (MALLOPHAGA:
HAEMATOMYZIDAE) OFF THE BUSH PIG, POTAMOCHOERUS
PORCUS, FROM ETHIOPIA, WITH COMMENTS
ON LICE FOUND ON PIGS
K. C. EMERSON AND ROGER D. PRICE
(KCE) 560 Boulder Drive, Sanibel, Florida 33957; (RDP) Department of Entomology,
University of Minnesota, St. Paul, Minnesota 55108.
Abstract. —A new species, Haematomyzus porci, is described and illustrated for speci-
mens taken off the Bush Pig, Potamochoerus porcus, in Ethiopia; this represents the third
species described in this genus. The known data of lice found on the eight species of pigs
are also summarized.
Key Words:
Only two species of Haematomyzus Pia-
get have been described to date: H. ele-
phantis Piaget from the African Elephant,
Loxodonta africana (Blumenbach), and the
Asiatic Elephant, Elephas maximus Lin-
naeus, and H. hopkinsi Clay from the Wart
Hog, Phacochoerus aethiopicus (Pallas), in
Kenya and Uganda. When Piaget (1869) de-
scribed H. elephantis, he established Hae-
matomyzus as a genus of Anoplura. Sub-
sequently, Enderlein (1904) erected the
family Haematomyzidae in the order An-
oplura for this species. Ferris (1931) pub-
lished results of a detailed study of the anat-
omy of H. elephantis, concluding that it had
‘biting mouth-parts” and should be in the
order Mallophaga; he, therefore, described
the suborder Rhynchophthirina for the
family Haematomyzidae. External mor-
phology features found only in the genus
Haematomyzus are illustrated in Figs. 1-3;
these include the prolonged mouthparts
which are mandibulate and have no piercing
mechanism, the thorax which is different in
shape from any known in the Anoplura or
other Mallophaga genera, and the legs which
are distinct from those found on any Ano-
Mallophaga, Haematomyzus, Haematopinus, Suidae
plura or other Mallophaga species. Clay
(1963) examined four hypotheses as to the
host for the original ancestral stock which
resulted in the genus Haematomyzus and
could not conclude which was most prob-
able, the Wart Hog or the African Elephant.
We recently received a series of Hae-
matomyzus collected from the Bush Pig in
Ethiopia; these lice represent a third species
of this group of Mallophaga. All measure-
ments are in millimeters. Details common
to Haematomyzus species will not be re-
peated here, since Ferris (1931) and Clay
(1963) have adequately treated them.
Haematomyzus porci Emerson and Price,
New SPECIES
Figs. 1-3
Type host: Potamochoerus porcus (Lin-
naeus) [Artiodactyla: Suidae].
Male.—As in Fig. 1. Each abdominal
pleuron II with 2 dorsal medial setae con-
sisting of short fine seta and longer thicker
seta; dorsal posterior seta on pleuron III 0.03
mm long. Dorsal abdomen with 5 clear cir-
cular areas on each side encompassing me-
dium seta in each; additional minute setae
VOLUME 90, NUMBER 3
Fig. 1.
lateral and medial to these. Abdominal ster-
na II-VIJ each with 2 unmodified setae and
following number of stout modified setae:
MS 12s I 20:1V=V.. 2a2 Vi 205 Vill. 7.
Genitalia (Fig. 3) with long prominent
rounded basal plate, closely apposed para-
meres with truncate apices each bearing 2
setae, and with spinose sac and associated
structures as shown. Dimensions: head
339
Haematomyzus porci, male, dorsal (left}-ventral (right) view. Scale line = 0.1 mm.
width, 0.43 mm; head length (including pro-
boscis), 0.65 mm; proboscis length, 0.38
mm; pterothorax width, 0.68 mm; abdo-
men width (at V), 1.02 mm; total length,
1.95 mm; genitalia length, 0.65 mm; geni-
talia width, 0.11 mm.
Female.—Head and thorax as for male
(Fig. 1). Abdomen as in Fig. 2. Width of 8
median tergal plates, respectively, from an-
340
Figs. 2-3.
dorsal view. Scale lines = 0.1 mm.
terior to posterior: 0.31—0.33 mm, 0.39-0.40
mm, 0.35-0.36 mm, 0.46 mm, 0.47-0.49
mm, 0.44-0.47 mm, 0.36-0.41 mm, and
0.28-0.31 mm. Abdominal sterna II-VII
each with 2 unmodified setae and following
number of stout modified setae: II, 12-13;
III, 20-22; IV, 25; V, 22-26; VI, 22; VII,
11. Dimensions: head width, 0.40-0.43 mm;
head length (including proboscis), 0.66-0.7 |
mm; proboscis length, 0.42-0.43 mm;
pterothorax width, 0.68 mm; abdomen
width (at V), 1.37—1.40 mm; total length,
2.30-2.34 mm.
Discussion. — Clay (1963) provides excel-
lent features for distinguishing H. hopkinsi
from H. elephantis. Haematomyzus porci is
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Haematomyzus porci. 2, Female abdomen, dorsal (left)}-ventral (right) view. 3, Male genitalia,
morphologically closer to the former, dif-
fering from H. elephantis in many of the
same ways as does H. hopkinsi. The prin-
cipal feature separating H. porci from H.
hopkinsi is the presence of 5 pairs of clear
circular areas, each surrounding a medium
seta, on the male dorsal abdomen as op-
posed to only 4 pairs of such areas on H.
hopkinsi (lacking the medioanterior pair)
and none on H. elephantis. The separation
of H. porciis further supported by the dorsal
posterior seta on male pleuron III only a
third the length of that of H. hopkinsi, the
female with a tendency for 1-5 fewer mod-
ified setae on each of abdominal sternites
II-VII, and smaller female dimensions for
VOLUME 90, NUMBER 3
the pterothorax width (0.68 mm vs 0.73-
0.76 mm) and total length (2.30-2.34 mm
vs 2.42-2.52 mm).
Material examined.—Holotype ¢ from
Potamochoerus porcus collected on July 16,
1964, near Addis Ababa, Ethiopia, by C. T.
O’Connor; in the collection of Oklahoma
State University, Stillwater. Paratypes: 7 9,
same data as holotype; distributed among
Oklahoma State University, University of
Minnesota, Field Museum of Natural His-
tory, and U.S. National Museum of Natural
History.
LicE FOUND ON Pics
Most mammalogists place the eight
species of living pigs into five genera (No-
wak and Paradiso 1983). The known data
of the lice on these eight species of pigs are
summarized here.
The genus Sus contains S. scrofa Lin-
naeus, the Wild Boar, found originally in
Europe, parts of Asia, and North Africa, but
now widely introduced by man. The louse
found on this host is Haematopinus apri
Goureau, a species of sucking louse in the
order Anoplura which has been reviewed
by Ferris (1951). The domestic pig is also
included in S. scrofa; domestic pigs, how-
ever, have a different anopluran louse, Hae-
matopinus suis (Linnaeus). We have not
been able to obtain lice off S. salvanius
(Hodgson), the Pigmy Hog, S. barbatus
Muller, the Bearded Pig, or S. verrucosus
Muller and Schlegel, the Javan Pig, but we
suspect that the lice of these south Asian
pigs will also be species of Haematopinus.
The remaining four genera of pigs each
contains a single species: Babyrousa baby-
russa (Linnaeus), the Babirusa, is found in
the Celebes and probably has a species of
sucking louse; Phacochoerus aethiopicus, the
Wart Hog, is found in Africa and has Hae-
matomyzus hopkinsi, a species of chewing
louse, as well as a sucking louse, Haema-
topinus phacochoeri Enderlein:; Potamo-
choerus porcus, the Bush Pig, is found in
341
Africa and Madagascar and has both Hae-
matomyzus porci, the new species of chew-
ing louse described here, and Haematopinus
latus Neumann, a sucking louse; Hylocho-
erus meinertzhageni Thomas, the Giant
Forest Hog, is found in Liberia, southwest-
ern Ethiopia, and northern Tanzania and
has the sucking louse Haematopinus mei-
nertzhageni Werneck, described by Wer-
neck (1952) subsequent to the review by
Ferris (1951).
A key to separate these three species of
Haematomyzus is as follows:
1. Male dorsal abdomen lacking clear circular
areas each surrounding medium seta; female
dorsal last abdominal segment with both short
and medium setae elephantis
— Male dorsal abdomen with 4-5 pairs of clear
circular areas each surrounding medium seta
(Fig. 1); female dorsal last abdominal segment
with all setae similar, of medium length (Fig.
2) ie EAP dT ATNaA Specie ytee
Male dorsal abdomen with 4 pairs of clear cir-
cular areas each surrounding medium seta and
posterior seta of pleurite III over 0.08 long;
female with pterothorax width over 0.72
te
: ; ee See hopkinsi
— Male dorsal abdomen with 5 pairs of clear cir-
cular areas each surrounding medium seta and
posterior seta of pleurite ITI less than 0.06 long;
female with pterothorax width less than 0.70
porci
ACKNOWLEDGMENTS
We thank J. S. Ashe for the loan of critical
specimens from the Field Museum of Nat-
ural History, Chicago. This research was
partially supported by Project No. Min-17-
015, Minnesota Agricultural Experiment
Station, St. Paul, Minnesota 55108, and has
been assigned Paper No. 15,703, Scientific
Journal Series.
LITERATURE CITED
Clay, T. 1963. A new species of Haematomyzus Pia-
get (Phthiraptera, Insecta). Proc. Zool. Soc. Lon-
don 141: 153-161.
Enderlein, G. 1904. Lausestudien. I. Uber die Mor-
phologie, Klassifikation und systematische Stel-
lung der Anopluren nebst Bemerkungen zur Sys-
342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tematik der Insektenordnungen. Zool. Anz. 28: | Nowak, R. M. and J. L. Paradiso. 1983. Walker’s
121-147. Mammals of the World, Vol. II. xxv + 569-1362.
Ferris, G. F. 1931. The louse of elephants. Haema- Piaget, E. 1869. Description d’un parasite de l’ele-
tomyzus elephantis Piaget (Mallophaga: Haema- phant. Tijdschr. Ent. 12: 249-254.
tomyzidae). Parasitology 23: 112-127. Werneck, F. L. 1952. Contribuicao ao conhecimento
1951. The Sucking Lice. Mem. Pacific Coast dos Anoplura. II. Rev. Brasil. Biol. 12: 201-210.
Entomol. Soc., Vol. I. San Francisco. 320 pp.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 343-355
BIOLOGY AND IMMATURE STAGES OF THE RHODODENDRON
GALL MIDGE, CLINODIPLOSIS RHODODENDRI FELT
(DIPTERA: CECIDOMYIIDAE)
DONALD R. SPECKER AND WARREN T. JOHNSON
(DRS) Research Support Specialist, Department of Entomology, Cornell University;
(WTJ) Professor, Department of Entomology, Cornell University.
Abstract. —Clinodiplosis rhododendri (Felt) is an occasionally serious pest of Rhododen-
dron catawbiense hybrids grown in nurseries in the Northeast. Biological and ecological
information is presented for the first time. Immature stages are described and illustrated.
The larva was examined via scanning electron microscopy. The species is multivoltine,
passes through 3 instars, and overwinters both in the soil and in flower buds as mature
larvae. Larval aestivation and natural enemies are reported. Damage is described.
Key Words:
Cecidomyiids, or gall midges, are a di-
verse group biologically and ecologically and
are represented by more than 1200 species
in North America. Although a few members
of the family, such as the Hessian fly and
alfalfa gall midge, are serious agricultural
pests and well studied, most species have
received very little attention.
White (1933) described damage caused
by an unidentified gall midge on rhododen-
dron. It was reported to cause considerable
deformation of the leaves of Rhododendron
ponticum L., R. maximum L., and hybrid
varieties in nurseries. White also reported
finding it on wild R. maximum in the Po-
cono mountains of Pennsylvania. Felt (1939)
described the gall midge as a new species,
Giardomyia rhododendri. Gagné (1973)
placed G. rhododendri in the genus Clino-
diplosis Kieffer. By our petition, the Ento-
mological Society of America has approved
the common name rhododendron gall
midge. The gall midge is recognized in var-
ious books dealing with insect pests of or-
namental plants, including those by Pirone
(1978) and Westcott (1973). It is also rec-
Insecta, rhododendron gall midge, Clinodiplosis rhododendri, insect pest
ognized as a pest in several books on rho-
dodendron culture such as those by Leach
(1961), Bowers (1960), and Van Veen
(1969).
MATERIALS AND METHODS
Site |1—a commercial nursery, East
Hampton, NY; Site 2—a commercial nurs-
ery, Melville, NY. Emergence of adults from
overwintering larvae was monitored with
the aid of traps. Traps consisted of standard
6-in.-diameter white plastic flowerpots
coated on the inside with Tanglefoot®. Traps
were placed in an inverted position on the
soil beneath branches of plants showing
damage from the previous year. Twenty-
eight traps were placed throughout two fields
at Site 2 on 10 May 1980. Traps were
checked by removing them and recording
the number and sex of all gall midges pres-
ent. Traps were checked nine times at in-
tervals of 3-5 days from 19 May to 15 June.
Above-ground plant parts were examined
with a 10x hand lens to ascertain ovipo-
sition sites. Eggs were collected with a
moistened camel-hair brush and placed in
344
70% ethanol. Plant parts bearing eggs were
removed and brought to the laboratory for
further observation. Foliage bearing evi-
dence of damage was removed and exam-
ined for larvae by carefully pulling it apart.
The number of larvae, their size, color, lo-
cation in the damaged tissue, and activity
were noted. Larvae were transferred to 70%
ethanol or a KAAD mixture with a moist-
ened camel-hair brush. Infested foliage was
removed and placed in paper bags for trans-
port to the laboratory. Petioles of 10 in-
fested leaves were sprayed with Tangle-
foot® on 26 June 1979 at Site 1 to determine
if mature larvae crawl down the plant to
reach the soil for pupation. These petioles
were checked on 2 July.
A random sample of 21 flower buds of
the cultivar ‘Nova Zembla’, and 32 flower
buds of ‘Roseum Elegans’, were taken on
10 April 1980 at Site 2 to ascertain whether
mature larvae had overwintered in them,
and whether there was a varietal preference.
The buds were dissected on 17 April 1980
and the number and condition of mature
larvae in them noted. Soil samples were tak-
en on 10 May 1981 at Site 2 to determine
the stage of development of overwintering
larvae and their distribution within the soil.
Fifteen saples were taken with an Oakfield ®
soil sampler from beneath plants that were
heavily damaged the previous summer.
Samples were divided into five depth inter-
vals: 1-2 cm, 2-4 cm, 4-6 cm, 6-8 cm, and
8-10 cm.
Eggs on plant parts were observed with a
dissecting microscope to determine color
changes associated with development; lo-
cation, orientation, and distribution on the
host plant; and to aid in illustration. Eggs
were placed directly in Euparal® on micro-
scope slides for detailed observation and
measurement. This was advantageous in that
the minute, translucent eggs were difficult
to recover from ethanol. An ocular microm-
eter was used to determine egg dimensions.
A small camel-hair brush was used to ma-
nipulate and transfer eggs.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Larvae were easily removed from leaves
by placing infested foliage in plastic bags;
in a few hours the larvae vacated the leaves.
Larvae crawling on the inner surface of the
bags were then collected with a moistened
camel-hair brush.
Larvae killed in KAAD were transferred
to 95% ethanol after 2 hour. Some larvae
were dehydrated in an alcohol series, cleared
in a terpineol solution (Hood 1953), and
mounted in Canada balsam on microscope
slides for detailed observation. Other larvae
were treated in cold 5% NaOH for 24 hours,
dehydrated, cleared as above, and mounted
in Canada balsam. Larval dimensions, in-
cluding head capsuls widths, were deter-
mined with the aid ofan ocular micrometer.
Larvae to be viewed with the scanning
electron microscope were dehydrated in an
alcohol series, washed twice in pure acetone,
then transferred to small plastic capsules that
had been perforated with a No. 00 insect
pin. The larvae were critical-point dried us-
ing pure acetone. Each capsule was then
gently opened and inverted over a mounting
stub covered with double-sided tape. Lar-
vae were positioned on the tape with the aid
ofa small camel-hair brush and then coated
with 200-250 4 of gold-palladium. All mi-
crographs were made at 10 kV with an AM-
RAY 1000® SEM.
An estimate of the time required for ma-
ture larvae to complete development and
emerge as adults was made by placing 20
mature larvae in a Syracuse watch glass con-
taining moist sand and checking it daily for
adult emergence. To more precisely deter-
mine the length of this period, mature lar-
vae were placed in Thunderbird® No. 111
clear plastic cups of about 30 ml capacity
containing sand moistened with distilled
water. A single larva was placed in each cup
after being examined for ectoparasites or
any readily visible pathological condition.
Each cup was covered with a plastic lid and
placed in a rearing room (25°C, 75% RH).
Cups were checked each morning and night
starting on the ninth day. Forty cups were
VOLUME 90, NUMBER 3
Table 1.
345
Dimensions (mm) of immature stages of C. rhododendri.
Larva
Egg Ist Instar 2nd Instar 3rd Instar Pupa
an Cements eae L Ww L W if W E W
Mean 0.29 0.08 0.64 0.17 1.32 0.35 DeD, 0.56 1.80 0.62
Maximum 0.30 0.09 1.17 0.47 2.10 0.51 3.03 0.76 2a 0.76
Minimum 0.28 0.06 0.27 0.07 0.75 0.17 Ie Pe 0.46 1S 0.53
SD 0.01 0.01 0.27 0.08 0.29 0.08 0.29 0.08 0.21 0.08
n 25 24 47 37 7
set up on 25 August 1979 with larvae col-
lected at Site | the day before. Fifty larvae
collected on 26 August 1980 at Site 2 were
set up on 2 September 1980.
Cocoons were recovered from thoroughly
dried soil samples with the aid of a No. 18
USA Standard Testing Sieve® (1.0 mm
opening, Tyler equivalent = 16 mesh). Soil
aggregates not passing the sieve were care-
fully broken apart under a dissecting mi-
croscope.
The longevity of adults was estimated by
recording mortality for individuals emerg-
ing in the plastic cups. Sex was determined
after death. Fecundity was measured by dis-
secting the ovaries of newly emerged gall
midges and counting the number of eggs per
female with the aid of a dissecting micro-
scope. Cecidomyiids emerge from the pupal
stage with their full complement of ova ma-
tured and ready for oviposition (Barnes
1946). Pairs of newly emerged adults were
confined on swollen vegetative buds in the
laboratory with the aid of small plastic cages
(10 cm tall, 8 cm diam.). Unfortunately, all
adults placed in cages on plants died within
a few hours without having oviposited.
DESCRIPTION OF IMMATURE STAGES
Felt (1939) published a description of the
adult. Immature stages have not been pre-
viously described.
Egg ellipsoidal, nearly round in cross sec-
tion, anterior end slightly wider than pos-
terior; ca. 3 times longer than wide, 0.29 x
0.08 mm (Fig. 9; Table 1). Chorion smooth,
shiny, transparent and unsculptured; sticky
and resilient. Vitelline membrane visible,
especially at posterior end. Newly deposited
egg nearly colorless. As egg develops it be-
comes reddish-orange. A large, diffuse, red-
dish area appears slightly posteriad of cen-
ter. Vitelline membrane becomes constricted
at both ends of egg. Incipient segmentation
evident as white patches appear along sides.
A dark-red eyespot appears in the anterior
eighth of egg during late stages of devel-
opment.
Larva spindle-shaped with a distinct but
minute head; ca. 3° times longer than wide,
length varies from ca. 0.5 mm in first instar
to ca. 2mm at maturity (Figs. 1, 3, 4; Table
1). Larva creamy white throughout most of
its development, becoming orange-yellow
at maturity. Head capsule weakly sclero-
tized, with prognathous mouthparts (Fig. 5).
Mouthparts consist of mandibles, maxillae,
a labrum and labium (Fig. 6). Mandibles
inserted internally. Papillae present on
maxillae and labium; pits present on la-
brum. Antennae two-segmented, conical,
conspicuous. Head separated from thorax
by well-developed cervix. Dark-red eyespot
present within cervix in all instars. Spiracles
present on prothoracic segment and abdom-
inal segments 1-8 of third-instar larva; those
on prothoracic segment and eighth abdom-
inal segment prominent. Spatula present on
prothoracic venter of third-instar larva. In-
tegument rugose (Figs. 3, 4). Rows of spi-
nules prominent on venter of meso- and
metathoracic segments and abdominal seg-
346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
EYE-SPOT
SPATULA
PROTHORACIC STERNALIS
SPIRACLE
PLEURAL
SETAE
DORSAL SETA
TERMINAL SETA ANUS
Figs. 1-2.
AS
L2
2
Clinodiplosis rhododenari. |, third-instar larva (dorsal and ventral views). 2, Pupa (A = antenna,
AP = antennal process, AS = apical seta, CL = clypeus, CX! = prothoracic coxa, LB = labella, LE = labrum-
epipharynx, L2 = mesothoracic leg, MP = maxillary palp, PSC = prothoracic spiracular cornicle, TR! =
prothoracic trochanter). Scale bar represents 0.5 mm.
ments 1-9. Barren patches within spinule
rows present on abdominal segments 1-7
(Fig. 7). Spinules surround anus. Six dorsal
and 4 pleural setae present on each thoracic
segment and on abdominal segments 1-7.
Terminal segment with 8 papillae; 4 with
setiform setae, 4 with corniform setae.
Pupa spindle-shaped, exarate (Fig. 2; Ta-
ble 1). Teneral pupa orange-yellow. In ma-
ture pupa (pharate adult) eyes black, wings
dark brown to black, legs and antennae dark
yellow-brown, legs lighter distally. Abdo-
men retains larval color. Sclerotized process
located on base of each antennal sheath.
Apical seta situated mediad and posteriad
to each process. Prothoracic spiracular cor-
nicles prominent, curved outward, tapering.
Abdomen 8-segmented; minute spiracles on
segments 2-6. Prominent setulae present
dorsally along anterior margins of segments
2-8.
Adult yellowish (Fig. 10).
LiFE HIstoRY
Eggs may be deposited on swollen, partly
opened, or fully opened vegetative buds
throughout most of the growing season but
may also be placed on dormant flower buds
in early autumn. Clutch size is variable
(mean = 6.66, SD = 5.51, range = 1-29, n
= 35). Eggs are most often deposited on the
undersurfaces or rolled edges of leaves as
soon as they are free from the bud but before
they have fully separated from each other.
Eggs are not deposited in a specific orien-
tation but may adhere to a leaf along their
lengths, on end, or obliquely, and are often
attached to each other in irregular clumps.
VOLUME 90, NUMBER 3 347
Figs. 3-7. Clinodiplosis rhododendri (mature larva). 3, Anterior segments, ventral view (SS = spatula, SR =
spinule rows). 4, Posterior segments, ventral view (AN = anus). 5, Head. 6, Mouthparts (LM = labrum, MX =
maxilla, LB = labium). 7, Rows of spinules. The barren area within the rows shows the location of an asetose,
anterior ventral papilla. Scale bars for Figs. 3, 4 represent 0.25 mm; all others represent 10.0 um.
348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 8-11. Clinodiplosis rhododendri. 8, Damage to very young leaves of a Rhododendron catawbiense hybrid.
Larvae are present in the rolled leaf margins. Note swollen, necrotic areas and necrotic lesions. 9, Eggs on
undersurface of an expanding leaf. 10, Adult female. 11, Mature pupa with broken cocoon. Scale bars for Figs.
8, 9 represent 10.0 mm, 0.5 mm, respectively; those for Figs. 10, 11 represent 1.0 mm.
A female may deposit several clutches on
the same developing leaf, distribute them
among other leaves in the same whorl, or
deposit them on different plants. Females
readily oviposit on leaves and buds already
bearing clutches from other females. In an
outbreak situation 272 eggs were found on
one swollen vegetative bud. Oviposition was
VOLUME 90, NUMBER 3
NUMBER
iN
Nv
NUMBER
9 10 11 12 13 14 15 16 17 18 19 20 21
DAYS
Figs. 12-13. Clinodiplosis rhododendri. 12, Adult
emergence following placement of mature larvae in
moist sand on day 0. 13, Emergence of C. rhododendri
(white bars) and Platygaster sp. (shaded bars) following
placement of mature larvae in moist sand on day 0.
never witnessed during frequent trips to Site
2 between 10:00 AM and 7:00 PM. Eggs
hatch in ca. 3 days.
Upon hatching a larva crawls into a loose-
ly inrolled margin of a developing leaf and
begins feeding on the abaxial surface; this
feeding usually induces a strong inrolling of
the leaf margin. Most leaf rolls are moist to
wet inside. Larvae that have hatched on
swollen vegetative buds crawl into the bud
to feed on all surfaces of developing leaves.
Larvae infesting flower buds usually feed on
the outer bud scales. Larvae do not move
from leaf to leaf but are restricted to feeding
on the leaf on which they hatched.
349
3° instar
1St instar
2"d instar
PERCENTAGE FREQUENCY
S38 37 ae 38 ao ay a
MICROMETERS
Fig. 14. Clinodiplosis rhododendri. 14, Frequency
distribution of larval head capsule widths. Note over-
lap between second and third instars.
Larvae pass through three instars (Fig. 14).
Head capsule widths of the second and third
instars overlap. The third-instar larva is dis-
tinctive, however, due to the presence of the
spatula.
All larval stages are motile but require a
film of water for sustained locomotion. Lar-
val development takes ca. 7 days. Mature
larvae crawl out of damaged leaves or buds
and either drop to the ground or descend
from the leaves on silken threads (pers. ob-
servation). They burrow into the top 2 cm
of soil and construct a flimsy, silken cocoon
(Fig. 11). Cocoons, which were not found
at depths exceeding 2 cm, are covered with
soil particles that adhere tightly; in sandy
soils cocoons and adhering particles are ap-
proximately ellipsoidal, ca. 2.5 mm in
length. Larvae moult into pupae ca. 7 days
after burrowing into the soil. The mature
pupa breaks out of the larval cocoon, prob-
ably with the aid of its antennal processes,
and wriggles to the soil surface where ec-
dysis to the adult occurs. Laboratory rear-
ings suggest that ca. 11 days elapse from the
time mature larvae drop to the soil until
adults emerge (Fig. 12). A few adults
emerged at 14-15 days.
Adults emerge during the night or early
morning; peak emergence of 27 laboratory-
reared adults occurred between 9:00 PM and
350
8:00 AM. Five adults were observed in the
field between 4:50 AM and 6:20 AM on 25
August 1979 at Site 2, but none were seen
during frequent visits between 10:00 AM
and 7:00 PM. Adults were observed to walk
upon plant surfaces with quick, jerky move-
ments. They frequently flew from plant part
to plant part. The longevity of adults in the
laboratory was ca. 3 days for females and |
day for males. The sex ratio is female-pre-
dominant; 76% of 25 laboratory-reared
adults were female. A binomial proportion
test (Snedecor and Cochran 1967) indicated
that the observed proportions were signifi-
cantly different from a 1:1 ratio (x? = 5.76,
df = 1, P < 0.025). Fecundity was variable
(mean = 34.5 eggs/female, SD = 11.5, range
= 10-51, n = 10).
Throughout this study the only nursery-
grown plants observed to be hosts of C. rho-
dodendri were hybrids of the Catawba rho-
dodendron, R. catawhiense Michaux. All
such hybrids observed in the field served as
host plants. The two most widely grown hy-
brids, ‘Nova Zembla’, and ‘Roseum Ele-
gans’, appeared to be equally susceptible to
attack; a binomial proportion test (Snedecor
and Cochran 1967) indicated that the pro-
portion of damaged flower buds was not
significantly different for these two hybrids
(x? = 0.153, df = 1, P= 0.70). Clinodiplosis
rhododendri was found on wild R. maxi-
mum at Livingston Manor, Sullivan Coun-
ty, NY during the summer of 1982.
SEASONAL HISTORY
On 10 May 1980, plants at Site 2 exhib-
ited slight swelling of vegetative buds.
Adults, eggs, or larvae of C. rhododendri
could not be found on any plant part. On
19 May most vegetative buds were swollen
(a small number had opened) while flower
buds were beginning to swell. A female of
C. rhododendri was found stuck to a swollen
flower bud anda male and female were found
stuck to a partly opened vegetative bud. By
24 May many plants were beginning to flow-
er. Although no evidence of C. rhododendri
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
was found ina random sample of 25 swollen
and opened vegetative buds, an adult was
found in an emergence trap for the first time.
By 27 May most vegetative buds had opened
and leaves were enlarging. Most plants were
in full bloom. The number of adults caught
in emergence traps peaked at this time (n =
6). Leaves damaged by C. rhododendri were
first found on 2 June. These contained
mostly first-instar larvae but also second and
third instars, and in some cases leaves had
already been vacated. The last adults from
the overwintering generation were found in
an emergence trap on 5 June.
Eggs were first observed on 29 May 1980
at Site 2. A single clutch of 16 eggs was
found in a random sample of 40 expanding
vegetative buds. On 2 June, two buds (of
20 randomly sampled) bore small clutches.
Three days later six of 20 buds yielded a
total of 120 eggs. Oviposition during this
period increased logarithmically (For log %
of vegetative buds bearing eggs vs. time, r
= (0.993). The observed increase in the mag-
nitude of the infestation could have been
due to two phenomena: |) a prolonged, in-
creasing emergence of overwintering adults,
or 2) oviposition by a subsequent, larger
generation of gall midges.
On 19 June 1980, a much greater number
of mature, brightly colored larvae were not-
ed than had been found previously. Many
of these larvae were present in severely
damaged leaves that had become necrotic
and dry and which thus provided little or
no food. On 25 June, only mature larvae
occupied the leaves. During this period the
weather was dry, the last rain having oc-
curred on 10 June. On 28 June, 10 damaged
leaf whorls were randomly sampled for lar-
vae; nine of these had 48 brightly colored
mature larvae distributed among them. Eggs
or immature larvae were not found. Heavy
rain fell on 29 June. The next day a random
sample of 10 damaged leaf whorls yielded
no larvae of any age. The larvae had ap-
parently been in aestivation in the leaves
for at least 10 days. Dry necrotic leaves con-
VOLUME 90, NUMBER 3
taining mature larvae were brought into the
laboratory and stored in a paper bag under
dry conditions for 16 days. At the end of
this period larvae were removed from the
dried leaves and observed with a dissecting
microscope. The larvae were motionless and
appeared dead, but with the addition of
water began moving. A number of these lar-
vae developed into adults. It appears that
free water, such as is supplied by rain, is
required for larvae to vacate the leaves. In
the prolonged absence of rain, larvae aes-
tivate.
The early seasonal history of C. rhodo-
dendri is characterized by a close synchrony
with the development of its host plant.
Adults of the overwintering generation be-
gin to emerge as vegetative buds swell, and
peak emergence appears to occur during full
bloom. During this time there is no intra-
specific competition for the many enlarging
vegetative buds as very few insects are pres-
ent. Subsequent generations of the gall midge
more fully utilize the large number of en-
larging buds. Intraspecific competition for
oviposition sites begins at this time and may
continue for the remainder of the season.
R. catawbiense hybrids usually produce two
flushes of growth per season but only the
spring flush is seasonally synchronized: the
second flush may occur at any time between
July and September. The gall midge popu-
lation, which increases greatly during the
spring flush, is faced with a reduced and
temporally discontinuous resource for the
remainder of the season, resulting in a loss
of synchrony between insect and host plant.
For example, on 28 June, 272 eggs were
found on one of the few remaining vegeta-
tive buds present in the nursery.
As autumn approaches, an increasing per-
centage of the larval population does not
pupate following cocoon construction but
instead enters diapause until following sea-
sons. Twelve percent of 50 last-instar larvae
collected at Site 2 on 25 August 1980 did
not pupate. In contrast, all of 20 last instar
larvae collected at Site 1 on 26 June 1979
35
developed into adults within 12 days. On
20 September, almost every plant had com-
pleted growth for the season although a few
enlarging vegetative buds could still be
found. Surprisingly, eggs occurred on these
buds. Based on field observations and lab-
oratory rearings, by this time five genera-
tions of the gall midge may have developed.
On 14 October 1979, mature and penulti-
mate-instar larvae were found although most
of these were in necrotic, dry leaves and
probably were not feeding. An examination
of dormant flower buds revealed small ne-
crotic lesions in the outer tissues of some
buds. A few of these damaged buds con-
tained mature larvae. Oviposition had ap-
parently occurred on these flower buds in
the absence of suitable vegetative buds, and
some of the larvae had matured and dropped
to the soil to overwinter. Mature larvae re-
maining in flower buds would be expected
to overwinter there because of the time of
year and their marked inactivity. A random
sample of 53 flower buds taken on 10 April
the following spring contained a total of 12
larvae, five of which were alive. Seventeen
buds showed evidence of damage. Four of
the living larvae were found in one bud.
NATURAL ENEMIES
No natural enemies of C. rhododendri
have been previously identified. Field and
laboratory observations suggest that natural
enemies are relatively insignificant as con-
trol agents during most of the seasonal his-
tory of the insect. A random sample of 40
mature larvae collected at Site 1 on 25 Au-
gust 1979 yielded only two parasitized in-
dividuals. No other incidences of parasitism
or predation were observed at Site 1. The
effect of natural enemies was greater at Site
2, but appeared to be confined to the late
seasonal history of the insect.
A male pteromalid, possibly of the genus
Callitula Spinola, was reared from an ec-
toparasitic larva found on a mature gall
midge larva collected on 25 August 1979 at
Site 1. The adult parasitoid eclosed ca. 9
352 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
days after the gall midge larva was placed
in moist sand, or ca. | day before the peak
emergence of the gall midge cohort. The
parasitized larva failed to construct a co-
coon or pupate.
Two male parasitic wasps of the genus
Platygaster Latreille were reared from a ma-
ture gall midge larva collected on 25 August
1979 at Site 1. The platygasterids were en-
doparasitic, and pupated in a head-to-head
position within the cuticle of their host. The
parasitized larva successfully constructed a
cocoon. The parasitoids emerged ca. 8 days
after the peak emergence of the gall midge
cohort.
A single encyrtid wasp, probably of the
genus Copidosoma Ratzeburg, emerged from
a sample of infested leaves collected on 26
August 1980 at Site 2. A pteromalid, pos-
sibly of the genus Mesopolobus Westwood,
was also recovered from the same sample.
A species of Platygaster was commonly
recovered from mature larvae collected on
26 August 1980 at Site 2. Ten of 30 gall
midge larvae in one sample and 15 of 50 in
another were parasitized by this wasp. In
all cases one individual pupated per host,
in contrast to the Platygaster sp. reared from
Site 1, where two individuals pupated with-
in one larva.
Sixteen of 25 parasitized larvae success-
fully constructed cocoons, although all died
shortly thereafter. All of the parasitoids
emerged after the peak emergence of the gall
midge cohort (Fig. 13).
The anthocorid bug Orius insidiosus (Say)
was abundant in infested leaves collected
on 26 August 1980 at Site 2. It was probably
feeding on mature larvae present in the
leaves, although feeding was not observed.
No other predators were observed.
INFESTATION AND DAMAGE
The rhododendron gall midge is an oc-
casionally serious, but not commonly en-
countered pest in nurseries on Long Island.
It has not been reported to be a pest in home
plantings.
The distribution of this insect at both
study sites was strongly clumped. At Site 2,
groups of non-infested and very lightly in-
fested plants could be found within 10 m of
heavily infested ones. Five emergence traps
were placed 2-3 m apart in a 72 m? bed of
4-yr old plants on 25 June 1980. Three days
later the number of adults in each trap was
as follows: 4, 0, 1, 0, 156. Barriers such as
woods, fence rows, and non-host plantings
seem to limit infestations. The infestation
at Site 1 was limited to a 2-ha field of stock
plants. A large number of container-grown
plants, separated from the infested field by
ca. 275 m of wooded fence-rows and non-
host plantings, were not infested. It appears
that the active dispersal capability of this
insect is small.
Infestations usually remain at a low level,
but under favorable conditions can increase
dramatically. A 72 m? bed of 2-yr old plants
growing in a screen house at Site 2 was in-
fested so severely that no growth occurred,
resulting in the destruction of the plants by
the nurseryman.
Serious infestations of the rhododendron
gall midge tend to be short-lived. This may
be a function of variables such as rapid turn-
over of nursery-grown plants, use of insec-
ticides, adverse environmental conditions
caused by weather and cultural practices,
and natural enemies. The owner of the nurs-
ery at Site | first noticed gall midge damage
in 1976. The severity of damage increased
each year until 1980, when it was markedly
diminished. Various insecticides were ap-
plied to the plants during this period. The
owner of the nursery at Site 2 first noticed
damage in 1979. Severity increased greatly
in 1980. The insecticide diazinon, applied
as a soil-drench in the spring of 1981, re-
portedly gave good control of the pest.
Whitman Wholesale Nurseries, Suffolk Co.,
NY, sustained damage by this pest on con-
tainer-grown plants in 1978. The insecticide
lindane was applied to the plants three times
during the year. The gall midge could not
be found at this nursery in 1979.
VOLUME 90, NUMBER 3
Various types of damage may result from
the feeding of C. rhododendri depending on
the number of larvae present and the de-
velopment stages of affected parts. Leaves
that are heavily attacked while still in the
bud are severely damaged and normally die
before attaining a length of 6 cm. Leaves
thus affected exhibit swollen, chlorotic areas
and necrotic lesions (Fig. 8). Leaves at-
tacked while free of the bud may reach full
size but become deformed. On these leaves
small (ca. 1 mm?), chlorotic lesions are pro-
duced in affected areas. All infested leaves
develop an inrolling of one or both margins
and become contorted. Damaged areas of
leaves become necrotic with time. Larvae
feeding within flower buds produce necrotic
lesions in the outer bud tissues.
DISCUSSION
Bionomics: The biology of C. rhododendri
is similar to that of the leaf-curling pear
midge, Dasineura pyri (Bouche). Both
species are multivoltine, oviposit on newly
expanding leaves, cause leaf margins to roll
on woody plants, and overwinter as mature
larvae in the soil (Barnes 1935). The ten-
dency toward restriction of emergence, mat-
ing, and oviposition activities of gall midges
to the night and early morning hours has
been reported by many workers, including
Weigel and Sanford (1920), Walter (1941),
and Coutin and Harris (1968). This restric-
tion of adult activities is advantageous for
at least two reasons: 1) it minimizes the
disruptive effects of wind, the primary dis-
persal agent for the higher gall midges (Ma-
maev 1968), and 2) it minimizes the poten-
tial for desiccation, especially during
emergence. The short adult life of C. rho-
dodendri is common to other cecidomyiids
(Weigel and Sanford 1920, Walter 1941,
Azab et al. 1965, Brewer 1971, Ranasinghe
1977). Another aspect of the biology of C.
rhododendri that is shared by other ceci-
domyiids is a female-predominant sex ratio
(Sasscer and Borden 1919, Barnes 1935,
Walter 1941, Redfern 1975, Ranasinghe
353
1977). Clinodiplosis rhododendri appears to
be native to the northeastern United States
as it has been found on wild R. maximum
in both the Catskill mountains of New York
and the Pocono mountains of Pennsylvania.
It has not been recorded to occur outside of
this area on a wild host, nor has it been
recorded from any other wild host. It has
been found on cultivated hosts in nurseries
from Maryland north to Massachusetts. R.
maximum, R. catawbiense, R. ponticum, R.
caucasicum Pallas, R. arboreum Smith, and
R. smirnowli Trautvetter are all reported to
be parent species of R. catawhbiense hybrids.
C. rhododendri has been reported to occur
on R. ponticum in nurseries (White 1933).
Phenology: The seasonal history of C.
rhododendri is similar to that of other ceci-
domyiids. The emergence of adults in spring
coincides closely with the development of
their host plant. Synchrony between gall
midges and host plants has been reported
by Bishop (1954), Gable et al. (1959), and
Coutin and Harris (1968). Coutin (1964)
reported that for many species the behavior
of ovipositing females and the duration of
Oviposition are governed by the floral bi-
ology of the host. The seasonal buildup of
large populations of C. rhododendri is com-
mon to other gall midges as well. Barnes
(1940) reported that over 2000 individuals
of Diarthronomyia chrysanthemi Ahlberg
may develop in one chrysanthemum plant.
The number of larvae of Contarinia pseu-
dotsugae Condrashoff developing in a single
Douglas-fir shoot 1s usually over 2000 (Con-
drashoff 1962). The aestivation of mature
larvae of C. rhododendri during dry weather
also resembles the behavior of other gall
midges. Bishop (1954) reported that mature
larvae of Dasineura gentneri Pritchard re-
main in clover florets for variable intervals,
dependent on weather conditions. During
damp or rainy weather there is a tendency
for the larvae to vacate the florets shortly
after reaching maturity. During dry weather
some larvae may remain in the florets for a
week or more. Barnes (1952b) showed that
354
fully fed larvae of Contarinia tritici (Kirby)
need moist conditions in order to descend
from damaged wheat plants to the soil. Dry
weather during the maturation of larvae re-
sults in an assemblage of larvae in damaged
plants that vacate the plants en masse when
rain comes. These same conditions are re-
quired by C. rhododendri. Diapause of ma-
ture larvae during autumn and winter is a
common aspect in the seasonal history of
many gall midge species, including C. rho-
dodendri. Barnes (1935) noted that in the
multivoltine species D. pyri, varying pro-
portions of the larvae of the second, third,
and fourth generations entered diapause un-
til the following spring. He also demonstrat-
ed that larvae of C. tritici could spend two
winters in diapause, those of Sitodiplosis
mosellana (Gehin) three winters (Barnes
1943), and later (Barnes 1952a) showed that
S. mosellana may diapause for up to 12
years. The late season utilization of flower
buds as an alternate resource by C. rhodo-
dendri appears to be unrecorded for other
cecidomyiids.
Natural enemies: Species of Platygaster
have been recorded from other cecido-
myiids by a number of workers (Brewer
1971, Coutin and Harris 1968, Houseweart
and Brewer 1972, and Ranasinghe 1977).
Most platygasterids are parasitoids of ceci-
domyiid larvae (Borror et al. 1976). Both
Coutin and Harris (1968) and Barnes (1935)
observed predation of gall midge larvae by
Anthocoridae.
ACKNOWLEDGMENTS
We thank Richard Vrana, Springs Nurs-
ery, L.I., and Kenneth Graham, Graham’s
Nursery, L.I., for granting permission to use
their nurseries as study sites. Raymond J.
Gagné (Systematic Entomology Labora-
tory, USDA) identified C. rhododendri and
Norman F. Johnson (Ohio State University)
its parasitoids. Donald Steinkraus (Cornell
University) was of valuable assistance in
preparing the plates and in microscopy.
Robert G. Mower (Cornell University) pro-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vided information on cultivated host plants.
We especially are indebted to Donald B.
Zepp (Rhone-Poulene, Inc.) for his constant
assistance in carrying out this study. We
thank E. Richard Hoebeke (Cornell Uni-
versity) for reviewing the manuscript.
LITERATURE CITED
Azab, A. K., M. F. S. Tawfik, and I. I. Ismail. 1965.
Morphology and biology of the aphidophagous
midge, Phenobremia aphidivora Rubsaamen. Bull.
Soc. Entomol. Egypte. 49: 25-45.
Barnes, H. F. 1935. Studies of fluctuations in insect
populations. V. The leaf-curling pear midge, Dasy-
neura pyri(Cecidomyiidae). J. Anim. Ecol. 4: 244-
253.
1940. The biology of the chrysanthemum
midge in England. Ann. Appl. Biol. 27: 71-91.
1943. Studies of fluctuations in insect pop-
ulations. X. Prolonged larval life and delayed sub-
sequent emergence of the adult gall midge. J. Anim.
Ecol. 12: 137-138.
. 1946. Gall Midges of Economic Importance.
Vol. II. Gall Midges of Fodder Crops. Crosby
Lockwood and Sons, Ltd., London. 160 pp.
1952a. Studies of fluctuations in insect pop-
ulations. XII. Further evidence of prolonged larval
life in the wheat-blossom midges. Ann. Appl. Biol.
39: 370-373.
1952b. Studies of fluctuations in insect pop-
ulations. XIII. An improved method of ascertain-
ing the correct date to sample when assessing lar-
val infestations of the wheat-blossom midges. Ann.
Appl. Biol. 39: 374-378.
Bishop, G. W. 1954. Life history and habits of a new
seed midge, Dasyneura gentneri Pritchard. J. Econ.
Entomol. 47: 141-147.
Borror, D. J., D. M. DeLong, and C. A. Triplehorn.
1976. An Introduction to the Study of Insects.
4th ed. Holt, Rinehart, and Winston, New York.
852 pp.
Bowers, C. G. 1960. Rhododendrons and Azaleas:
Their Origins, Cultivation, and Development. 2nd
ed. The Macmillan Co., New York. 525 pp.
Brewer, J. W. 1971. Biology of the pinyon stunt needle
midge. Ann. Entomol. Soc. Am. 64: 1099-1102.
Condrashoff, S. F. 1962. Bionomics of three closely
related species of Contarinia Rond. (Diptera: Ceci-
domyiidae) from Douglas-fir needles. Can. Ento-
mol. 94: 376-394.
Coutin, R. 1964. Le comportement de ponte chez
plusieurs Cécidomyies en relation avec l’état de
développement chez la plante-hdte des organes re-
cherchés pour l’oviposition. Rev. Zool. Agric. Appl.
63: 45-55.
Coutin, R. and K. M. Harris.
1968. The taxonomy,
VOLUME 90, NUMBER 3
distribution, biology and economic importance of
the millet grain midge, Geromyia penniseti (Felt),
gen. n. (Dipt., Cecidomyiidae). Bull. Entomol. Res.
59: 259-273.
Felt, E. P. 1939. A new gall midge on rhododendron.
J. N.Y. Entomol. Soc. 47: 41-42.
Gable, C. H., W. A. Baker, and L. C. Woodruff. 1959.
The sorghum midge, with suggestions for control.
U.S. Dep. Agric. Farmers’ Bull. 1566. 9 pp.
Gagné, R. J. 1973. A generic synopsis of the nearctic
Cecidomyiidae (Diptera: Cecidomyiidae: Ceci-
domyiinae). Ann. Entomol. Soc. Am. 66: 857-
889.
Hood, J. D. 1953. Microscopical Whole-Mounts of
Insects. 4th mimeographed ed. Entomology stock-
room, Cornell Univ., Ithaca, N.Y. 53 pp.
Houseweart, M. W. and J. W. Brewer. 1972. Biology
of a pinyon spindle gall midge (Diptera: Cecido-
myiidae). Ann. Entomol. Soc. Am. 65: 331-336.
Leach, D.G. 1961. Rhododendrons of the World and
How to Grow Them. Charles Scribner’s Sons, New
York. 544 pp.
Mamaevy, B. M. 1968. Evolution of Gall Forming
Insects— Gall Midges. The British Library, Lend-
ing Division, West Yorkshire, England. 317 pp.
Translated from the Russian by A. Crozy, 1975.
Pirone, P. P. 1978. Diseases and Pests of Ornamental
Plants. 5th ed. John Wiley and Sons, New York.
566 pp.
355
Ranasinghe, M. A. S. K. 1977. Bionomics of a sub-
terranean gall midge (Diptera: Cecidomyiidae) from
Artemesia ludoviciana. Great Basin Nat. 37: 429-
442.
Redfern, M. 1975. The life history and morphology
of the early stages of the yew gall midge Taxomyia
taxi (Inchbald) (Diptera: Cecidomyiidae). J. Nat.
Hist. 9: 513-533.
Sasscer, E. R. and A. D. Borden. 1919. The rose
midge. U.S. Dep. Agric. Bull. 778. 8 pp.
Snedecor, G. W.and W.G. Cochran. 1967. Statistical
Methods, 6th ed. Iowa State Univ., Ames.
Van Veen, T. 1969. Rhododendrons in America.
Sweeney, Krist, and Dimm, Portland, OR. 176
pp.
Walter, E. V. 1941. The biology and control of the
sorghum midge. U.S. Dep. Agric. Tech. Bull. 778.
26 pp.
Weigel, C. A. and H. L. Sanford. 1920. Chrysanthe-
mum midge. U.S. Dep. Agric. Bull. 833. 25 pp.
Westcott, C. 1973. The Gardener’s Bug Book. 4th
ed. Doubleday and Co., Inc., Garden City, N.Y.
689 pp.
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PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 356-368
MALE FORETARSAL VARIATION IN LYCAENIDAE AND
RIODINIDAE, AND THE SYSTEMATIC PLACEMENT
OF STYX INFERNALIS (LEPIDOPTERA)
RoBERT K. ROBBINS
Department of Entomology, NHB STOP 127 Smithsonian Institution, Washington,
DC 20560 USA.
Abstract. —I describe and illustrate male forelegs, particularly the tarsus, of Lycaenidae,
Riodinidae, and Styx infernalis, and note seven characters: (1) whether the male foreleg
is used for walking, (2) presence or absence of pretarsal claws, (3) number of tarsomeres,
(4) distribution of scales on the tarsus, (5) whether scales lie flat on the tarsus, (6) presence
or absence of tarsal spines (A-Type trichoid sensilla) and (7) distribution of B-Type trichoid
sensilla on the tarsus when present. Previous descriptions of the male foretarsus and
pretarsus of riodinids and Styx were inaccurate, including a reported pretarsal claw in
Styx. Characters of the Styx male foreleg are either shared with riodinids or are unique.
This conclusion supports Harvey’s classification of Styx as a riodinid, and is inconsistent
with Ehrlich’s and Scott’s phylogenies to the butterfly families.
Key Words:
Morphology of the male foretarsus and
pretarsus in the butterfly, Styx infernalis
Staudinger, has been disputed. Ehrlich
(1958) distinguished the male foretarsus and
pretarsus of Styx from those of Riodinidae.
He reported that male Styx, a monobasic
genus from the eastern Andes of Peru, has
a segmented foretarsus whereas riodinids
have a fused male foretarsus. He noted a
pretarsal claw (and possibly a second one
that had been broken in the one foreleg that
he had available for study) in Styx but not
in riodinids (with rare exceptions). Forbes
(1960:138), on the other hand, stated, “‘It is
said that the South American Styx infernalis
has developed legs in both sexes; in fact. . .
the true male ... has the proper reduced
legs” of a riodinid. Harvey (1987) partially
resolved this controversy by reporting that
male riodinids may have a segmented tarsus
and that male Styx lacks pretarsal claws.
The systematic position of Styx has like-
leg characters, butterfly classification, Lycaenidae, Riodinidae, Styx
wise engendered controversy. Ehrlich (1958)
erected a monobasic subfamily for Styx of
rank equal to the Riodinidae (his Riodini-
nae) and Lycaenidae (his Lycaeninae) on the
basis of differences in the male forelegs and
some other characters. This classification has
been followed by many subsequent authors
(e.g. Common and Waterhouse 1982, Ack-
ery 1984). Further, using some of Ehrlich’s
results, Scott (1985) proposed a cladogram
in which Styx is the first taxon to split off
from a lineage leading to the Lycaenidae
and Riodinidae.
The classification and phylogenies of Ehr-
lich (1958) and Scott (1985) have been ques-
tioned. Harvey (1987) placed Styx in the
Riodinidae on the basis of two shared, de-
rived character states: female foretarsal
trichoid sensilla clustered centrally, not lat-
erally, and lack of apophyses posteriores on
the female genitalia. I (Robbins 1988) re-
ported that Styx and Riodinidae (with the
VOLUME 90, NUMBER 3
exception of the Old World genus Laxita
Butler) are the only butterflies that share the
loss of a cluster of trichoid sensilla on the
dorsal posterior inner face of the male fore-
leg trochanter, which supports Harvey’s
placement of Styx in the Riodinidae. Scott
and Wright (1988) placed Styx in the Rio-
dinidae (their Riodininae) “for the mo-
ment.” I (Robbins 1987b) reported that
Scott (1985) did not analyze the distribution
of male lycaenid and riodinid foretarsal
character states parsimoniously, casting
doubt on his phylogeny.
The purposes of this paper are (1) to de-
scribe and illustrate the male foretarsal
morphology of the Lycaenidae (sensu Eliot
1973), Riodinidae (sensu Stichel 1910-
1911), and Styx, (2) to resolve the differing
results of Ehrlich, Forbes, and Harvey, (3)
to detail the qualitative differences in male
foreleg morphology of Riodinidae and Ly-
caenidae, and (4) to use this information to
assess the classifications and phylogenies of
Ehrlich, Scott, and Harvey.
The scanning electron microscope (SEM)
provides an opportunity to study leg struc-
tures in a detail not available to many pre-
vious authors, and I have made extensive
use of it. I describe morphology of the male
lycaenid and riodinid foretarsi by citing pre-
vious results and adding my new findings.
I then report the morphology of the male
foretarsus of S. infernalis.
MATERIALS AND METHODS
I used specimens in the collection of the
National Museum of Natural History for
study except for males of Styx infernalis.
Gerardo Lamas (Lima, Peru) loaned me a
male specimen with both forelegs intact, and
Phil Ackery (British Museum of Natural
History) sent me Ehrlich’s dissection of a
male with one unbroken foreleg.
I prepared forelegs for study by briefly
wetting them in 80% ethanol, soaking them
in 10% potassium hydroxide for 24-48
hours, and rinsing them in water or ethanol.
In some cases, I removed some or all scales
357
using fine watchmaker forceps and a brush
with stout bristles. Specimens for the SEM
were soaked for 10 minutes in absolute eth-
anol before being mounted on stubs, which
were coated with carbon and gold.
MALE FORETARSUS AND PRETARSUS
Lycaenidae (sensu Eliot 1973).—The ly-
caenid male foreleg is unique among the
butterflies. With few exceptions (see below),
it lacks pretarsal claws (Bates 1861) (Figs.
2, 3), and its tarsus is fused (Bates 1861)
(Fig. 1), ends in a stubby (Fig. 2) or down-
curved point (Clench 1955, Eliot 1973) (Fig.
3), and possesses A-Type and B-Type trich-
oid sensilla (terminology from Ma and
Schoonhoven 1973). A number of authors
(Sibatani 1974, Higgins 1975, Miller and
Brown 1979) reported a single pretarsal claw,
but I found no structure that fits Snodgrass’s
(1935) description of a pretarsal claw, and
believe that these authors misconstrued the
down-curved point at the tarsal tip reported
by Clench and Eliot in some lycaenids (Fig.
3). Scales cover the dorsal and lateral sur-
faces of the lycaenid male foretarsus, but
not the distal ventral surface, where many
trichoid sensilla occur (Fig. 4). Further,
scales lie relatively flat on the tarsus surface
so that B-Type trichoid sensilla are not cov-
ered by scales (Fig. 5). Lycaenid forelegs are
used for walking (Bates 1861, Ford 1945)
although a number of popular books mis-
takenly state the opposite (Howe 1975, Pyle
1981). Scales usually have longitudinal
ridges with shingled and distally tapered
scutes (Downey and Allyn 1975) (Fig. 6).
These structures are not reported in trichoid
sensilla, and in lieu of better evidence, I use
them to distinguish scales from trichoid
sensilla. I describe below the A-Type and
B-Type trichoid sensilla on lycaenid male
forelegs as well as distinctive setae that also
may be trichoid sensilla.
(1) A-Type trichoid sensilla (spines).—
Spines are stout trichoid sensilla, some-
times called “bristles,” that have fluted sides
and occur primarily on the ventral surface
3
Figs. 1-7. Male lycaenid foretarsus. 1. “Theritas” augustinula Strand, dorsum, scales removed to show lack
of segmentation. Scale line 600 microns. 2. Calycopis cecrops Fabricius, lateral view, stubby tip, spine (A-Type
trichoid sensillum, labelled s), “‘macrotrichion” (m), B-Type trichoid sensilla (t), and lack of pretarsal claws.
Scale line 60 microns. 3. Lycaena editha Mead, lateral view, down-curved point at tip. Scale line 75 microns.
4. “Theritas” theocritus Fabricius, ventro-lateral aspect, no scales on ventral surface, empty sockets of removed
scales on lateral surface. Scale line 200 microns. 5. C. cecrops, lateral view, spine (A-Type trichoid sensillum,
s), “macrotrichion” (m), and B-Type trichoid sensillum (t), which extends beyond scales that lie flat on the
tarsus. Scale line 176 microns. 6. “7.” theocritus, scale showing scutes of longitudinal ridges, base of scale to
right. Scale line 3 microns. 7. “7.” Theocritus, fine structure spines. Scale line 12 microns.
VOLUME 90, NUMBER 3
of the lycaenid foretarsus (Bates 1861) (Figs.
1-5, 7, 8) and other legs. Histology of spines
in nymphalids (Eltringham 1933) and neu-
rophysiology from nymphalids and pierids
indicate that they are mechanoreceptors
(Morita et al. 1957, Ma and Schoonhoven
1973). Those on female butterfly foretarsi
are associated with clustered B-Type trich-
oid sensilla, and are apparently used to
abrade leaves (reviewed in Chew and Rob-
bins 1984), but may also be mechanorecep-
tors. Superficially similar spines occur on
the ventral abdomen of some lycaenids and
riodinids (Clench 1955, Inoue and Kawazoe
1966, Eliot 1973, Harvey 1987) and on but-
terfly antennae (Myers 1968, Grula and
Taylor 1980), where they are presumed to
be mechanoreceptors (Odendaal et al. 1985).
Spines on legs and antennae always seem to
occur in association with B-Type trichoid
sensilla, as they do on the venter of the fe-
male abdomen in the lycaenid, Curetis Hiib-
ner (Robbins unpubl.).
(2) B-Type trichoid sensilla.— These sen-
silla are scattered over the dorsal, lateral,
and ventral sides of the lycaenid male fore-
tarsus (Figs. 1, 2, 4, 5). At magnifications
below about 1000 times, B-Type trichoid
sensilla on male lycaenid foretarsi appear to
be smooth-walled (Robbins 1987a) (Fig. 2),
but at magnifications above about 8000
times, they have a variable fine structure
(Figs. 13, 14). It is unclear in the absence
of histological and neurophysiological data
whether there is one kind of B-Type trichoid
sensillum on male lycaenid foretarsi with
variable surface structure or if there are sev-
eral morphologically and functionally dif-
ferent types. Another fine structure, in which
there are ringed indentations (Fig. 15), oc-
curs on female lycaenid foretarsi, but I have
not found them on male lycaenid foretarsi.
Undoubtedly, histological and neurophysi-
ological techniques will be needed to estab-
lish homologies.
B-Type trichoid sensilla on butterfly legs
are chemosensory and mechanosensory.
Lycaenids extend their proboscis when their
Si)
tarsi are exposed to water or sugar water
(Anderson 1932), and Hodgson (1958)
showed that trichoid sensilla respond neu-
rophysiologically to sodium chloride, su-
crose, and tactile stimulation. It has not been
specifically demonstrated, however, that the
B-Type trichoid sensilla on the lycaenid male
foreleg are chemosensory, although this is
a reasonable inference. Similar sensitivity
among B-Type trichoid sensilla to water,
sugars, sodium chloride, and tactile stimu-
lation has been shown for a variety of but-
terfliles (reviewed in Fox 1967), and El-
tringham (1933) presented a histological
description of these sensilla. Ma and
Schoonhoven (1973) described the histol-
ogy of a clustered B-Type sensillum on the
female pierid foretarsus, and demonstrated
that these clusters are sensitive to plant sec-
ondary compounds, water, sodium chlo-
ride, and tactile stimulation.
(3) ““Macrotrichia.” — Clench (1955) not-
ed that a pair of long setae, which he termed
““macrotrichia,” occurs on the dorsal sur-
face of the male lycaenid foretarsus just bas-
al to the tip (Figs. 2, 5). Their surface struc-
ture at higher magnifications is distinctive
(Figs. 9, 10), and these markings are often
more pronounced towards their distal end.
Whether these setae are scales or sensilla is
currently unknown. Superficially similar
structures are found in most other butter-
flies, even male Ithomiinae (Nymphalidae)
that have the tibia and tarsus fused into a
short segment (Fig. 11). In Phoebis Hiibner
(Pieridae), the analagous structures have
scale-like longitudinal ridges and scutes (Fig.
12), similar to some piliform scales (Brown
& Miller 1983). Kuznetsov (1967) illustrat-
ed similar structures in a sphingid and arc-
tid, termed them “setae” or “ungal bris-
tles,’’ and reported that their number varies
in Lepidoptera from 2-10 and is “of definite
taxonomic importance.”
Some male lycaenids have a five-seg-
mented foretarsus and pretarsal claws, which
apparently have evolved at least four times
in the Theclinae and perhaps once in the
360 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 8-14. Male foretarsi. Socketed base of sensillum to left in figures of their fine structure. 8. Aflides
halesus Cramer (Lycaenidae), fine structure spine (A-Type trichoid sensillum). Scale line 6 microns. 9. C. cecrops
(Lycaenidae), fine structure “macrotrichion.” Scale line 2 microns. 10. A. halesus (Lycaenidae), fine structure
“‘macrotrichion.” Scale line 2 microns. 11. Pagyris cymothoe Hewitson (Nymphalidae: Ithomiinae), apex of
fused foretarsus and tibia with ‘“‘macrotrichia.” Scale line 33 microns. 12. Phoebis sennae Linnaeus (Pieridae),
fine structure ‘macrotrichion.”’ Scale line 2.5 microns. 13. A. halesus (Lycaenidae), surface structure B-Type
trichoid sensillum. Scale line 2 microns. 14. C. cecrops (Lycaenidae), surface structure B-Type trichoid sensillum.
Scale line 2 microns.
VOLUME 90, NUMBER 3
Liphyrinae + Miletinae (Eliot 1973). Eliot
(1973) suggested that if the “genes” for a
segmented foretarsus and clawed pretarsus
were on the Y chromosome, then crossing
over with the X chromosome might account
for its repeated evolution in males. How-
ever, the segmented male foretarsus lacks
the clusters of B-Type trichoid sensilla found
on female foretarsi, at least in Theclopsis
Godman and Salvin (Figs. 16, 17), casting
some doubt on this hypothesis.
Riodinidae (sensu Stichel 1910-1911).—
Unlike the male lycaenid foretarsus, the male
riodinid foretarsus is not used for walking
(Bates 1861, Ford 1945), and is covered
dorsally, laterally, and ventrally with elon-
gate scales (the so-called “‘brush foot”’) (Figs.
18-21). It has from 1 to 4 tarsomeres (God-
man and Salvin 1879-1886, Scott 1985)
(Figs. 18-21), contrary to Ehrlich’s findings,
but segment partitions are sometimes in-
complete (Godman and Salvin 1879-1886,
Powell 1975), making a count of the number
of tarsomeres somewhat arbitrary. The male
riodinid pretarsus is like that of lycaenids
in that it lacks claws (Figs. 18-21). Godman
and Salvin (1879-1886) reported male pre-
tarsal claws in Apodemia nais Edwards, but
the three specimens that I examined lacked
them. I have not seen any riodinid with
male foreleg pretarsal claws, but remnant
ones may occur in some species.
The occurrence of spines (A-Type trich-
oid sensilla) on male foretarsi differs be-
tween lycaenids and riodinids. Male riodin-
ids lack foreleg spines (Bates 1861) (Figs.
18-20) although I examined one male of
Emesis with a spine on one foretibia and
none on the other. A striking exception is
males of Sarota Westwood (Harvey 1987),
which have many foretarsal spines (Figs. 21,
22).
The distribution of B-Type trichoid sen-
silla differs markedly between lycaenid and
riodinid male foretarsi. B-Type trichoid
sensilla are absent from the male riodinid
foretarsus except at the distal end of the
tarsus where “macrotrichia” also occur.
These sensilla are difficult to discern be-
361
cause they do not extend beyond the scales
covering the tarsus as they do in lycaenids.
I usually find one or two “macrotrichia” at
the tarsal tip (Figs. 23, 25-27), and some-
times a few B-Type trichoid sensilla (Figs.
23, 24). Many setae on the tarsus resemble
B-Type trichoid sensilla at lower magnifi-
cations, but at higher magnifications appear
to be scales with longitudinal ridges and
scutes (Fig. 28). It is not known whether the
B-Type trichoid sensilla on the riodinid
foretarsus are neurophysiologically active.
All riodinid male foretarsi may have
B-Type trichoid sensilla and “macrotrich-
ia” at the tip even though I could not find
them in all preparations. Some scales have
to be removed to see them, and in these
cases, I may have removed them with the
scales. However, the restricted occurrence
of B-Type trichoid sensilla on the distal tar-
sus where they are intermixed with elongate
scales, whether or not these sensilla are pres-
ent in all riodinid species, is quite different
from their distribution in lycaenids, as de-
scribed above. Further, even though male
Sarota have spines similar to lycaenids, their
B-Type trichoid sensilla are distributed ac-
cording to the riodinid pattern (Figs. 22-
24).
Styx infernalis.— The structure of the male
S. infernalis foretarsus and pretarsus is sim-
ilar to those of riodinids in some respects,
and is unique in others. The Styx male fore-
leg pretarsus lacks claws (Harvey 1987) (Figs.
29, 30), and in that respect, is the same as
riodinids and most lycaenids. There is a
lightly sclerotized structure at the tip of the
tarsus that may be a remnant of the pretar-
sus (Figs. 29, 30), perhaps homologous with
the arolium. No riodinids have such a struc-
ture. The distal edge of the lightly sclero-
tized structure at the tip of the tarsus ap-
pears dark under a light microscope, and
may account for Ehrlich’s report of a single
pretarsal claw.
The male riodinid foretarsus segmenta-
tion is unusual. One specimen (the Ehrlich
dissection) has two tarsomeres with an in-
dication of two other partitions while the
362 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 15-23. Foretarsi. 15. female Liptena libyassa Hewitson (Lycaenidae), fine structure clustered B-Type
trichoid sensillum. Scale line 10 microns. 16. male Theclopsis murex Druce (Lycaenidae), ventral aspect, seg-
mented, but no clustered B-Type trichoid sensilla. Scale line 100 microns. 17. female 7. murex Druce (Lycaeni-
dae), ventral aspect with lateral cluster of B-Type trichoid sensilla (circled). Scale line 100 microns. 18. male
Melanis pixe Boisduval (Riodinidae), lateral aspect with most elongate scales on lateral and ventral surfaces
removed (sockets visible), four tarsomeres. Scale line 380 microns. 19. male Emesis mandana Cramer (Riodin-
idae), lateral view with most elongate scales on lateral surface removed (sockets visible), three tarsomeres. Scale
line 430 microns. 20. male Stalachtis magdalenae Westwood (Riodinidae), lateral aspect with most elongate
VOLUME 90, NUMBER 3
second specimen has four complete tarso-
meres on both forelegs (Figs. 31, 32). Powell
(1975) reported similar intraspecific varia-
tion in the number of tarsomeres in the male
foretarsus of the riodinid 4. nais. Segmen-
tation in Styx appears to be more complete
and to allow more intersegmental move-
ment than in riodinids, but like riodinids
and lycaenids, it has less than 5 tarsomeres.
The distribution of setae on the male fore-
tarsus of S. infernalis is similar to that of
riodinids. It has a sparse covering of elon-
gate scales on all sides (Figs. 31, 32) with
two “‘macrotrichia” on the dorsal end of the
tarsus (Figs. 31, 32). One foreleg had no
spines (A-Type trichoid sensilla) on the tar-
sus (Fig. 31), whereas the other foreleg from
the same specimen had a spine on the sec-
ond tarsomere (Fig. 33). B-Type trichoid
sensilla occur on the last tarsomere, pri-
marily on the ventral surface (Figs. 34, 35)
except that one leg had one trichoid sensil-
lum on the third tarsomere. The fine struc-
ture of these trichoid sensilla is superficially
more similar to that of “macrotrichia” (Fig.
36) than to that of lycaenid and riodinid
B-Type trichoid sensilla.
It is not known whether male S. infernalis
use their forelegs for walking. Ehrlich (1958)
noted that its male foretarsus is less than
half the size of the pterothoracic legs and is
doubtfully functional. Because other but-
terflies with “brush feet’ do not use their
forelegs for walking, I agree that the same
is probably true for Styx.
DISCUSSION
The male foreleg of Lycaenidae differs
from that of Riodinidae in a number of
characters. (1) The foreleg is used for walk-
ing in lycaenids, but not in riodinids. (2)
The distal, ventral surface of the tarsus lacks
scales in lycaenids but not in riodinids. (3)
Scales lie flat on the tarsus in lycaenids but
not in riodinids. (4) The tarsus is wholly
fused in lycaenids (with some five-seg-
mented exceptions) whereas it is partially
or wholly fused in riodinids with 1-4 tar-
someres. (5) The lycaenid tarsus possesses
spines (A-Type trichoid sensilla) over much
of the ventral surface whereas the riodinid
tarsus does not, with the notable exception
of Sarota. (6) The lycaenid tarsus has scat-
tered B-Type trichoid sensilla that protrude
beyond the scales while the riodinid tarsus
has B-Type trichoid sensilla restricted to the
tip where they are intermixed with elongate
scales. (7) The foreleg is more than half the
length of the pterothoracic legs in lycaenids
and less than half this length in riodinids
(Ehrlich 1958). (8) The coxa does not extend
beyond its articulation with the trochanter,
or if it does, it is arched upwards in lycae-
nids whereas it extends beyond the tro-
chanter in a blunt process without being
arched upwards in riodinids (Robbins 1988).
(9) The trochanter has a cluster of small
trichoid sensilla on its anterior inner surface
whereas this cluster is lacking in riodinids
(Robbins 1988).
The male foreleg of Styx is structurally
that of a riodinid. It shares the riodinid
character state for each of the 9 characters
above except that data for the first character
are lacking. Ehrlich (1958) was mistaken in
finding similarity between the male foreleg
of Styx and that of male lycaenids with a
five-segmented foretarsus and clawed pre-
tarsus.
The Styx male foreleg differs from that of
riodinids in several characters. (1) The light-
ly sclerotized structure at the tarsus tip is
—
scales removed (sockets visible) to show lack of segmentation. Scale line 200 microns. 21. male Sarota dematria
Westwood (Riodinidae), lateral view with most elongate scales removed (sockets visible), spine (A-Type trichoid
sensillum, s) on ventral surface. Scale line 430 microns. 22. male S. dematria (Riodinidae), detail of ventral
surface showing spines with fluted walls. Scale line 60 microns. 23. male S. dematria (Riodinidae), tip showing
position of “macrotrichion” (m) and B-Type trichoid sensillum (t). Scale line 60 microns.
364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 24-30. Male foretarsi. 24. Sarota dematria (Riodinidae), fine structure B-Type trichoid sensillum at
apex. Scale line 3 microns. 25. S. dematria (Riodinidae), fine structure “‘macrotrichion” at apex. Scale line 2
microns. 26. Hades noctula Staudinger (Riodinidae), apex showing position of “‘macrotrichion” (m) and piliform
scale (c). Scale line 87 microns. 27. H. noctula (Riodinidae), fine structure ‘“‘macrotrichion” at apex. Scale line
2.5 microns. 28. M. pixe (Riodinidae), fine structure piliform scale at apex. Scale line 3 microns. 29. Styx
infernalis, lateral aspect, lightly sclerotized structure at tip, no pretarsal claws. Scale line 60 microns. 30. S.
infernalis, posterior aspect of 29. Scale line 50 microns.
VOLUME 90, NUMBER 3
Figs. 31-36. Male foretarsus of Styx infernalis. 31. Lateral view, four tarsomeres, “macrotrichia.” Scale line
400 microns. 32. Ventral view, four tarsomeres, “macrotrichion” (m). Scale line 380 microns. 33. Spine (A-
Type trichoid sensillum) on ventral surface of second tarsomere. Scale line 20 microns. 34. B-Type trichoid
sensillum (t) on ventral surface last tarsomere. Scale line 38 microns. 35. Fine structure B-Type trichoid sensillum
on ventral surface of last tarsomere. Note longitudinal lines instead of broken latitudinal lines in Lycaenidae
and Riodinidae. Scale line 2.5 microns. 36. Fine structure ““macrotrichion” dorsal surface of last tarsomere.
Scale line 2.5 microns.
unique to Styx. (2) The tarsomere partitions
in Styx appear to allow greater interseg-
mental movement. (3) The forecoxa ex-
tends a shorter distance beyond the tro-
chanter in Styx than in most riodinids
(Robbins 1988). (4) The trochanter of Styx
lacks a cluster of trichoid sensilla on the
dorsal, outer posterior surface whereas it 1s
366
present in riodinids (Robbins 1988). These
four differences show that Forbes’s descrip-
tion of the male Stvx foreleg as typically
riodinid was partially incorrect.
SYSTEMATIC POSITION OF STYX
Ehrlich (1958) placed Styx in its own
subfamily (Styginae) of rank equal to the
Riodinidae (his Riodininae) and Lycaeni-
dae (his Lycaeninae). His evidence was (1)
the occurrence of two recurrent veins in the
forewing cell, (2) the form of the labial scler-
ite, (3) a strongly convex mesothoracic an-
episternum, and (4) the morphology of the
male foreleg. Ehrlich remarked that the first
two character states are unique to Styx and
the third is also “unique but close to the
riodinines.” These unique character states
by themselves provide no evidence on the
systematic placement of Styx. Either they
evolved on the lineage leading to Styx only
or they are part of a transformation series
for which information from other charac-
ters is necessary to show the order of trans-
formation.
Ehrlich noted that the male foreleg of Styx
is close to lycaenids whose males have a
five-segmented foretarsus with pretarsal
claws. This comparison was incorrect be-
cause Styx lacks pretarsal claws and has less
than five tarsomeres. Further, lycaenids,
whose males possess a five-segmented tar-
sus and clawed pretarsus, have the lycaenid
pattern of scales, spines, and B-Type trich-
oid sensilla (Fig. 16), not the one shared by
riodinids and Styx. Thus, Ehrlich’s evi-
dence did not justify giving the Styginae rank
equal to the Lycaenidae and Riodinidae.
Scott (1985) proposed that Styx branched
from the lineage that then evolved into the
Lycaenidae and Riodinidae. His evidence
is that Styx possesses (1) a large anepister-
num that became “‘slightly smaller” in the
remainder of the lineage, and (2) eyes that
are not notched at the antennae whereas the
remainder of the lycaenids and riodinids
have notched eyes.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
While Ehrlich noted that the shape of the
mesothoracic anepisternum is unique to
Styx (but close to the riodinids), Scott con-
sidered its size to be a “primitive” character
state, but did not indicate his evidence for
this hypothesis. He did not measure the an-
episternum nor indicate whether its size is
allometrically correlated with body size.
Further, the mesothoracic anepisternum is
not a separate sclerite in Libytheidae and
Pieridae, and is present in only some species
of Papilionidae and Nymphalidae (Ehrlich
1958). Since this sclerite may be present or
absent in potential outgroups and its size
unmeasured when present, there is no evi-
dence that a large anepisternum is “primi-
tive” on the lineage leading to the Lycaeni-
dae and Riodinidae.
Scott’s statement that the Lycaenidae and
Riodinidae (exclusive of Styx) have eyes
notched at the antennae is inaccurate. Al-
though it is true for many Lycaenidae and
Riodinidae, some (Hades Westwood, Eu-
selasia Hiibner) have the same arrangement
of compound eyes and antennae as Styx. In
short, Scott provides no evidence for his
systematic placement of Styx.
Harvey (1987) put Styx in the Riodini-
dae. He characterized the Riodinidae as
those butterflies (1) with B-Type trichoid
sensilla on the female foretarsus clustered
centrally and (2) lacking apophyses posteri-
ores on the female genitalia. Styx and rio-
dinids (with the exception of Laxita) also
share the loss ofa trichoid sensillum cluster,
which is present in all other butterfly fam-
ilies, on the male foretrochanter (Robbins
1988).
My results in this paper are consistent
with Harvey’s placement of Styx in the
Riodinidae. The differences between Styx
and other riodinids in male foretarsus struc-
ture are character states that are unique to
Styx (such as the slightly sclerotized struc-
ture at the tip of the tarsus) and that provide
no evidence on its systematic position. On
the other hand, riodinids and Styx share 8
VOLUME 90, NUMBER 3
character states of the male forelegs. These
results are consistent with Harvey’s classi-
fication of Styx as a riodinid.
ACKNOWLEDGMENTS
I am grateful to Gerardo Lamas for loan-
ing me the only male specimen of Styx in
the Museo de Historia Natural (Lima) and
for allowing me to examine its forelegs with
the SEM. I thank Phil Ackery for loaning
me Ehrlich’s dissection of a male Styx and
for trying to find another male in the British
Museum collection. For sharing informa-
tion on butterfly phylogeny with me over
the years, I thank Don Harvey. For allowing
me to read an advance copy of his co-au-
thored paper on butterfly phylogeny, I thank
James Scott. For reading and commenting
upon the manuscript, I am grateful to Don
Harvey, Gerardo Lamas, James Scott, and
Adrienne Venables. I particularly thank an
anonymous referee whose comments great-
ly improved the manuscript. For technical
help with the SEM, I thank Brian Kahn,
Susann Braden, and Heidi Wolff.
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PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 369-372
OBSERVATIONS ON THE TRUE BUGS EMESA TENERRIMA, A
POSSIBLE SPIDER MIMIC, AND GHILIANELLA BORINCANA
(HEMIPTERA: REDUVIIDAE: EMESINAE) FROM PUERTO RICO
JorGce A. SANTIAGO-BLAY AND JENARO MALDONADO-CAPRILES
(JASB) Biology Department, University of Puerto Rico, Rio Piedras, Puerto Rico 00931
(Current address, Department of Entomological Sciences, 218 Wellman Hall, University
of California, Berkeley, California 94720); (JMC) Urbanizacion Aponte 61 1, Cayey, Puerto
Rico 00633.
Abstract.—Emesa tenerrima and Ghilianella borincana were observed in a semi-ever-
green forest of northern Puerto Rico during day hours for eleven months. Emesa tenerrima
lives on the webs of the pholcid spider Modisimus signatus with which it shares the
elongated body form and banded coloration pattern of its legs. Mild disturbances induce
mantid-like displays; stronger disturbances produce easy movement over the web or flight
escape. Live pholcids were not observed coinhabiting the webs occupied by E. tenerrima.
Ghilianella borincana is not a web inhabitant, shows no interest in offered pholcids but
seems to eat spiders. It intermingles with dark vegetation debris and, with the aid of
catalepsis, is highly cryptic. Disturbed individuals drop from the substrate.
Key Words:
Hinton (1973, 1976) noted that while nu-
merous spiders mimic insects, no insects
indisputably mimic spiders. Only recently
has the occurrence of spider mimicry been
confirmed (Greene et al. 1987, Mather and
Roitberg 1987, Whitman et al., in press).
Emesine true bugs (Hemiptera: Reduviidae:
Emesinae) are frequently associated with
spiders (Gagné and Howarth 1975, Hick-
man 1971, Wygodzinsky 1966).
One of us (JASB) noticed a ‘“‘stick-like
spider” inhabiting a pholcid web in the
understory of a semi-evergreen forest of
northern Puerto Rico. Upon manipulation
with forceps, the arthropod disclosed its true
identity: Emesa tenerrima, an emesine (Fig.
1). Several specimens, including a second
emesine, Ghilianella borincana (Fig. 2), were
observed in dark vegetation debris sus-
pended in the forest. This paper documents
some biological features of Emesa tenerri-
Emesinae, Pholcidae, spider mimicry, mimicry
ma and Ghilianella borincana, and suggests
their possible biological significance.
METHODS
The study site, a semi-evergreen, sub-
tropical moist, premontane forest (Ewel and
Whitmore 1973, Holdridge 1982), is locat-
ed in a limestone hill, a short walk off road
2, km. 21.4 (near junction road 165), Toa
Baja. Observations were done, usually dur-
ing the mornings of 12 different days, from
September 1983 to August 1984. Each ob-
servation period lasted a minimum of ten
minutes. Observations were made on dif-
ferent individuals of each species. Six ob-
servation periods were devoted to E. tener-
rima, seven to G. borincana. Detection of
G. borincana was facilitated by placing a
white background behind possible residence
places. Simple manipulations were per-
formed to observe the reactions of the eme-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
370
. \
\
\
\
\
Fig. 1. mesa tenerrima. Pin length about 40 mm.
Fig. 2. Ghilianella borincana. Pin length about 35 mm.
sines to artificial stimuli. Voucher speci-
mens are deposited in the Entomology
Museum, Agricultural Experiment Station,
Univ. Puerto Rico, Mayagiiez Campus (at
Rio Piedras), Puerto Rico.
RESULTS
Emesa tenerrima (Dohrn, 1860)
(Fig. 1)
This winged, silvery grayish-black eme-
sinine was always collected on webs of Mo-
disimus signatus (Banks) (Pholcidae). Webs
of this spider were found on the branches
of Quararibaea turbinata (Bombacaceae), a
common shrub on the study site. The webs
were located 0.25—2.00 m above ground in
shady areas (50-75% leaf cover). As noted
for other emesines, FE. tenerrima was usually
observed on the web, hanging upside down,
and did not entangle, even if suddenly forced
to move. Emesa tenerrima resembles M.
signatus in its overall elongate body form
VOLUME 90, NUMBER 3
and silver/black transverse banded legs.
However, the body, excluding the legs, of
E. tenerrima is much longer and more ro-
bust than that of M7. signatus. Remnants of
dead spiders, including a Modisimus sig-
natus, were found twice in the emesine in-
habited web, but E. tenerrima and M. sig-
natus were not observed coinhabiting a web.
When warm air was blown directly to-
ward E. tenerrima, or when the branch sup-
porting the web was repeatedly touched, ad-
duction of the meso- and metathoraxic legs
followed. However, the forelegs adducted
only when they were touched, then the fem-
ora and tibiae apposed in a mantid-like pos-
ture. Gentle touch with forceps of the an-
tennae, the meso- and metathoraxic femora,
or tibiae prompted quick walking and es-
cape from the web to surrounding vegeta-
tion. Subsequent manipulation was fol-
lowed by escape to the ground where the
insect was extremely difficult to detect. Es-
cape was also achieved by flight (speed = 3
min 10s,n= 1).
Ghilianella borincana
(Maldonado, 1960)
(Fig. 2)
This wingless, dark-brown metapterinine
lives on vegetation debris in very shady areas
that hangs vertically from Coffea arabica
(Rubiaceae) or Q. turbinata. There it hangs
by four legs, with the fore legs adducted and
apposed, as in EF. tenerrima, but with the
head directed downwards. At times, rem-
nants of other spider species were noticed
on the debris. Ghilianella borincana was ob-
served once on the border of a pitfall trap
with a small unidentified spider held be-
tween a fore femur and tibia.
Ghilianella borincana exhibited catalep-
sis. When a branch with G. borincana was
touched over 30 times, only the antennae
moved, becoming perpendicular to the lon-
gitudinal axis of the body. At times speci-
mens were left unattended for more than 10
min, either on vegetation debris or on a
white piece of paper, and no change in po-
371
sition was noted. However, rubbing a pencil
in front of an emesine placed on a piece of
paper made the insect move away. Escape
occurred by dropping from the substrate.
When living pholcids were brought close to
a G. borincana, no reaction was observed.
DISCUSSION
Reports of emesine/spider associations are
summarized by Wygodzinsky (1966). Ne-
sidiolestes ana and Empicoris rubromacu-
latus have been observed using webs as a
place to feed (Gagné and Howarth 1975,
Hickman 1971), whereas Stenolaemus ed-
wardsii is known to feed on its host spider-
lings (Hickman 1971, Maldonado-Capriles
and van Doesburg 1966).
Twelve emesines are known for Puerto
Rico, but their biology is unknown (Mal-
donado-Capriles 1986). Our data are insuf-
ficient to indicate the nature of the biolog-
ical association between EF. tenerrima and
M. signatus. One possibility is that the re-
semblance of the emesine leg pattern to that
of the spider short-circuits intraspecific
communication (Greene et al. 1987, Mather
and Roitberg 1987, Whitman et al., in press),
thus, possibly representing a case of aggres-
sive mimicry. The fact that we did not see
live pholcids coinhabiting the web with E.
tenerrima questions this hypothesis, unless
spiders actually were eaten at the time of
observation. Another possibility is that E.
tenerrima simply uses the web as a place to
live, at least during the day hours.
Ghilianella borincana, in contrast, may
find protection by remaining on vegetation
debris, at least during the day hours. Re-
semblance to other objects has been re-
ported for a predatory ant-mimic mirid
(Wheeler and Henry 1980), a termite-eating
(McMahan 1983) and medically important
reduviids (Harwood and James 1979), and
an asopine pentatomid nymph that mimics
its chrysomelid prey (Bourdouxhe and Jo-
livet 1981). We have not found comments
concerning similarity of emesines to a sub-
strate habitat.
372
ACKNOWLEDGMENTS
We wish to thank L. Orsak (Scientific
Methods, Inc.; Chico, CA), Dr. S. Medina
(Univ. Puerto Rico, Mayagiiez) and two
anonymous reviewers for their careful re-
view of an earlier version of this paper. Drs.
G. O. Poinar, Jr. and G. M. Thomas (Univ.
California, Berkeley) provided materials and
instructions, respectively, to JASB toward
completion of the photographic process. Dr.
Willis Gertsch (Portal, AZ), Dr. M. J. Vélez,
Jr. and Mr. R. Rodriguez (Univ. Puerto
Rico, Rio Piedras) loaned specimens of M.
signatus.
LITERATURE CITED
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platys cincta (Coleoptere Chrysomelidae) in Sen-
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
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vida. Instituto Americano de Cooperacion para la
Agricultura. San José, Costa Rica. 216 pp.
Maldonado-Capriles, J. 1960. Assassin bugs of the
genus Ghilianella in the Americas (Hemiptera,
Reduviidae, Emesine). Proc. U.S. Natl. Mus. 112:
393-450.
1986. The Reduviidae of Puerto Rico: An
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Maldonado-Capriles, J. and P. H. van Doesburg, Jr.
1966. On some Emesinae from Dutch Guiana
(Surinam), with a new subspecies. Proc. Entomol.
Soc. Wash. 68: 325-329.
Mather, M. H. and B. D. Roitberg. 1987. A sheep in
wolf's clothing: Tephritid flies mimic spider pred-
ators. Science 236: 308-310.
McMahan, E. A. 1983. Adaptations, feeding prefer-
ences, and biometrics of a termite-baiting assasin
bug (Hemiptera: Reduviidae). Ann. Entomol. Soc.
Amer. 76: 483-486.
Wheeler, A. G., Jr. and T. J. Henry. 1980. Seasonal
history and host plants of the ant mimic Barber-
iella formicoides with description of the Sth instar
(Hemiptera: Heteroptera: Miridae). Proc. Ento-
mol. Soc. Wash. 82: 269-275.
Whitman, D. W., L. Orsak, and E. Greene. Spider
mimicry in fruit flies (Diptera: Tephritidae): Fur-
ther experiments on jumping spider predation
avoidance by Zoonemata vittigera (Coquillet). Ann.
Entomol. Soc. Amer. (In press.)
Wygodzinsky, P. 1966. A monograph on the Eme-
sinae (Reduviidae, Hemiptera). Bull. Amer. Mus.
Nat. Hist. 133: 1-614.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 373-392
DISTRIBUTION AND HABITAT COMPARISONS FOR CARABODES
COLLECTED FROM CONIFER BRANCHES WITH DESCRIPTIONS
OF BREVIS BANKS AND HIGGINSIN. SP.
(ACARI: ORIBATIDA: CARABODIDAE)
R. MARCEL REEVES
Department of Entomology, University of New Hampshire, Durham, New Hampshire
03824.
Abstract. —Seven species of Carabodes (brevis Banks, dendroetus Reeves, higginsi n.
sp., willmanni Bernini, granulatus Banks, niger Banks and labrynthicus (Michael)) were
collected from spruce and fir branches using the caustic soda wash technique. A rede-
scription of C. brevis, a description of C. Aigginsi and distribution within North America
for each species is included. Comparisons of branch samples with leaf litter/rotten wood
samples from the forest floor clearly show that C. brevis and C. dendroetus prefer arboreal
habitats. Additional evidence from literature and other collections indicate that C. gran-
ulatus, C. labrynthicus and C. niger may be arboreal depending on the presence of lichens,
moss or fungi but C. willmanni may be arboreal only where lichens are present. Habitat
preferences for C. higginsi could not be determined except that most specimens were
collected in association with conifers.
Key Words:
Oribatid mites, Carabodes brevis, C. dendroetus, C. higginsi, C. willmanni,
C. granulatus, C. niger, C. labrynthicus
Oribatid mites are difficult to remove from
arboreal habitats, especially evergreens with
needles, with the conventional method of
searching bark, branches or foliage. Often
lichens, moss or various other organic ac-
cumulations adhere to branch surfaces ob-
scuring vision. Fungi and lichens on the
trunks of trees offer an easier habitat from
which to collect. In Europe C. labrynthicus
(Michael) has sometimes been collected
from dry plants on rock surfaces (Seyd and
Seaward 1984) and Bellido (1979) found all
stages of C. willmanni Bernini burrowing
into lichens. Travé (1963) found C. /abryn-
thicus and minusculus Berlese more com-
mon in moss and lichens on rock and tree
surfaces than in soil. The presence of certain
species of Carabodes on the bark surfaces
of trees or in lichens, fungi, moss, etc. at-
tached to these surfaces is thus known, yet
the extent of their distribution into the for-
est canopy has not been documented.
The caustic soda wash technique that re-
sulted in the mite collections reported in
this paper is used for second instar larval
surveys of spruce budworm (Choristoneura
fumiferana (Clemens)) (Miller and Mc-
Dougal 1968). Briefly this technique re-
quires clipping branches from the mid-
crowns of host trees and immersing the
branches in a 1% hot caustic soda solution
for several hours to free larvae from their
hibernacula. Much of the organic debris
(needles, bark, etc.) is removed through
sieving before the eventual sample is filtered
ina Biichner funnel. Large numbers of mites
on filter papers from branch samples sub-
jected to the above process were brought to
374
my attention. I proceeded to remove mites
as the samples were processed and time per-
mitted. These samples were clipped on Sep-
tember 3/4 or 9/11, 1983 from red spruce
(Picea rubens Sarg.) and balsam fir (Abies
balsamea (L.) Mill.) at 10 different locations
all within approximately 13 miles of Errol,
Coos Co., New Hampshire. The amount of
branch surface and whether lichens, fungi
or moss were present on the branches is
unknown.
Among the mites collected from the
branch samples were varying numbers of
seven species of Carabodes. | had rarely en-
countered the two most abundant species,
C. brevis Banks and C. dendroetus Reeves,
in leaf litter/rotten wood collections indi-
cating that they may prefer an arboreal hab-
itat. Thus a comparison of species present
in both habitats was appropriate even though
differences in sampling methods did not al-
low statistical comparisons. Descriptions of
C. brevis Banks and C. higginsi n. sp. are
included as well as North American distri-
butional records and habitat preferences for
all seven species based on the literature, the
author’s collection, and the Canadian Na-
tional Collection (CNC), Biosystematics
Research Centre, Ottawa.
Terminology and abbreviations are those
developed by F. Grandjean, as summarized
by Balogh and Mahunka (1983). All mea-
surements are in micromillimeters, and ex-
cept for the holotype of C. brevis, are made
from unmounted specimens. Specimen
measurements are as follows: total length—
tip of rostrum to posterior edge of notogas-
ter, width—widest part of notogaster,
height — from between genital and anal plates
to highest point of notogaster, prodorsal
length—tip of rostrum to posterior edge of
dorsosejugal depression. Line drawings were
made primarily from dissected specimens
and may be a composite of more than one
specimen. SEM’s were made from mites
stored in 70% ethyl alcohol, air dried, placed
onto sticky tape on '2 inch aluminum stubs
and coated with 200 A AuPd in a Hummer
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
IV Sputter Coater before observation in an
AMR1000 Scanning Electron Microscope.
Abbreviations of collectors names as fol-
lows: DSC—D. S. Chandler, RMR—R. M.
Reeves.
Carabodes brevis Banks 1896
Figs. 1-17
Diagnostic characters: Notogastral setae
long, nearly uniform in diameter with only
a few distal barbs; dorsosejugal depression
of nearly uniform width between bothridia;
sensillus short, capitate; notogastral pits of
moderate size (S—12 diameter) and usually
separated by less than the diameter of larg-
est pit; two pairs of aggenital setae present.
Adult. — Measurements: Total length: ho-
lotype 4 500; “‘cotypes” 2° 520, 6 545; other
material examined (mean (range), n = 13
unless otherwise noted) 2 524 (470-580), 2
481 (450-520). Notogastral width: holotype
4270; “cotypes”’ 2 300, 6 305; other material
examined 2 296 (270-330), 6 274 (250-300).
Height: holotype (not measured); ““cotypes”
2 240, 4 240; other material examined 2 238
(210-270) (n = 12), 6 209 (185-245). In-
tegument: Yellow brown, distinctly lighter
in color than C. niger Banks. Prodorsum:
Prodorsal length: holotype 4 175; “‘cotypes”
2 180, 6 190; other material examined 2? 170
(140-185), ¢ 158 (135-185). Prodorsal sur-
face (Figs. 1, 10, 11) with generally smaller
pits than on center of notogaster, most uni-
form in area between lamellae, becoming
indistinct on lamellar surfaces, the lamellar
surface with weakly developed anterolat-
erally directed striations. Base of prodor-
sum between interlamellar setae with trans-
verse folds which terminate in an inverted
V-shaped fold near middle of prodorsum.
Dorsosejugal groove present on posterior
edge of prodorsum, width uniform between
bothridial bases. Rostral setae (ro) arched
medially, minutely barbed, 32-37 long. La-
mellar setae (/e) inserted near tip of lamella,
arched medially, more strongly barbed than
ro, 45-48 long. Interlamellar setae (in) in-
serted close to where inverted V-shaped fold
VOLUME 90, NUMBER 3 375
Figs. 1-3. Carabodes brevis, adult. 1, Dorsal view. 2, Ventral view. 3, Lateral view. Scale bar in um.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 4-7.
Femur and genu. Antiaxial views. Scale bar in um.
begins, arched medially, acuminate distally,
82-88 long. Sensillus (ss) (Figs. 1, 13) short
(length beyond bothridial opening 28-38),
capitate, head spinose. Bothridial wall en-
tire. Seta ex absent. Notogaster: Notogaster
(Figs. 1, 8, 12) covered with round pits of
variable size (S—12 diameter), inter-pit dis-
tance usually less than largest pit diameter.
Pits less uniform in shape near edges of no-
togaster. Circumnotogastral depression (a
depression parallel with lateral and poste-
rior margins of notogaster) present (Fig. 8).
Ten pairs of notogastral setae present (Banks
(1896) erroneously notes that there are elev-
Carabodes brevis, adult. 4, Leg I. 5, Leg II. Femur, genu, tibia and tarsus. 6, Leg III. 7, Leg IV.
en pairs). All notogastral setae similar, long
(ta, ti and ms extending at least half way to
insertion of next posterior seta), nearly uni-
form in diameter, a few small barbs present
at or near tip (minute bars on shaft seen
only in SEM, Fig. 9), posteromarginal setae
smoother at tip in dorsal view than central
notogastral setae. Seta fa inserted anterior
to ti, thus fa, ti, ms and r, form longitudinal
row with te and r, forming a second parallel
row laterally. (Banks’ (1904, 1915) figures
of C. brevis show only nine pairs of noto-
gastral setae, one of the posteromarginal se-
tae lacking, and the fe-r, row is erroneously
VOLUME 90, NUMBER 3
Figs. 8-12.
continued posteriorly in line with the pos-
teromarginal setae.) Anterior notogastral
setae slightly longer than posterior (50-55
vs. 40-45 respectively in “‘cotypes,” 55-65
vs. 48-52 respectively in four specimens ex-
tracted from spruce branches). Position of
im normal but gland g/a sometimes ventrad
of im (Fig. 3) or posterior to 1m (Fig. 1).
Gnathosoma: Setal positions and pitting of
377
Carabodes brevis, adult. 8, Dorsal view (221 =). 9, Seta t7 (1910 =). 10, Prodorsum, anterior view
(394). 11, Dorsosejugal groove detail (760 =). 12, Pits on notogaster between f/ and ms (580). Scale bars in
um.
ventral surface shown in Fig. 14. Palpal setal
formula 0-2-1-3-9 (+ 1 solenidion). Ventral
surface: Pits on epimera similar in size to
those on prodorsum while pits on ventral
plate similar in size to those on center of
notogaster (Fig. 2). Pits on genital and anal
plates much smaller than other ventral sur-
face pits. Epimeral plates divided by fur-
rows, enlarged depression in center of first
378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 13-16. Carabodes brevis, adult. 13, Sensillus (2530). 14, Ventral view of gnathosoma and epimera
(570). 15, Genital plates (1114). 16, Anal plates (760 x). Scale bars in um.
VOLUME 90, NUMBER 3 379
Fig. 17. Known North American distribution of Carabodes brevis.
380
two furrows (Figs. 2, 14). Epimeral setal for-
mula 3-1-3-3, seta /b longest. Four pairs of
genital setae, first pair directed posteriorly,
others laterally, the latter extending *%4 dis-
tance to lateral edge of plate (Figs. 2, 15).
Two pairs of aggenital setae (ag) on ventral
plate (only one was found on one side of the
male “‘cotype’’). Two pairs of anal (an) and
three pairs of adanal setae (ad) with ad,
longest (length 35-40) and ad; shortest
(length 20-25) (Fig. 16). Setal diameter in-
creases from ad; to ad, with diameter of ad,
similar to posteromarginal notogastral se-
tae. Lyrifissure iad positioned anterior to
ad;. Pedotecta I and II (pd, and pd,) and
discidium (di) as shown in Fig. 2. Lateral
surface: Generally surface below lamella,
bothridium and edge of notogaster and
above leg insertions with small tubercles;
remainder with variously sized pits (Figs.
3). Area posterodorsad to insertion of leg
IV with tubercles joined into groups of four
or more. Pedotectum I covering base of ac-
etabulum I, widest ventrally, tapering dor-
sally to near bothridial base. Ridge present
above acetabulum II extends anteriorly onto
pd, where it becomes palmate and demar-
cates tuberculate area dorsally and pitted
area ventrally. Lyrifissures ih and ips on
ventral edge of notogaster below r;, p; and
p> (Figs. 2, 3). Lyrifissure ip posterior and
dorsal to insertion of p, (Fig. 3). Edge of
ventral plate adjacent to notogastral plate
with band of small tubercles. Legs: Pits
present on antiaxial surfaces of trochanters
III and IV, enlarged distal portion of femur
I, lower half of enlarged distal portion of
femur II, on all but the ventral flange of
femur III, and all of femur IV (Figs. 4-7).
Setation (I-IV, solenidia in brackets), tro-
chanters (1-1-2-1), femora (4-4-3-2), genua
(3(1)-3(1)-1(1)-2), tibiae (4(2)-3(1)-2(1)-
2(1)), tarsi (15(2)-15(2)-15-12). Ventrodis-
tal edge of femora III and IV with spur, that
of femur III strengthened basally. Unguinal
setae (uv) of tarsi I-IV short, wide basally,
abruptly tapered at tip. Tactile setae of all
legs barbed except for those on the dorsal
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and lateral surfaces of tibiae and tarsi. Distal
setae of tarsi, in particular fc, it, a and p,
attenuate at tip.
Immatures.— Unknown.
Material examined.—Holotype: 6, Sea
Cliff, Long Island, New York, May, N.
Banks, from dead fungus (Polyporus),
mounted in balsam. ‘“‘Cotypes”: 1 9, 1 4,
same collection data as holotype, in alcohol.
The holotype slide has on the label “(3 left)”
and probably refers to a third “‘cotype”
mounted ventrally and mislabelled accord-
ing to Norton (1978) as Carabodes dorsalis.
This specimen was not seen. The holotype
and “‘cotypes” are in the Museum of Com-
parative Zoology, Harvard University,
Cambridge, Massachusetts. Additional
specimens were examined from USA: Geor-
gia, North Carolina, West Virginia, Penn-
sylvania, New Jersey, Vermont, New
Hampshire; CANADA: Ontario, Québec,
New Brunswick, Nova Scotia, Newfound-
land. This species is thus widely distributed
in eastern North America from the southern
Appalachians to Newfoundland (Fig. 17).
The collections examined which contain
C. brevis are from a wide variety of forest
habitats including leaf litter, rotten wood,
moss, lichens, bark and fungi. Rarely has a
sample of leaf litter or rotten wood con-
tained more than three specimens while
samples which include Polyporus fungi may
contain as many as eight specimens and the
largest collection examined (25 specimens)
came from “loose bark with moss, lichens
on birch trunk” from Newfoundland. Car-
abodes niger is present and usually in greater
abundance in nearly all of my collections of
C. brevis.
Carabodes higginsi Reeves
New SPECIES
Figs. 18-35
Diagnostic characters: Prodorsum with
medium sized pits (S—7 diameter); dorso-
sejugal depression a narrow furrow; noto-
gaster covered with tubercles (7-10 diam-
VOLUME 90, NUMBER 3 381
Figs. 18-20. Carabodes higginsi, adult. 18, Dorsal view. 19, Ventral view. 20, Lateral view. Scale bar in um.
382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 21-24.
eter); all body setae setiform, notogastral
setae arise from tubercles, ta, ti and ms 1
distance to next setal insertion; sensillus
capitate, invaginated distally; 0-2 aggenital
setae.
Adult.— Measurements (mean (range), n
= 9 for 2, n = 2 for 4): Total length: 2 378
(355-390), 3 338 (315-360). Notogastral
width: 2 221 (205-230), 4 192 (180-205).
Height: 2 183 (170-195), 6 162 (145-180).
Integument: Light brown. Thin cerotegu-
ment seen only on prodorsum. Prodorsum:
Prodorsal length: 2 103 (95-110), 4 88 (85-
90). Dorsosejugal depression a narrow fur-
row. Prodorsal surface (Figs. 18, 27) cov-
ered with pits (diameter 5-7), pits generally
more circular in middle of prodorsum, ex-
tend to lateral edge of lamellae. Rostral (ro)
Carabodes higginsi, adult. 21, Leg I. 22,
Leg II. Femur, genu, tibia and tarsus. 23, Leg III. 24,
Leg IV. Femur and genu. Antiaxial views. Scale bar in um.
and lamellar (/e) setae setiform, bent me-
dially, closely appressed to prodorsal sur-
face, very minutely barbed at 1000x, le
(length 27) longer than ro (length 17). In-
terlamellar setae (in) (length 32) longer than
le, setiform, slightly arched. Sensillus (ss)
(Figs. 18, 30) capitate with distal surface
invaginated, forming an irregularly edged
cup-like opening. Edge of bothridium with
ventral notch (Figs. 20, 30). Notogaster: No-
togaster (Figs. 18, 25) covered with tuber-
cles (diameter 7-10), sometimes so closely
appressed so as to partially flatten adjacent
surfaces (Fig. 28). Circumnotogastral
depression not apparent dorsally and only
barely visible in lateral view (Fig. 20). Two
short ridges present on lateral edge of no-
togaster just posteriad of humeral angle (Fig.
VOLUME 90, NUMBER 3
Figs. 25-29.
18). Lyrifissure 7a present on posterior ridge.
Notogastral setae (Figs. 18, 29) setiform,
erect, seta fa longest (35-38), others de-
creasing in length posteriorly to postero-
marginal setae (p,—p;, 15-20); each seta
arising from middle of tubercle (Figs. 28,
29); ta and ti extending approximately '4 to
’o distance to insertion of ti and ms, re-
spectively. Seta ta positioned anterior to fi,
thus notogastral setae forming two rows: fa,
w
co
w
Carabodes higginsi, adult. 25, Dorsal view (300 x). 26, Lateral view (253 x). 27, Prodorsum,
anterior view (504 x). 28, Notogastral tubercles around seta ti (2270 x). 29, Seta ms (6400 x). Scale bars in um.
ti, ms and r, medially and fe and r, laterally.
Gnathosoma: Pits and setae on mentum as
shown in Fig. 31. Palpal formula: 0-2-1-3-9
(+ 1 solenidion). Ventral surface: Pits pres-
ent on all ventral surfaces, pits variable in
size and shape, generally smaller than those
on center of prodorsum, smallest on genital
and anal plates. Epimeral plates (ep,—-ep,,
Figs. 19, 31) divided by furrows, epimeral
setal formula 3-1-3-3, setae setiform, short
384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 30-34. Carabodes higginsi, adult. 30, Sensillus (3670 x). 31, Ventral view of gnathosoma and epimera
(460 x). 32, Genital plates (1100). 33, Lateral view of genital plates (1390). 34, Anal plates (890 x). Scale
bars in um.
VOLUME 90, NUMBER 3
Fig. 35. Known North American distribution of Carabodes higginsi (&) and C. willmanni (@).
386
(8-9). Genital, anal and aggenital setae se-
tiform, short (8-9). One pair of aggenital
setae (Figs. 19, 33) present in 8 of 9 females
and 3 of 6 males, only one ag present on
one female and one male and ag absent in
two males. Adanal setae setiform, length of
ad, similar to ag, ad, and ad, longer with
ad, approximately twice length of ad;. Lyr-
ifissure iad lateral to ad;. Pedotecta I and
II and discidium present as shown in Figs.
19, 20. Lateral surface: Most of lateral sur-
face below edges of lamellae, bothridium
and notogaster but above leg insertions
without pits (Fig. 20). Small tubercles pres-
ent in area immediately above acetabulum
I and partially hidden by pedotectum I and
on posterior edge of integumental fold that
extends from humeral angle ventrally to
acetabulum II. Alveolar remnant of exo-
bothridial setae (ex) present immediately
below bothridium. Circumnotogastral
depression faintly indicated below im and
seta ms. Legs: Pits present on antiaxial sur-
faces of femora I and IJ and trochanters and
femora III and IV (Figs. 21-24). Setation
(I-IV, solenidia in brackets), trochanters 1-1-
2-1, femora 4-4-3-2, genua 3(1)-3(1)-1(1)-
2, tibiae 4(2)-3(1)-2(1)-2(1), tarsi 15(2)-
15(2)-15-12. Ventral flange of femur III
weakly developed. More leg setae are gla-
brous in C. higginsi than in C. brevis with
the barbed condition retained on paraxial
setae d’ on femora I and II, v’ on genu I, v’
and /' on genu II, v’ on tibia I and antiaxial
setae v” on tibiae III and IV. Solenidion (¥)
of tibia II shorter than in C. brevis and sim-
ilar in size and shape to tarsal II solenidia
w, and w>, a condition also noted by Bernini
(1976) for C. minusculus. Setae (p) of tarsus
I short, blunt-tipped and only slightly longer
than s. Distal setae of all tarsi, in particular
tc, it, aand p (except leg I), attenuate at tip.
Immatures.— Unknown.
Material examined: Holotype: adult 8,
USA, NEW HAMPSHIRE, Carroll Co., 2.5
mi. NW Wonalancet, The Bowl, VHI-6-85,
RMR, sifted spruce litter; deposited in Ca-
nadian National Collection. Paratypes: | °,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
same data except X-1-85, DSC, sifted co-
nifer leaf litter; 1 9, same data except VII-
30-85, DSC, sifted hemlock/fir leaf litter; 1
6, same data except VI-21-85, DSC; 1 4,
same data except VIII-6-85, sifted litter by
stream; | 6, Strafford Co., College Woods,
Durham, IV-8-65, RMR, moss on log; 1 8,
same data except VIII-7-85, sifted leaf litter,
rotten wood and stumps; | 9, 1 6, 3 sex
undetermined, Coos Co., 17 mi. N Crystal,
IX-9/11-83, A. Godfrey, extracted from
spruce branches; 2 2, same data except ex-
tracted from fir branches; 1 6, same data
except 3 mi. NE Errol, [X-3/4-83, extracted
from fir branches; 1 9, PENNSYLVANIA,
Huntington Co., Alan Seeger Natural Area,
V-30-85, DSC, sifted pine leaf litter; 1 9,
VIRGINIA, Giles Co., White Pine Lodge
nr. Mountain Lake, [X-12-67, J. M. Camp-
bell, white pine duff, 1 9, CANADA, NOVA
SCOTIA, Cape Breton Highlands Natl. Pk.,
Paquette Lake Trail, VIII-29-84, V. Behan,
lichens on rocks on Glasgow Lake Trail; 1
6, Kejimkuyik Natl. Pk., VIII-17-68, E. E.
Lindquist, Polyporus on dead fir and spruce
trunks; | 4, 12 sex undetermined, same data
except VIII-18-68, 1’ white pine duff. Para-
types deposited in United States National
Museum, Washington, D.C., Museum of
Comparative Zoology, Cambridge, Massa-
chusetts, Canadian National Collection,
Biosystematics Research Centre, Ottawa and
the personal collections of R. A. Norton and
the author.
This species is distributed from the Ca-
nadian Maritime Provinces southwest to
Virginia. It has been collected most often
from conifer leaf litter, bark and branches
and usually as single specimens. Fir and
spruce branches and white pine duff have
provided the largest number of specimens/
sample.
Remarks. — Carabodes higginsi belongs to
the minusculus group by having the noto-
gaster covered with tubercles, a narrow dor-
sosejugal furrow and by the shape of the
sensillus. It has longer and more tapered
notogastral setae than C. wi//manni, which
VOLUME 90, NUMBER 3
is the only other species of this group known
from North America. The absence of ag-
genital setae in this group has been noted
only in C. pulcher Bernini (Bernini 1976).
This species is named after an oribatol-
ogist and friend, the late Harold G. Higgins,
of Salt Lake City, Utah.
DISTRIBUTION AND HABITAT
COMPARISONS
The distribution and habitat preferences
for the remaining five species are based on
literature, material in the CNC and my own
personal collection.
Carabodes granulatus is known from II-
linois, Kentucky, New York and North Car-
olina (Marshall et al. 1987). The additional
specimens seen are from Newfoundland,
Nova Scotia, New Brunswick and Ontario
in Canada and New Hampshire, Vermont,
Massachusetts, Pennsylvania, Virginia,
South Carolina, Georgia, Florida, Missis-
sippi, Missouri and Oklahoma in United
States. It has possibly the widest distribu-
tion in eastern North America of any Car-
abodes species and is found from New-
foundland south to northern Florida and
west to southern Ontario, Missouri and
Oklahoma (Fig. 36). It is also one of the
most abundant species in leaf litter and rot-
ten wood samples from Pennsylvania and
New Hampshire but may also be found in
sphagnum and other mosses, lichens, bark,
grass sod and fungi.
Carabodes labrynthicus is a commonly
collected Holarctic species in Europe and
North America. It has the most northerly
distribution of any Carabodes in the Nearc-
tic zone (Fig. 37). Marshall et al. (1987) lists
this species from Québec, Northwest Ter-
ritories, Yukon Territory, Alaska and Vir-
ginia. The CNC and my own collection have
additional specimens from CANADA: Lab-
rador, Newfoundland, Prince Edward Is-
land, Nova Scotia, New Brunswick, Ontario
and Manitoba and USA: Maine, New
Hampshire, Vermont, New York, Pennsyl-
vania and New Jersey. The CNC also has
387
specimens from Magadan, Magadan Re-
gion, USSR. It is the only species of Car-
abodes 1 have collected above 1220 m ele-
vation on Mt. Washington, New Hampshire
with the highest collections at 1585 m ele-
vation. The highest numbers I have seen in
samples have come from leaf litter and rot-
ten wood from New England, although
specimens were also present in samples from
moss, lichens, grass, fungi and wrack. Seyd
and Seaward (1984), in their review of orib-
atid mite/lichen associations, consider that
this species, while showing a preference for
lichens, is also adapted for existence on oth-
er plants.
Carabodes niger has been recorded from
New York, Ohio, Virginia and North Car-
olina (Marshall et al. 1987). Additional
specimens in the CNC and my own collec-
tion extend this distribution to Newfound-
land, Prince Edward Island, Nova Scotia,
New Brunswick, Québec, Ontario and Man-
itoba in Canada and Maine, New Hamp-
shire, Vermont, Massachusetts, Pennsyl-
vania, New Jersey and Maryland in the
United States (Fig. 38). It 1s difficult at this
time to determine the southerly distribution
of this species below Pennsylvania and
Maryland because material I have seen from
further south contain many as yet unde-
scribed species closely related to C. niger.
Norton (1978) has pointed out that the
length of the posteromarginal setae on spec-
imens from Ohio are longer, nearly reaching
the insertion of the next seta, while on type
specimens from New York these setae are
less than half this distance. I have also noted
this and find the longer setal lengths pre-
dominate in samples from northern New
England and Canada while shorter setal
lengths predominate in samples from Mas-
sachusetts (Martha’s Vineyard), New York
(Long Island), New Jersey, Maryland and
Pennsylvania. A detailed study of C. niger
and closely related species will be necessary
to resolve whether C. niger is a complex of
species or simply showing clinal variation
with latitude.
388 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pel
gree
Fig. 36. Known North American distribution of Carabodes granulatus.
Banks (1895) collected the type speci-
mens from Polyporus fungi, and Norton
(1978) noted that most collections from New
York and Ohio have been from fungi, es-
pecially Polyporus. There are many collec-
tions in the CNC labelled “bracket fungi”
and ‘Polyporus fungi” in association with
logs, stumps and tree trunks. The dominant
VOLUME 90, NUMBER 3
389
Fig. 37. Known North American distribution of Carabodes labrynthicus.
species in these samples is nearly always C.
niger while the species most often found
with niger is C. brevis. In my collection
nearly all samples containing C. brevis also
contain C. niger with the latter species dom-
inant numerically. Thus, a close habitat as-
sociation between these two species is evi-
dent. However, I have usually found C. niger
is more abundant in forest leaf litter and the
most commonly encountered Carabodes
390 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 38. Known North American distribution of Carabodes niger.
species from forest leaf litter, rotten wood, Carabodes willmanni is Holarctic and is
bark and fungi in Pennsylvania and north- distributed in Europe from Scandinavia
ern New England. It has also been collected south to Italy and Spain and known in North
from moss, lichens and grass sod. America only from central New Hampshire
VOLUME 90, NUMBER 3
Table 1.
rotten wood collections.
391
Percent representation of Carabodes species from branch extractions and from sifted leaf litter/
Branch Extractions
Sifted Leaf Litter/Rotten Wood
Species of Spruce-fir Spruce-fir Mixed forest
brevis a2 <1 <1
dendroetus 37, 0) =<
higginsi 5 0 al
willmanni 2 0 0
granulatus 2 <1 41
niger | 11 46
labrynthicus eal 87 2
Other spp. _ 0 pail 11
Total specimens 129 3546 1602
(Bernini 1975). I have seen additional ma-
terial from Labrador, Newfoundland, Nova
Scotia, New Brunswick, New Jersey and
from spruce and fir branches in northern
New Hampshire (Fig. 35). Samples with the
largest number of specimens (43 maximum)
are from lichens or lichens mixed with
woody shrubs while samples from leaf litter,
moss, bark or grass contain one to three
specimens only. In Europe C. willmanni is
a common inhabitant of lichens (Bellido
1979, Colloff 1983, Seyd and Seaward 1984)
and Bellido (1979) has reported on the in-
fluence of temperature and moisture on the
seasonal abundance of larvae, nymphs and
adults of this species while feeding in li-
chens.
Table | gives the percentages of individ-
uals recovered from branch samples pro-
cessed with caustic soda and compares them
with leaf litter/rotten wood samples from a
virgin spruce-fir stand at Norton Pool, 3 mi.
E of East Inlet Dam, Coos Co., NH, and a
virgin mixed forest at “The Bowl,” 2.5 mi.
NW Wonalancet, Carroll Co., NH. A more
complete description of these areas may be
found in Lyon and Reiners (1971). The
dominant Carabodes species on branches
were brevis and dendroetus while leaf litter/
rotten wood was preferred by C. granulatus,
niger and labrynthicus. Carabodes willman-
ni and higginsi were absent or poorly rep-
resented in all these habitats suggesting that
these are not favored habitats for these
species. All seven species were collected from
spruce branches while only four (C. brevis,
dendroetus, higginsi and granulatus) were
collected from fir branches. Also 85% (56
specimens) of C. brevis were collected from
spruce branches. Thus spruce branches seem
to be preferred to those of fir, possibly as a
result of the rougher bark surface or the po-
sition of the needles around the twigs.
In summary the above evidence indicates
considerable variation in habitat prefer-
ences for certain species. Carabodes brevis
and dendroetus are primarily arboreal with
C. brevis often associated with C. niger in
fungal fruiting bodies. Carabodes granula-
tus, labrynthicus and niger, while more
common in leaf litter/rotten wood, may be
arboreal, with granulatus and labrynthicus
usually associated with lichens and moss
and niger with fungi. Carabodes willmanni
is probably limited to bark and branches
where lichens are present. The caustic soda
wash technique did not indicate habitat
preferences for C. higginsi but other collec-
tion information point toward a preference
for coniferous litter or bark.
ACKNOWLEDGMENTS
Scientific Contribution Number 1546
from the New Hampshire Agricultural Ex-
periment Station.
The Central University Research Fund,
392
University of New Hampshire, provided fi-
nancial support for scanning electron mi-
crographs used in the descriptions. I thank
Stanley S. Swier and April Godfrey for pro-
viding the extracted branch samples, Don-
ald S. Chandler for collecting many of the
leaf litter/rotten wood samples and Valerie
Behan-Pelletier for providing access to the
CNC material. Iam grateful to Ardis John-
ston, Museum of Comparative Zoology,
Cambridge for the loan of the type speci-
mens of C. brevis. I thank Valerie Behan-
Pelletier, Biosystematics Research Centre,
Ottawa, Canada and Donald S. Chandler
and John S. Weaver III, University of New
Hampshire, for review of the manuscript.
LITERATURE CITED
Balogh, J. and S. Mahunka. 1983. The Soil Mites of
the World. Vol. 1: Primitive Oribatids of the Pa-
laearctic Region. Elsevier: New York. 372 pp.
Banks, N. 1895. On the Oribatoidea of the United
States. Trans. Am. Entomol. Soc. 22: 1-16.
1896. New North American spiders and
mites. Trans. Am. Entomol. Soc. 23: 57-77.
1904. A treatise on the Acarina, or mites.
Smithsonian Institution. Proc. U.S. Nat. Mus.
28(1382): 1-114.
. 1915. The Acarina or mites. A review of the
group for the use of economic entomologists.
USDA Rep. 108. 153 pp.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Bellido, A. 1979. Ecologie de Carabodes willmanni
Bernini, 1975 (Acari, Oribatei) dans les forma-
tions pionniéres de la lande armoricaine. Rev. Ecol.
Biol. Sol. 16: 195-218.
Bernini, F. 1975. Notulae Orbatologicae XII. Una
nuova specie di Carabodes affine a C. minusculus
Berlese 1923 (Acarida, Oribate1). Redia 56: 455-
471 + pls. 144.
. 1976. Notulae Onbatologicae XIV. Revision
di Carabodes minusculus Berlese 1923 (Acarida,
Oribatei). Redia 59: 1-49 + pls. 1-9.
Colloff, M. J. 1983. Oribatid mites associated with
marine and maritime lichens on the island of Great
Cumbrae. Glasg. Nat. 20: 347-359.
Lyon, C. J. and W. A. Reiners (Eds.). 1971. Natural
areas suitable for ecological research. Dept. Biol.
Sci. Publ. No. 4, Dartmouth College, Hanover,
NH. 75 pp.
Marshall, V. G., R. M. Reeves, and R. A. Norton.
1987. Catalogue of the Oribatida (Acari) of con-
tinental United States and Canada. Mem. Ent. Soc.
Can. 139. 418 pp.
Miller, C. A. and G. A. McDougal. 1968. A new
sampling technique for spruce budworm larvae.
Can. Dep. For. Rur. Dev., Ottawa, Ont. Bi-Mon.
Res. Notes 24: 30-31.
Norton, R. A. 1978. Notes on Nathan Banks species
of the mite genus Carahodes (Acari: Oribatei). Proc.
Entomol. Soc. Wash. 80: 611-615.
Seyd, E. L. and M. R. D. Seaward. 1984. The asso-
ciation of oribatid mites with lichens. Zool. J. Linn.
Soc. 80: 369-420.
Travé, J. 1963. Ecologie et biologie des Oribates
(Acariens) saxicoles et arboricoles. Vie et Milieu
14: 1-267.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, p. 393
NOTE
The Newly Discovered Spring Crown Gall of Asphondylia
rudbeckiaeconspicua (Diptera: Cecidomytiidae) on Rudbeckia
laciniata (Asteraceae) in Pennsylvania
Asphondylia rudbeckiaeconspicua Osten
Sacken (Diptera: Cecidomyiidae) forms a
conspicuous, apical summer gall on the
meristems and flower discs of Rudbeckia
laciniata L. (Asteraceae) (Felt, E. P. 1940.
Plant Galls and Gall Makers. Comstock
Publishing Co., Ithaca, N.Y. 364 pp.). The
gall is large, globular, usually about 5 cm
across, and always polythalamous. Until
now, only the summer galls of 4. rudbeck-
laeconspicua were known.
Because all of the known Asphondaylia spp.
overwinter as larvae in living plant tissue
(R. J. Gagné, personal communication) A.
rudbeckiaeconspicua must either lay its eggs
in the crown of R. laciniata, the only part
of the plant that survives winter above
ground, or use another host. Assuming the
former to be the more likely case, I searched
the crowns of R. /aciniata in early June 1985
at Pittsburgh, Pennsylvania and there lo-
cated six basal bud galls. They were ap-
proximately 4 cm high and 2.5 cm wide,
and they originated at the base of the 1985
stems. This indicated that the eggs or young
larvae overwintered in these buds. The galls
were analogous to summer galls in that they
were polythalamous, with one larva devel-
oping per cell.
Pupae taken from these spring galls, and
adults reared on 10 June 1985 were iden-
tified as 4. rudbeckiaeconspicua by Dr. R.
J. Gagné of the Systematic Entomology
Laboratory, USDA, Washington, D.C. This
confirms that A. rudbeckiaeconspicua has
two generations per year, one in spring crown
galls, and the second in summer apical galls.
From this discovery it now seems likely
that other 4sphondylia, known only from
apical summer galls on Asteraceae, also have
another gall elsewhere on their host plant
where they overwinter.
John D. Plakidas, 8220 Walter Martz
Road, Frederick, Maryland 21701.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 394-395
NOTE
A Polynomial Riley Name in Cecidomyiidae (Diptera) and
Implications of Such Names for Cynipidae (Hymenoptera)
In an article on insects injurious to grape
vines, Riley (1873, Fifth Ann. Rept. Nox-
ious and Beneficial Insects of the State of
Missouri, p. 117) coined the name Vitis to-
matos for tomato-like galls on grape. This
name has not to my knowledge been used
in the scientific literature since then. Riley
evidently intended to use the name to refer
to the gall only, because he wrote that the
gall was made by the gall midge Lasioptera
vitis Osten Sacken 1862. A name proposed
before 1931 for the work of an animal may
be considered available in zoological no-
menclature (Int. Code. Zool. Nomen. 1984,
3rd ed., Art. la), but Riley did not explain
why he used separate names for the gall and
gall maker. The question now is whether
the generic or specific names of Vitis to-
matos are available for taxonomic use. The
answer is important because L. vitis is not
the actual gall maker. Instead, that species
appears to be only an occasional inquiline
in galls caused by Janetiella brevicauda Felt
1908 (pers. observ.; R. B. Johnson unpub.
thesis, Cornell University, 1952). If Vitis is
an available generic name, it could be used
for a group of grape-feeding species that in-
cludes brevicauda and that are now in the
catch-all genus Janetiella; if tomatos is
available, it would be a senior synonym of
brevicauda.
Judged solely from the contents of Riley’s
paper (ibid.), Vitis tomatos would appear to
be available for use. To find out what Riley
might have intended, one needs to look else-
where. Earlier in Riley’s paper (p. 114), the
name Vitis pomum Walsh and Riley is used
for another gall on grape. That species had
been described previously as [Cecidomyia]
Vitis pomum (Walsh and Riley 1869, Am.
Entomol. 1: 106). The name Cecidomyia
was understood, being the heading (p. 105)
of the section in which several gall midges
and their galls were described. Walsh and
Riley (ibid.) coined many other names in
that formula: the generic name understood
and not repeated for each species; another
word capitalized and in the genitive form
of the plant name; and the final word de-
scriptive of the gall. For [Cecidomyia] Vitis
pomum that meant, “apple [gall] of grape
[formed by a Cecidomyia].”
Two separate reasons to invalidate Vitis
tomatos appear to be present: that Vitis to-
matos is in reality a polynomial and that
Vitis is in the genitive case. Polynomials are
not available according to binomial no-
menclature and so are not considered by the
International Code of Zoological Nomen-
clature (ICZN 3rd ed., 1985). There is a
provision of the ICZN (Art. 1 1h(v)) to ac-
cept species-group names that were pub-
lished as separate words referring to a single
entity. For example, Cecidomyia piniinopis
Osten Sacken was originally coined as Ce-
cidomyia pini inopis but, because pini inopis
is based on the host, then known as Pinus
inopis, the separate words are closed up and
the name considered available from its orig-
inal description. But Vitis pomum is not
available from 1869, when proposed by
Walsh and Riley, but from 1878, when Os-
ten Sacken (1878, Smithson. Misc. Colls.
270: 7) combined the two separate words
as Cecidomyia vitis-pomum, thus satisfying
the provisions of binominal nomenclature
and making Osten Sacken the author as of
1878.
The second point one notices when leaf-
ing through the paper by Walsh and Riley
is that the first word of any two-word name
is in the genitive case, e.g. Salicis brassi-
coides Walsh (p. 105) and Populi vagabunda
Walsh (p. 107). Vitis, too, if one assumes
VOLUME 90, NUMBER 3
the practice was continued, must be in the
genitive case, although that name, being in
the third Latin declension, takes the same
form for the nominative and genitive cases.
The International Code of Zoological No-
menclature (Art. 1 1g) requires that a generic
name be in the nominative singular.
Indications are that Riley formed Vitis
tomatos following the pattern used by Walsh
and Riley of using polynomial names and
using the penultimate word in the genitive
case. Vitis tomatos, then, appears to be in-
validly constructed and not available for
taxonomic use.
While researching this problem, I noticed
that the Catalog of Hymenoptera in Amer-
ica North of Mexico (Krombein, Hurd,
Smith, and Burks 1979, Smithson. Inst.
Press) improperly lists many cynipid names
as available from the date their specific
names were coined as two independent
words that do not refer to a single unit, un-
395
like pini inopis above. Consider Atrusca
quercuscentricola (ibid., p. 1090), which was
described as Cynips quercus centricola (Os-
ten Sacken 1861, Proc. Entomol. Soc. Phila.
1: 58): the name should be Afrusca centri-
cola and date from 1865 when Osten Sacken
(1865, Proc. Entomol. Soc. Phila. 4: 345)
first used a single word for the specific name
of that species.
I am grateful to L. G. Clark for sending
me a copy of the R. B. Johnson thesis and
to W. N. Mathis, R. V. Peterson, C. W.
Sabrosky, G. C. Steyskal, and F. C. Thomp-
son for their comments on an early draft of
this paper.
Raymond J. Gagné, Research Entomol-
ogist, Systematic Entomology Laboratory,
PST, Agricultural Research Service, USDA,
% U.S. National Museum NHB 168, Wash-
ington, D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 396-397
Book REVIEW
The Behavioural Ecology of Ants. By John
H. Sudd and Nigel R. Franks. Blackie,
Chapman and Hall, New York. 1987, x
+ 206 pp. $55 cloth, $23 paperback.
The common perception of ants as busy
automatons, harmoniously and altruistical-
ly functioning for the greater good of queen
and colony, should be shaken by a reading
of The Behavioral Ecology of Ants, by John
H. Sudd and Nigel R. Franks. Instead of
harmony and altruism, the authors describe
oligogynous and polygynous colonies (those
with few and many reproductive females),
intracolonial struggles for reproductive
dominance, workers killing their queens,
workers producing all of a colonies’ male
off-spring, ritualized antagonistic interac-
tions among workers, infiltration of colo-
nies by murderous socially parasitic ants,
and the use of propaganda pheromones; all
to maximize each ant’s reproductive suc-
cess. The book deals with the eusociality of
ants, wherein individual ants forgo repro-
ducing to enhance the reproduction of a
nestmate; how eusociality could evolve in
ants, and why, except for termites, it is re-
stricted to Hymenoptera. W. D. Hamilton’s
(1964) hypothesis of kin selection based on
haplodiploid reproduction in Hymenoptera
is used to explore and explain the social
activities of ants, particularly social altru-
ism to close kin.
Written for use by advanced undergrad-
uates, graduate students, and researchers in
entomology, ecology and behavior, the book
may prove to be a bit too challenging for
undergraduates. It is, however, admirably
suited for a post baccalaureate audience fa-
miliar with the specialized terminology.
While crucial terminology is defined, the
inclusion of a glossary would have been use-
ful.
In an undogmatic style, suited to a field
in which today’s truth can sometimes be-
come tomorrow’s fiction, Sudd and Franks
steer the reader through the intricacies of
ant communication, reproductive strate-
gies, foraging behavior, caste systems, social
parasitism and more. Not unexpectedly,
some subjects receive a more thorough
treatment than others, but for the most part
they are adequately presented. A reader ea-
ger for more information on an interesting
topic will notice how well, with few excep-
tions, the book is documented (over 300
references).
The illustrations are of generally high
quality; however, the cover drawing both
fascinates and irks me. It is a rather striking
computer-generated 3D line image of an
army ant worker. At first glance, something
seemed amiss about the ant. A more focused
inspection revealed that the antennae were
thread-like; each drawn with a single line,
egregiously and inexplicably out of propor-
tion.
Some minor slip ups: grammatical (e.g.
data was [sic]), factual (e.g. Trachymyrmex
septentrionalis is primarily an ant of some-
what xeric woodlands from Texas eastward,
rather than a desert ant, as stated), and in
the adaptation of figures from other publi-
cations (e.g. Fig. 2.1), can be found. Some
readers may disagree with some of the au-
thors’ statements, such as those about intra-
and interspecific competition, which seem
to ignore the phenomenon of invading
species like the red imported fire ant, Sole-
nopsis invicta. We are told about r and K
strategists in Chapter 7, and referred to Sec-
tion 1.4.1 for a prior explanation which is
not to be found. Such slight defects are more
than outweighed by the value of the book
to myrmecologists and other interested par-
ties, if only for the fact that it summarizes
much of the information explosion on ant
behavior and ecology, occurring over the
past two decades.
Overall there is much which can be said
VOLUME 90, NUMBER 3
favorably about The Behavioral Ecology of
Ants, and I especially recommend it for
myrmecologists and those teaching courses
in insect behavior and evolution. It is a rel-
atively slender book (206 pages), but packed
397
with facts and thought-provoking theories
which fascinated me.
J. F. Carroll, Livestock Insect Laboratory,
Rm. 120, B-307, BARC-East, ARS, USDA,
Beltsville, MD 20705.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, p. 397
Book REVIEW
The Metallic Wood-boring Beetles of
Canada and Alaska: Coleoptera: Bupres-
tidae. By Donald E. Bright, Biosystemat-
ics Research Centre, Agriculture Canada,
Ottawa, Ontario. Part 15. The Insects and
Arachnids of Canada. (Publication 1810).
335 pp., 264 figures, 87 maps. 1987. Cost:
$14.90 (Canada), $17.40 (Other Coun-
tries, Canadian funds).
This is another in the fine series of hand-
books for the identification of Canadian ar-
thropods written by systematists of the Bio-
systematics Research Centre and outside
collaborators. Although this volume is pri-
marily concerned with the Canadian fauna,
it will be useful to workers in the United
States since most of the species are found
in both countries. The author included some
of the species found in the northern states
that should extend into Canada but that are
not yet recorded north of the border.
The text is English but identification keys
to all taxa are in both English and French.
A generous diagnostic description for each
genus and species 1s given along with an
up-to-date nomenclature, the principal his-
torical taxonomic combinations and refer-
ences, lists of host plants, the general dis-
tribution, and identification notes. Habitus
drawings are provided for at least one species
in each genus, and male genitalia are illus-
trated for nearly all of the species. Distri-
bution maps for the Canadian records of
119 of the 189 species or subspecies are
included. A short glossary of morphological
terms, a complete list of references, and an
index complete the volume.
I can find little to criticize about this man-
ual. One might wish for more habitus illus-
trations, but the keys and descriptions sup-
ported by line drawings of male genitalia
appear to be adequate.
Taxonomists, museum workers, forest-
ers, pest control operators, port identifiers,
and students will find this a valuable ref-
erence for their libraries especially if they
do not specialize in the family Buprestidae.
John M. Kingsolver, Systematic Ento-
mology Laboratory, Agricultural Research
Service, USDA, % NHB 168, National Mu-
seum of Natural History, Washington, DC
20560.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, pp. 398-400
Book REVIEW
The Ixodid Ticks of Uganda. By John G.
Matthysse and Murray H. Colbo. Ento-
mological Society of America, 4603 Cal-
vert Road, College Park, MD 20740. xu
+ 426 pp., 166 plates. 1987. $35.00/pa-
per. ISBN 0-938522-31-0 (500 copies
printed).
In 1963, J. G. Matthysse was a member
of a U.S. Department of Agriculture team
sponsored by the U.S. Agency for Interna-
tional Development (USAID) concerned
with livestock development in Uganda with
special emphasis on improved cattle ranch-
ing in Ankole District. This was an area
being successfully cleared of tsetse flies
through USAID financial support. The final
report submitted by that team recommend-
ed that a taxonomic, biological and eco-
nomic survey of the ticks of Uganda be un-
dertaken as a basis for tickborne disease
control.
A joint research program between the
Government of Uganda and USAID was
initiated in July 1965 and later that same
year Murray H. Colbo joined the team. From
this work a book-length manuscript, in-
cluding plates and maps, was completed in
July 1969. As the Entomological Society of
America flyer for this book announces, “‘it
was twenty years in the making, but [xodid
Ticks of Uganda has finally arrived.” I will
discuss that delay a bit further in this re-
view.
At the time this book was conceived, all
East African countries and most countries
in the rest of Africa carried out tick and
tickborne disease control programs. How-
ever, in recent years, wars and other civil
disturbances, as well as natural disasters,
have combined to curtail control of ticks
and tickborne diseases over much of the
continent. Ticks and tickborne diseases are
among the most important factors in pre-
venting the improvement of livestock hus-
bandry in Africa. In much of East Africa,
breed improvement is prevented by the sus-
ceptibility of introduced cattle to tickborne
diseases. Throughout Africa, ticks and the
diseases they transmit cause great economic
loss in livestock through death (especially
in calves), stunting and poor growth, wound
exposure to other infections and infesta-
tions, interference in trade, and great ex-
pense in application equipment, acaricides,
immunizations and drug treatments.
The Ixodid Ticks of Uganda is the fifth
book to be published dealing with the ticks
ofan East African country and, to my mind,
it is the most useful. For those readers who
are interested, the others are Harry Hoog-
straal’s Ticks of the Sudan (1956), a mon-
umental piece of work but becoming out-
dated; Guy Yeoman and Jane Walker’s The
Ixodid Ticks of Tanzania (1967) and Walk-
er’s The Ixodid Ticks of Kenya (1974), both
very useful references but without keys or
figures; and Pierre Morel’s Etude sur les Ti-
ques d’Ethiopie (1976), again a useful work
but with only a few figures of Rhipicephalus
species and no keys.
The text of The Ixodid Ticks of Uganda
is roughly divided into two parts (there are
no chapters as such). The first part is intro-
ductory in nature, giving the reader a back-
ground of previous tick work conducted in
Uganda, followed by a detailed description
of the country, including physiography, cli-
mate, ecological zones, vegetation types, and
livestock and wild host distributions. This
is all important information and the authors
subsequently discuss tick species in relation
to these factors and show that tick distri-
bution is regulated by rainfall, temperature,
altitude and vegetation as well as by host
availability. The only problem I have with
this section is my inability to decipher some
of the maps. For example, trying to read the
VOLUME 90, NUMBER 3
legend on the ecological zone map (p. 13)
is impossible without the aid of a magni-
fying glass. There follows a methods section
giving procedures for mapping tick collec-
tions and preparing transects, together with
specific locality coordinates and a list of
hosts by district from which ticks were col-
lected. Again, the only difficulty I have with
this section is the map on page 19 where
Uganda collecting locations are marked with
black circles, triangles, and squares. What
do these three different symbols mean?
There is no symbol legend with this map to
guide us, and it is not until we reach the
bottom of page 25 that the symbols are ex-
plained.
The remainder of the book is devoted to
keys, descriptions, hosts, distributions, and
figures of the genera and species of Ixodidae
occurring or likely to be found in Uganda.
Eleven genera are discussed, with the genera
Amblyomma, Haemaphysalis, Ixodes and
Rhipicephalus occupying the major portion
of the book. The Amblyomma species are
illustrated with light microscopy photo-
graphs, Haemaphysalis species by pen and
ink drawings from Hoogstraal publications
in the Journal of Parasitology, Ixodes species
by both pen and ink illustrations and scan-
ning electron photomicrographs, and Rhipi-
cephalus species primarily by light micros-
copy photographs, but with a few SEM’s
and some pen and ink drawings of female
genital apertures. For some strange reason,
a few of the figures in the plates were printed
upside down—see plate 68, fig. 3; plate 129,
fig. 2; plate 155, fig. 5; and plate 164, fig. 4.
I have not tested all the keys, but for sev-
eral years I have used the authors’ Rhipi-
cephalus key. Before running specimens
through that key, any worker new to this
genus should read the warning on page 273
on how difficult it can be to correctly iden-
tify African species of Rhipicephalus. As
Harry Hoogstraal once said to me, “God
created the genus Rhipicephalus, and even
He can’t identify them.” I say that I have
used the Rhipicephalus key for several years
399
because a version of this book has been in
the hands of a few tick taxonomists for a
long time. Beyond the fact that this is a fine
addition to the literature on African ticks,
Matthysse and Colbo deserve medals for
their patience and perseverence during the
twenty years that preceded publication. A
brief history of this long path to press may
be of interest.
Originally, Matthysse planned to have this
book published by the Commonwealth In-
stitute of Entomology, and in August, 1969
delivered typescript to them in London.
While still there, he received a cable from
the State Department in Washington, D.C.
instructing him to deliver it to the Uganda
Government Printer in Entebbe, which he
did. It should be noted that the Uganda
Government Printer is part of the Presi-
dent’s office and high level priorities rule
the sequence of printing operations. It should
also be noted that in 1966 Milton Obote led
a revolution in Uganda and installed him-
self as President, so the government at that
time was relatively unstable, and frequent
border clashes were taking place with Tan-
zania. In 1971, Idi Amin Dada seized pow-
er, toppling the regime of Milton Obote, and
in 1972 Amin ordered the expulsion of
Uganda’s 60,000 Asians, among whom were
the skilled workers employed by the Uganda
Government Printer. Between 1969 and
1973, despite repeated attempts, no news
was received by Matthysse and Colbo on
the fate of their book. In 1974, they were
told that the book was scheduled to be out
in June. In February 1975, they were told
that a bound copy had appeared but without
illustrations. In fact, a few copies of this
book actually exist, though without figures,
maps, publisher information, or even the
authors’ names! Torture of this kind con-
tinued until 1980 when Matthysse received
a letter from Entebbe informing him that,
“nota trace remains of the manuscript” and
that the Uganda Government Printer had
been hard hit by the war. So it was back to
square one. The book was now out of date
400
and between 1980 and 1987 it was almost
completely rewritten and updated. I want
to emphasize that this is not the text of the
1969 manuscript; it is a true 1987 publi-
cation. Ultimately, the Entomological So-
ciety of America agreed to be the publisher.
Congratulations to the authors for writing
it and to the ESA for publishing it at a rea-
sonable cost.
Book
Revision of the Caddisfly Genus Psilotreta
(Trichoptera: Odontoceridae). By C. R.
Parker and G. B. Wiggins. Royal Ontario
Museum, Life Sciences Contribution
Number 144. 1987, v + 55 pp., 57 figs.,
1 table. $12.00 (CDN), from Royal On-
tario Museum, Publication Services, 100
Queen’s Park, Toronto MSS 2C6, Cana-
da.
The family Odontoceridae is a compar-
atively small family of caddisflies that con-
tains representatives on all continents ex-
cept the African. Their larvae, which live
in lotic habitats, construct rigid cases made
of small sand grains. In North America we
recognize 6 genera of which only Psilotreta
occurs in the east. In addition to the 6 species
which the authors recognize in this genus
from North America, another 14 are known
from Asia.
In this revision the genus is characterized
in its adult (including male and female gen-
italia), larval and pupal stages. Males, fe-
males and larvae of all North American
species are separated by key, and they are
grouped into 2 species groups: the indecisa
group with indecisa (Walker), frontalis Banks
and /abida Ross, and the rufa group with
rufa (Hagen), rossi Wallace and amera
(Ross). The usual full synonymy, descrip-
tions, distribution and complete illustra-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
James E. Keirans, Department of Health
and Human Services, Public Health Service,
National Institutes of Health, National In-
stitute of Allergy and Infectious Diseases,
Department of Entomology, Museum Sup-
port Center, Smithsonian Institution,
Washington, D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
90(3), 1988, p. 400
REVIEW
tions are given for each species. The Asian
species are less fully treated; males of only
10, females of 6 and larvae of 3 species being
known. Two new species are described from
Assam and Sikkim, India. The paper fin-
ishes with a cladistic and biogeographic
analysis of the American species.
Considering the overall exhaustive cov-
erage, I find it surprising that the authors
did not illustrate more of the Asian species.
The male genitalia are figured for only 5 of
the 14 species, but illustrations exist for 7
more species and the types of many are
readily available. Even if only the best avail-
able illustrations were copied photograph-
ically for these species, their identification
would be made much easier than having to
look them up in a number of journals, some
rather obscure.
This consideration aside, their work is well
done, with the printing and illustrations clear
and crisp. Now for the first time, it is pos-
sible for workers to identify all the adults
and larvae of this important component of
the North American benthos. This study is
a necessity for all Trichopterists and those
that need to identify benthic collections from
Eastern North America.
Oliver S. Flint, Jr., Department of Ento-
mology, Smithsonian Institution, Washing-
ton, DC 20560.
PUBLICATIONS FOR SALE BY THE
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The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939.
A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
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The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
253 pp. 1949
A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 19522000.
A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp.
The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
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Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
The North American Predaceous Midges of the Genus Pa/pomyia Meigen (Diptera: Cera-
topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979
The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200
Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982...
A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp.
The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
AEST LOS hs te ee eS Sa Ne eS ee ee ee Sh
An Identification Manual for the North American Genera of the Family Braconidae (Hy-
menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987
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the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
WHARTON, R. A. and D. L. J. QUICKE—A new species of Bracon (Hymenoptera: Braconidae)
parasitic on Eoreuma loftini (Dyar) (Lepidoptera: Pyralidae) ........................-. 288
NOTES
GAGNE, R. J.—A polynomial Riley name in Cecidomyiidae (Diptera) and implications of such
names,for Cynipidae (Hymenoptera) <2 2.25. se eee eee ee eee ee Oe 394
PLAKIDAS, J. D.—The newly discovered spring crown gall of Asphondylia rudbeckiaeconspicua
(Diptera: Cecidomyiidae) on Rudbeckia laciniata (Asteraceae) in Pennsylvania .......... 393
BOOK REVIEWS
CARROLIE; J. F: The Behavioural Ecology of ANts enki ds inch coho oa ee oe ats 396
FLINT, O. S., JR. Revision of the Caddisfly Genus Psilotreta (Trichoptera: Odontoceridae) ... 400
KEIRANS, J. E. The Ixodid Ticks of, Uganda © 80.55.28 washes oe tae Sa ee gree eee 398
KINGSOLVER, J. M. The Metallic Wood-boring Beetles of Canada and Alaska: Coleoptera:
Ruprestidae. 3". sos Fes oe Fe Ree Bae 2 i BIN NOTE ee Te Ce ee ORs ee ate ae 397
é VOL. 90 OCTOBER 1988 NO. 4
> | (ISSN 0013-8797)
PROCEEDINGS |
me
of the ERA
ENTOMOLOGICAL SOCIETY
« WASHINGTON
PUBLISHED
QUARTERLY
BARNES, J. K.—Notes on the biology and immature stages of Poecilographa decora (Loew)
WODPTECTE DEW CES ) Sik coal SARA cape Orn Shea ie el in nc ae I 474
CARROLL, J. F.— Worker size and piracy in foraging ants .......................0200.0.. 495
CLEMENT, S. L., T. MIMMOCCHI, R. SOBHIAN, and P. H. DUNN—Host specificity of
adult Eustenopus hirtus (Waltl) (Coleoptera: Curculionidae), a potential biological control
agent of yellow starthistle, Centaurea solstitialis L. (Asteraceae, Cardueae) .............. 501
DIATLOFF, G. and W. A. PALMER—The host specificity and biology of Aristotelia ivae Busck
(Gelechiidae) and Lorita baccharivora Pogue (Tortricidae), two microlepidoptera selected
as biological control agents for Baccharis halimifolia (Asteraceae) in Australia .......... 458
LANE, M.A., F.E. KURCZEWSKI, and R. B. HANNA—Antennal sensilla and setae of Evagetes
PALM AyIncnoniced:, F OMIpIGaG he hrs, Melee cay aati ick Ayla Ni, See ci ee Ca oa a 428
NAKAHARA, S.—Generic reassignments of North American species currently assigned to the
genus Sericothrips Haliday (Thysanoptera: Thripidae) ..........................0.0.-- 480
NORTON, R. A., W. C. WELBOURN, and R. D. CAVE—First records of Erythraeidae parasitic
on oribatid mites (Acari, Prostigmata: Acari, Oribatida) ......................2....-.. 407
POGUE, M. G.—Revision of the genus Lorita Busck (Lepidoptera: Tortricidae: Cochylini), with
BIA ESC nNOnOl AMeWiSDEGIES: 1745. yee Ne aitee ERE a eee gh HoT Ge eS 440
ROBINSON, W. H. and N. BAO—The pest status of Periplaneta fuliginosa (Serville) (Dictyop-
Fer ASO AeH He ttn CITIAN hetel toe Sc 2 EME ener Poke Swe eel A. oh ph eS) Fares aE 401
STRICKMAN, D.—Redescription of the holotype of Culex (Culex) peus Speiser and taxonomy
of Culex (Culex) stigmatosoma Dyar and Thriambus Dyar (Diptera: Culicidae) ......... 484
TRIPLETT, D. C. and A. R. GITTINS—Nesting, mating and foraging habits of Melissodes
(Melissodes) tepida tepida Cresson in Idaho (Hymenoptera: Anthophoridae) ............ 462
VAN DEN BUSSCHE, R. A., M. R. WILLIG, R. K. CHESSER, and R. B. WAIDE—Genetic
variation and systematics of four taxa of neotropical walking sticks (Phasmatodea: Phas-
IE TUIE CS) efi Meg ll @ As AS er ei ete o> Fue eg rhe ee aN ce ee ee AE 422
WHITFIELD, J. B.—Taxonomic notes on Rhyssalini and Rhysipolini (Hymenoptera: Bracon-
idae) with frst Nearctic Tecords, OF three PENETawe es. fe tl ele. «Ge ee ot Wewte Sele eel 471
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
OFFICERS FOR 1988
F. EUGENE Woon, President WARREN E. STEINER, JR. Program Chairman
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‘
Pele i
j 90(4), 1988, pp. 401-406
a
THE PEST STATUS OF PERIPLANETA FULIGINOSA (SERVILLE)
(DICTYOPTERA: BLATTIDAE) IN CHINA
WILLIAM H. ROBINSON AND NONGGANG BAO
Urban Pest Control Research Center, Department of Entomology, Virginia Polytechnic
Institute and State University, Blacksburg, Virginia.
Abstract.—There are approximately 19 pest cockroach species in China. The most
common household pests include the German cockroach, Blattella germanica (L.) and
six Periplaneta species, including the smokybrown cockroach, P. fuliginosa (Serville). The
pest status of P. fuliginosa in residences was determined by evaluating the attitudes and
knowledge of apartment residents in Hangzhou, China that regularly experience this
cockroach as a household pest. The questionnaire-survey indicated that a mean of 5
cockroaches was seen per day in the apartments. However, the residents considered the
presence of 2 per day a problem. A majority of residents considered cockroaches more
serious than a leaky faucet or a broken window, but considered having mice more serious
than having cockroaches. Approximately 8% of the people questioned linked household
cockroaches to the spread of disease. The apartment residents reported spending an average
of 0.38¥ ($0.11) (range 0-1.5¥) per month on cockroach control. Those interviewed in
April were willing to spend an average of 2.65¥ ($0.72) (range 0-5.5¥), and those inter-
viewed in August were willing to spend 5.27¥ ($1.44) (range 0-7.5¥) per month for
elimination of their cockroach infestation. The attitudes and knowledge of cockroach pests
by urban residents in China was similar to those in the U.S.
Key Words:
There are approximately 19 cockroach
species that are pests in households in the
People’s Republic of China (Table 1). The
major pest species are Blattella germanica
(L.), and six Periplaneta species: P. ameri-
cana (L.), P. australasiae (F.), P. brunnea
(Burmeister), P. fallax (Bienko), P. fuligi-
nosa (Serville), and P. japonica Karny. The
German cockroach, B. germanica, occurs
primarily in locations that are regularly
heated in the winter, such as hotels, restau-
rants, and transportation vehicles (Woo and
Guo 1984). Periplaneta americana and P.
fuliginosa are more generally distributed,
and occur in single- and multi-family dwell-
ings, hotels, restaurants, stores, and food
processing plants (Woo 1982). Periplaneta
cockroach pest status, China, survey
Japonica is a serious household pest in
northeastern China (Woo 1982). The ori-
ental cockroach, Blatta orientalis L., has
been recorded as a pest in Beijing, Xingjiang,
and Yunnan provinces (Woo 1987). Pyc-
noscelus surinamensis (L.) has been report-
ed from Yunan, Guangdong, and Fujian
provinces (Woo 1987). Eupolyphaga sinen-
sis Walker is a household pest in northern,
central, and southeastern provinces (Woo
1987). Eupolyphaga sinensis, Polyphaga
plancyi Bolivar, and Opisthoplata orientalis
Burmeister are often used in traditional
Chinese medicine (Woo 1987).
The urban population of the People’s Re-
public of China is approximately 230 mil-
lion. Although urban residents are only 22%
402
Table 1. The cockroaches species that are known
to be household pests in China.
Reference
Species
Family Blattidae
Periplaneta americana (L.) Woo 1982
Periplaneta australasiae
(F.) Woo 1982
Periplaneta brunnea
(Burmeister) Woo 1982
Periplaneta fallax (Bienko) | Woo 1982
Periplaneta fuliginosa
(Serv.) Woo 1982
Periplaneta japonica Karny Woo 1982
Blatta orientalis L. Woo 1987
Hebardina concinna (Haan) Chen et al. 1986
Neostylopyga rhombifolia
(Stoll) Woo 1987
Family Blaberidae
Pycnoscelus surinamensis
(L.) Woo 1987
Family Blattellidae
Blattella germanica (L.)
Blattella latistriga (Walker)
Blattella lituricollis (Walker)
Parcoblatta kyotensis
Woo and Guo 1984
Woo and Guo 1984
Woo and Guo 1984
Asahina Woo 1987
Family Corydiidae
Eupolyphaga yunnanensis
Kirby Woo 1981
Eupolyphaga sinensis
Walker Woo 1987
Eupolyphaga thibetana
Chopard Woo 1981
Polyphaga plancyi Bolivar Woo 1987
Family Epilampridae
Opisthoplata orientalis
Burmeister Woo 1987
of the total population, this segment of so-
ciety lives in more confined and crowded
conditions than the rural population, and is
often exposed to vertebrate and inverte-
brate pests. Information to homemakers on
the biology and control of pests in urban
areas 1s provided primarily by the Sanita-
tion and Anti-epidemic Stations in each of
the provinces and major cities.
Data on cockroach species biology and
habits, distribution, aspects of chemical and
non-chemical control, disease transmission,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and insecticide resistance in China are
available (Woo 1982, 1987, Woo and Guo
1984, Li and Nie 1984). However, there is
little information on the attitudes and
knowledge of urban residents toward cock-
roach pests. These data are important for
the design and implementation of urban pest
management programs (Zungoli and Rob-
inson 1984).
The objective of the research presented
here was to determine the pest status of the
smokybrown cockroach, P. fuliginosa, by
evaluating the attitudes and knowledge of
residents that regularly experienced this
cockroach as a household pest.
MATERIALS AND METHODS
The study was conducted in April and
August 1985, in the city of Hangzhou, Zhe-
jiang Province, China. The survey (Table 2)
consisted of 14 questions; of which nearly
all were open ended, i.e. responses were not
chosen from a list offered by the interview-
er. The topics covered in the questions in-
cluded, what causes cockroaches, what is
the best method of control, the number of
cockroaches seen daily, where in the apart-
ment cockroaches are seen, the amount of
money spent on cockroach control, and the
amount the residents were willing to spend
on control. The age of the person inter-
viewed and the number of years living in
the apartment were also recorded.
The survey method consisted of one in-
terviewer questioning individual residents
in their apartments. Five interviewers con-
ducted the surveys. A total of 105 people
were interviewed: 51 in April, when cock-
roaches were not active outside or inside
the apartments; and 54 in August, when
cockroaches were active inside and outside
the apartments.
The survey sites were apartment build-
ings adjacent to Zhejiang Agricultural Uni-
versity. The apartment buildings were con-
structed within the last five years. The
residents were primarily workers from near-
by factories. The monthly earning per fam-
VOLUME 90, NUMBER 4
Table 2. The questions in the interviews conducted
in Hangzhou, China to determine the pest status of
Periplaneta fuliginosa.
General Opinion of the Cockroach Problem.
Q. Are roaches a serious problem?
A. Yes—67% (April), 94% (August); No—33%
(April), 6% (August).
How many roaches do you see per day (when
you have them as a pest)?
¥ 4.5 (Apmil), * 5 (August).
Where do you see roaches in your apartment?
Bedroom — 63%; Bathroom — 54%; Kitchen—
98%
POP
Pest Status of P. fuliginosa.
Q. Which of these do you think is a worse prob-
lem?
A. April August
Roaches 65% 93%
Faucet leak 25% 6%
Equal 10% 2%
Roaches 63% 67%
Broken window 31% 30%
Equal 6% 4%
Roaches 43% 52%
Trash in hall 57% 44%
Equal _ 4%
Roaches 33% 4%
Mice 63% 87%
Equal 4% 9%
Q. What bothers you the most about having
cockroaches?
Found around food— 53%
Found everywhere— 10%
Cockroach feces— 10%
Spread disease — 8%
Smell— 8%
Other— 12%
Q. If you are visiting someone’s apartment and
you see 20 (15, 10, 5, 2) roaches, would you
think there was a problem with cockroaches?
A. 20 roaches considered a problem—97% yes
15 roaches considered a problem—97% yes
10 roaches considered a problem—97% yes
5 roaches considered a problem—88% yes
2 roaches considered a problem—59% yes
Knowledge of the Cause and Control of Cockroach-
es.
Q. What do you think causes roaches?
A. Food and filth— 43%
Don’t know— 28%
Fly into apt.— 10%
Other— 19%
403
Table 2. Continued.
Q. What is the best way to control roaches?
A
Insecticides— 48%
Sanitation— 29%
Don’t know— 14%
Other— 9%
Economic Impact of Cockroaches.
Q. Did you purchase insecticides to control
roaches?
Yes—87%, No—13%
How much did you spend on roach control
last year (summer)?
X spent—O.38p¥
How much are you willing to spend for elimi-
nation?
X 2.65¥ (April)
X¥ 5.27¥ (August)
> OF OP
ily (3-4 persons) was estimated to be be-
tween 150¥ and 300Y.
All data were coded and analyses were
performed using Statistical Analysis Sys-
tems programs (SAS Institute 1985). Statis-
tical procedures included analysis of vari-
ance, linear regression, and Chi-square
contingency analysis. For all analyses, an
alpha level of significance was set at 0.05.
RESULTS AND DISCUSSION
The survey questions are not presented
in the sequence they occurred on the survey
form, but are grouped to provide easier dis-
cussion of the results. Three questions were
not evaluated; they pertained to the smell
of insecticides, the rating of cockroach con-
trol, and the prospect of eliminating all
cockroach pests.
Background information. —The mean
number of years the residents lived in their
apartment was 4.5 (median 3 yrs). The
number of years of residence coincides with
the age of the apartment buildings; most of
the people surveyed were the first occupanis
of the apartments in the buildings.
General opinion of the cockroach prob-
lem.—The presence of P. fuliginosa in the
apartments was considered a serious prob-
lem by the majority of the people ques-
404
tioned in either April (x? = 5.78; df = 1; P
< 0.025) or August (x? = 38.72; df = 1; P
< 0.005). In August, a larger percentage of
residents (94%) considered cockroaches a
serious problem than residents questioned
in April (67%). Residents questioned in April
reported they had a mean of 4.5 cockroach-
es per day in their apartment in the summer;
and residents questioned in August reported
a mean of 5.0 cockroaches per day in their
apartment. Although residents were able to
recall the approximate number of cock-
roaches in their apartments during the sum-
mer, their perception of the seriousness of
the problem was influenced by whether
cockroaches were present at the time or not.
Residents questioned during the time cock-
roaches were actually present in their apart-
ment (August) indicated they were a serious
problem more often than when questioned
during the time cockroaches were absent
(April). Thoms and Robinson (1986) re-
ported that observations by apartment res-
idents on the distribution and seasonal
abundance of domiciliary cockroaches can
be accurate.
The reason residents consider the pres-
ence of approximately 5 cockroaches per
day to be a serious problem may be due to
the distribution of the cockroaches in the
apartment. Cockroaches were reported
throughout the apartments; 98% of the re-
spondents reported cockroaches in the
kitchen, and 63% reported cockroaches in
the bedroom. Five cockroaches in an apart-
ment may be considered few when com-
pared to German cockroach infestations of
similar structures (Akers and Robinson
1981). However, P. fuliginosa is the primary
household cockroach pest in Hangzhou
(Bao, unpublished data), and the residents
have little or no experience with cockroach
species that have a small body size or occur
in large numbers.
Pest status of P. fuliginosa. Several situ-
ations, such as trash in the hall, or other
pests, such as mice, were compared with a
cockroach infestation. The ‘‘equal’’ re-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
sponses were not more than 5% of the total
responses, and these responses were exclud-
ed from the statistical analysis. A majority
of residents considered cockroaches much
more serious than a leaky faucet, or a broken
window (April x? = 3.38; df= 1; P < 0.025;
August x? = 5.78; df = 1; P < 0.025). Re-
sponses were nearly evenly divided on the
comparison of cockroaches and the pres-
ence of trash in the hall. Sixty-three percent
of the residents questioned in April consid-
ered mice more serious than cockroaches,
and 87% of those questioned in August con-
sidered mice more serious.
Fifty-three percent of the people ques-
tioned reported being bothered by the pres-
ence of cockroaches around food, and some
respondents (8%) linked cockroaches to the
spread of disease. When responding to a
question on distribution, 98% reported
cockroaches in their kitchens. The response
“they are everywhere” may indicate the dif-
ficulty people have in expressing what they
disliked about cockroaches (Wood et al.
1981). The fecal pellets of Periplaneta species
are large (2.5—4.5 mm), and can be scattered
in cabinets and storage areas. Residents
(10%) reported disliking the presence of P.
fuliginosa fecal pellets in their apartments.
Fifty-nine percent of the people ques-
tioned considered the presence of just two
cockroaches (in the hypothetical situation)
to represent “‘a problem.” The attitudes in-
dicated in the questions regarding the num-
ber of cockroaches that constitute “‘a prob-
lem” indicate that pest control may not be
considered successful unless the number of
cockroaches seen per day is at least less than
two. When there are severe infestations,
maintaining low numbers of cockroaches
may be difficult. Robinson and Zungoli
(1985) suggested that the expectations of ur-
ban residents for cockroach elimination may
need to be tempered by pest control per-
sonnel and education programs.
Knowledge of the cause and control of
cockroaches. — When asked about the cause
of cockroach infestations, a large majority
VOLUME 90, NUMBER 4
(43%) of the residents considered food scraps
and filth to be the most important (x° =
23.76; df = 3; P < 0.005). Periplaneta fu-
liginosa is capable of flying short distances,
and 10% of the residents reported cock-
roaches flying into their apartments from
the outside as the cause of their apartments
being infested. Nearly one-third (28%) of
the residents did not know the cause of
cockroach infestations. When asked what
was the best way to control cockroaches,
only 29% of the residents mentioned sani-
tation, whereas 48% considered insecticides
the best method of control.
Economic impact of cockroaches. —Evi-
dence of the importance of the cockroach
infestations was found in the amount of
money the residents reported spending for
control, and on the amount they were will-
ing to spend for elimination. Residents re-
ported spending 0.38¥ ($0.11) (range O-
1.5¥) per summer for cockroach control,
but they were willing to spend considerably
more, from 2.65¥ ($0.72) (range 0-5.5Y)
for those questioned in April to 5.27¥
($1.44) (range 0-7.5¥) for those questioned
in August for elimination of the problem.
The 0.38¥ per summer probably represents
the purchase of several packets of cockroach
control tablets (boric acid) for the four
months that P. fuliginosa are present in
apartments. The amount of money the res-
idents reported they were willing to spend
for cockroach elimination was considerable,
and varied according to when they were
questioned (April or August). At the time
(August) cockroaches were most common
in their apartments, the people questioned
were willing to spend nearly twice the
amount of money for control than what they
stated they were willing to pay for control
when cockroaches were not present. Sawyer
and Casagrande (1983) stated that the se-
verity of a pest problem can be determined
by assessing the amount of money a person
is willing to spend to alleviate the problem.
The results of the survey of the apartment
residents in Hangzhou provide considerable
405
information on the pest status of P. fuligi-
nosa. Although the time this cockroach
species 1s active inside apartments 1s limited
to approximately one-third of the year, the
residents considered it a serious household
pest. The mean number of cockroaches the
residents reported seeing was only five. The
residents reported that the presence of just
two cockroaches would be considered a
problem. However, the apartments are small
and there is considerable opportunity for
interaction between cockroaches and peo-
ple. Thoms and Robinson (1986) reported
urban apartment residents intolerant of B.
orientalis when found indoors. They re-
ported that 82% of the apartment residents
questioned in Roanoke, VA considered two
oriental cockroaches indoors a problem, and
96% considered five a problem.
Thoms and Robinson (1986) reported that
the low tolerance of oriental cockroaches
reported by urban apartment residents may
result from the size of the oriental as com-
pared with the much smaller German cock-
roach, and the perception that the oriental
cockroach is an invader from outdoor hab-
itats. The residents in this study had little
or no experience with infestations of small-
sized cockroaches, such as the German
cockroach, because this species is not a
household cockroach pest in Hangzhou
(Bao, unpublished data).
The response of the residents of Hang-
zhou to the questions comparing cock-
roaches to other household pests (mice), and
to household problems (leaky faucet) was
very similar to the responses by U.S. resi-
dents to similar questions. Wood et al.
(1981) reported that urban apartment resi-
dents considered German cockroaches much
more important than a broken window,
leaky faucet, or trash in the yard, and they
were nearly evenly divided in their opinions
of cockroaches and mice. In the results re-
ported here, the responses of the Hangzhou
residents were nearly the same as those re-
ported in the United States.
This information collected in this survey
406
can provide a basis for the education com-
ponent of a cockroach pest control program
(Robinson and Zungoli 1985). The re-
sponses to the questions concerning the
causes and control of P. fuliginosa infesta-
tions indicate a need for some specific in-
formation. The dependence on chemical
control for cockroaches is apparent by the
large percentage (48%) of the residents that
considered insecticides to be the best way
to control these pests. Robinson and Zun-
goli (1985) reported a significant change in
resident understanding of cause and control
of household cockroaches as a result of an
education program.
ACKNOWLEDGMENT
We appreciate the help of Wan Xing-
sheng, Zhu Kunyan, Chen Yi, and Zou
Wanhe in conducting the interviews of the
residents. Patricia A. Zungoli, Department
of Entomology, Clemson University, re-
viewed and improved the manuscript.
LITERATURE CITED
Akers, R. C.and W. Robinson. 1981. Spatial patterns
and movement of German cockroaches in urban
low-income apartments (Dictyoptera: Blattelli-
dae). Proc. Entomol. Soc. Wash. 83: 168-172.
Chen, P., C. Huang, P. Su, and J. Huang. 1986. Re-
port on cockroache species, biology and habits in
Nanning, Guangxi, pp. 16-24. /n Z. Huang, ed.,
Studies on Household Cockroaches and Bedbugs.
Series 3., Medical Animal Pest Control 2(4).
Li, Jie and Nie Wei-qing. 1984. Bionomics of Te-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
trastichus hagenowii parasitizing in the ootheca of
Periplaneta fuliginosa. Acta Entomologica Sinica
27: 406-409.
Robinson, W. H.and P. A. Zungoli. 1985. Integrated
control program for German cockroaches (Dic-
tyoptera: Blattelidae) in multiple-unit dwellings. J.
Econ. Entomol. 78: 595-598.
Sawyer, A. J. and R. A. Casagrande. 1983. Urban
pest management: A conceptual framework. Ur-
ban Ecology 7: 145-157.
SAS Institute. 1985. SAS user’s guide: Statistics. SAS
Institute, Cary, N.C.
Thoms, E. M. and W. H. Robinson. 1986. Distri-
bution, seasonal abundance, and pest status of the
oriental cockroach (Dictyoptera: Blattidae) and an
evaniid wasp (Hymenoptera: Evaniidae) in urban
apartments. J. Econ. Entomol. 79: 431-436.
Woo, Fo-ching. 1981. Blattoptera: Corydidae, Blat-
tidae, Panesthiidae. Insects of Xizang (Tibet). 1:
57-61.
1982. Species of the genus Periplaneta Bur-
meister from China, with reference to their bio-
nomics and economic importance (Blattaria: Blat-
tidae). Acta Entomologica Sinica 25: 416-422.
1987. Investigations on domicilary cock-
roaches from China. Acta Entomologica Sinica.
30: 430-438.
Woo, Fo-ching and Yu-yuan Guo. 1984. The specific
identification, distribution, bionomics, and eco-
nomic importance of the genus Blattella Caudell
(Blattaria: Blattidae) from China. Acta Entomo-
logica Sinica 27: 439-443.
Wood, F. E., W. H. Robinson, S. K. Kraft, and P. A.
Zungoli. 1981. Survey of attitudes and knowl-
edge of public housing residents toward cock-
roaches. Bull. Entomol. Soc. Amer. 27: 9-13.
Zungoli, P. A.and W. H. Robinson. 1984. Feasibility
of establishing an aesthetic injury level for German
cockroach pest management programs. Environ.
Entomol. 13: 453-458.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 407-410
FIRST RECORDS OF ERYTHRAEIDAE PARASITIC ON
ORIBATID MITES (ACARI, PROSTIGMATA: ACARI, ORIBATIDA)
Roy A. Norton, W. CALVIN WELBOURN, AND RONALD D. CAVE
(RAN) Faculty of Environmental and Forest Biology, S.U.N.Y. College of Environ-
mental Science & Forestry, Syracuse, New York 13210, and Research Associate, Field
Museum of Natural History, Chicago, Illinois 60605; (WCW) Acarology Laboratory, The
Ohio State University, 484 W. 12th Ave., Columbus, Ohio 43210; (RDC) Departamento
de Proteccion Vegetal, Escuela Agricola Panamericana, P.O. Box 93, Tegucigalpa, Hon-
duras.
Abstract. —Larvae of two undescribed species of the mite genus Leptus (Erythraeidae)
were found attached to the heavily sclerotized cuticle of ten oribatid mites, representing
four species and three families. Parasitized mites, each carrying a single Leptus larva, were
collected from forest soils in Massachusetts, Mississippi, and Alabama. This is the first
reported association between these taxa, but the specificity of the relationship is unknown.
Key Words:
Oribatid mites, most of which are sa-
prophagous or fungivorous inhabitants of
soil organic horizons, commonly serve as
hosts for a variety of parasitic and com-
mensal organisms. From the early work of
Nicolet (1855) to the detailed studies of K.
Purrini and colleagues (e.g. Purrini 1980),
we have learned that these mites are para-
sitized by bacteria, fungi, virus-like organ-
isms, various protozoans (amoebae, eugreg-
arines, microsporidians, helicosporidians),
and nematodes. Their servitude as inter-
mediate hosts for anoplocephalid tape-
worms is perhaps the most widely known
parasitic relationship (see review by Seng-
busch 1977). Records of commensal asso-
ciations are fewer. Phoretic transport of
nematodes (e.g. Travé 1956) or deuto-
nymphs of the mite suborder Astigmata is
not uncommon (unpublished obervations,
R.A.N.), but is rarely reported. Ciliate pro-
tozoans, apparently related to Conidi-
ophryidae and similar to those carried by
soil-dwelling mesostigmatic mites (Dindal
Leptus, ectoparasites, oribatid mites
1973), are also commonly attached to leg
or body setae of oribatid mites (unpublished
observations, R.A.N.); they are probably
commensals.
To this list of relationships we can now
add ectoparasitism by mites of the family
Erythraeidae. These mites are protelean ec-
toparasites whose larvae utilize a wide va-
riety of insect and arachnid hosts (Welbourn
1983 and included references), and whose
deutonymphs and adults are free-living
predators. Their red color and wide host
range make them the most obvious of ter-
restrial protelean parasites.
METHODS AND RESULTS
From three different localities we ob-
served multiple cases of parasitism by lar-
vae of the erythraeid genus Leptus on adult
oribatid mites which had been extracted by
Berlese funnels from forest soil and litter.
Each of four specimens of Oribatella exten-
sa Jacot (Oribatellidae) from Mississippi
(kudzu litter, Ecru, Pontotoc Co., 18-III-
408
1981, R. L. Brown, col.) carried a single
Leptus larva attached dorsally on the pos-
terior half of its notogaster (Figs. 1, 2). Four
specimens of Damaeus verticillipes (Nico-
let) (Damaeidae) from Massachusetts (white
pine plantation litter, Babson College cam-
pus, Wellesley, Norfolk Co., 6-XI-1985, J.
R. Philips, col.) also carried a single Leptus
each. In three cases the larva was attached
to the notogaster, behind the stacked exu-
vial scalps which this species usually carries
(Figs. 3, 4); in one case attachment was on
the ventral plate, immediately posteriad of
the anus, but the general posture of the larva
was as in Fig. 3. In the third collection, from
Alabama (forest litter, Conecuh Co., 0.2 mi.
W of junction of Sepulga River and Rt. I-65,
15-III-1986, R. D. Cave, col.), one speci-
men each of Xenillus occultus Banks (Xenil-
lidae) and Damaeus grossmani Wilson (Da-
maeidae) carried a Leptus larva. In the
former case, the parasite was positioned near
the notogastral margin, midway along its
left side; in the latter, attachment was sim-
ilar to that of Fig. 1, but on the nght side.
None of the attached Leptus larvae were
engorged when collected.
The host mites from Massachusetts were
observed alive for one week to compare their
behavior with that of non-parasitized in-
dividuals. Despite the relatively large size
and sometimes non-axial positioning of the
parasite (Fig. 1), the oribatid hosts were quite
mobile, and showed no abnormal activity.
Noticeable engorgement did not take place
during this time. Although not observed
alive, the oribatid mite hosts from the other
two collections must have been active
enough to respond to increasing desiccation
and move through the litter column in the
Berlese extractor.
DISCUSSION
These examples illustrate the ability of
Leptus larvae to attach almost anywhere on
an arthropod integument. Whereas other
erythraeid larvae, with short cheliceral bas-
es, tend to attach along molting sutures
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(Young and Welbourn 1987), Leptus larvae
have long, flask-shaped cheliceral bases
(Baker 1982, Southcott 1961, Treat 1975)
which facilitate the penetration of even well
sclerotized cuticle, such as the notogaster of
oribatid mites. Baker (1982) reported a
“viscous substance” that was secreted just
prior to attachment of Leptus larvae, and
suggested that upon hardening it formed a
cement which secured the mite to the host.
It can be seen in Fig. 2, where the gnatho-
soma contacts the oribatid mite cuticle. Al-
though the material seems to play a role in
attachment (no stylostome is formed), it also
forms a seal around the feeding lesion (Abro
1988). In contrast to other erythraeid lar-
vae, the gnathosoma of Leptus species lacks
the buccal fringe or ring which usually per-
forms the latter function (unpublished ob-
servations, W.C.W.).
Systematic studies of Leptus, which in-
cludes nearly 90 named species worldwide,
have concentrated on the larval instar (e.g.
Southcott 1961, Beron 1975, Fain and Elsen
1987, Fain et al. 1987). The American fauna
is still poorly known, and all of the larvae
from the oribatid mites represent unde-
scribed species whose degree of host spec-
ificity is unknown. The Leptus larvae from
Alabama and Mississippi are conspecific,
and their occurrence on representatives of
three oribatid genera, from three distinct
families of Brachypylina, suggests little host
specificity. Yet, no other parasitized arthro-
pods were observed in the Mississippi sam-
ple, despite the presence of unattached Lep-
tus larvae and numerous potential arthropod
hosts (including other taxa of oribatid mites).
Similarly, unattached larvae of the Massa-
chusetts Leptus (representing a second un-
described species) were found in the original
sample, but only individuals of D. verticil-
lipes were parasitized.
Can a Leptus larva complete its devel-
opment to a free-living deutonymph using
a single oribatid mite host? We have no
evidence at present, since no observed spec-
imen had started to engorge. If we speculate
VOLUME 90, NUMBER 4 409
Yr»,
AN
Pon iy
a) 4 z a
Figs. 14. Leptus larvae parasitizing adult oribatid mites. Fig. 1. Oribatella extensa Jacot, from Mississippi
(dorsal aspect), with Leptus sp. larva attached to notogaster. Fig. 2. As in Fig. 1, except dorsolateral aspect. Note
hardened “cementing” material at point of attachment. Fig. 3. Damaeus verticillipes (Nicolet), from Massachu-
setts (posterolateral aspect), with Leptus sp. larva attached to notogaster. Fig. 4. As in Fig. 3, except posterior
aspect. All scanning electron micrographs; scale bars on Figs. | and 3 represent 45 um, those on Figs. 2 and 4
represent 25 um.
410
that it can, the parasitism would probably
be lethal to the host, considering the relative
size of the two mites. This would be con-
sistent with the absence from the samples
of hosts with engorged parasites, since de-
creased mobility in the Berlese funnels would
be expected once engorgement begins; it
would also be consistent with the absence
of oribatid mites having noticeable lesions
resulting from prior parasitism.
ACKNOWLEDGMENTS
Facilities for scanning electron micros-
copy were provided by the N. C. Brown
Center for Ultrastructure Studies, S.U.N.Y.,
C.E.S.F. We are grateful to V. M. Behan-
Pelletier (Biosystematics Research Centre,
Agriculture Canada, Ottawa), J. B. Kethley
(Field Museum of Natural History, Chica-
go), and S. C. Palmer (S.U.N.Y.-C.E.S.F.),
for providing constructive criticism of the
manuscript, and to Dr. J. R. Philips for con-
tributing specimens. Partial support for this
work was provided by a National Science
Foundation grant to R.A.N. (BSR 84 15747).
LITERATURE CITED
Abro, A. 1988. The mode of attachment of mite lar-
vae (Leptus spp.) to harvestment (Opiliones). J.
Natural History 22: 123-130.
Baker, G. T. 1982. Site attachment of a protelean
parasite (Erythraeidae: Leptus sp.). Experientia 38:
923.
Beron, P. 1975. Erythraeidae (Acariformes) larvaires
de Bulgarie. Acta Zool. Bulgarica 1: 45-75.
Dindal, D. L. 1973. Review of soil invertebrate sym-
biosis, pp. 227-256. Jn D. L. Dindal, ed., Proc.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ist Soil Microcommunities Conference. U.S.
Atomic Energy Comm., publ. CONF-711076.
Fain, A. and P. Elsen. 1987. Observations sur les
larves du genre Leptus Latreille, 1795 (Acari: Er-
ythraeidae) d’Afrique centrale. Rev. Zool. Afr. 101:
103-140.
Fain, A., G. L. Gummer, and J. O. Whittaker, Jr.
1987. Two new species of Leptus Latreille, 1796
from the USA. Intern. J. Acarology 13: 135-140.
Nicolet, M. H. 1855. Histoire naturelle des acariens
qui se trouvent aux environs de Paris (premiére
partie). Arch. Mus. Hist. Natur. 7: 381-482.
Purrini, K. 1980. On the incidence and distribution
of parasites of soil fauna of mixed coniferous for-
ests, mixed leaf forests, and pure beech forests of
lower Saxony, West Germany, pp. 561-582. In
D. L. Dindal, ed., Soil Biology as Related to Land
Use Practices. U.S. Environ. Protection Agency
Publ. no. EPA-560/13-80-038.
Sengbusch, H. G. 1977. Review of oribatid mite-
anoplocephalan tapeworm relationships (Acari;
Oribatei: Cestoda; Anoplocephalidae), pp. 87-102.
In D. L. Dindal, ed., Biology of Orbatid Mites.
S.U.N.Y. College of Environ. Sci. & Forestry, Syr-
acuse.
Southcott, R. V. 1961. Studies on the systematics
and biology of the Erythraeoidea (Acarina), with
a critical revision of the genera and subfamilies.
Australian J. Zool. 9: 367-610.
Travé, J. 1956. Le nématode phorétique Cheilobus
quadrilabiatus Cobb sur des oribates (acariens).
Vie et Milieu 7: 110-112.
Treat, A. E. 1975. Mites of moths and butterflies.
Cornell University Press, Ithaca. 362 pp.
Welbourn, W. C. 1983. Potential use of trombidioid
and erythraeoid mites as biological control agents
of insect pests. Univ. Calif. Spec. Publ. 3303. Pp.
103-140.
Young, O. P. and W. C. Welbourn. 1987. Biology of
Lasioerythraeus johnstoni (Acari: Erythraeidae)
ectoparasitic and predacious on the tarnished plant
bug, Lygus lineolaris (Hemiptera: Miridae), and
other arthropods. Ann. Entomol. Soc. Amer. 80:
243-250.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 411-421
NOTES AND REDESCRIPTIONS OF SOME ANOPHELES SERIES
ARRIBALZAGIA HOLOTYPES (DIPTERA: CULICIDAE) IN
THE BRITISH MUSEUM (NATURAL HISTORY)
RICHARD C. WILKERSON
Department of Entomology, Walter Reed Army Institute of Research. Mailing address:
Walter Reed Biosystematics Unit, Museum Support Center, Smithsonian Institution,
Washington, D.C. 20560.
Abstract. — Notes and redescriptions are given for four holotypes of Anopheles (Anoph-
eles) Series Arribalzagia in the British Museum (Natural History). Anopheles mediopunc-
tatus, An. maculipes and An. venezuelae (=punctimacula) are described and illustrated
and notes given for 4n. amazonicus (=mattogrossensis). Dissection of male genitalia of
An. mediopunctatus shows that this species is probably commonly misidentified through-
out South America.
KeyWords: Anopheles, Arribalzagia, mediopunctatus, maculipes, venezuelae, punctimacu-
la, amazonicus, MattOgrossensis, taxonomy
Series Arribalzagia (Theobald 1903, as
genus) 1s a neotropical group of Anopheles
(Anopheles) mosquitoes containing 33
nominal species, among which are known
or suspected vectors of malaria parasites. In
the only objective attempt to define this
group Reid and Knight (1961) listed 19 val-
id names. Two additional species, 47. ve-
ruslanei Vargas and An. anchietai Correa
and Ramalho, have been added since by
their respective authors. Reid and Knight
considered all New World Anopheles
(Anopheles) with laticorn pupal trumpets to
belong in the Arribalzagia Series.
The following is part of a review of Series
Arribalzagia. This review demands revali-
dation of present species concepts. Below
are redescriptions and/or comments on the
four nominal species present in the British
Museum (Natural History). Two of these,
An. maculipes (Theobald) and An. medi-
opunctatus (Theobald) are valid species and
are described and illustrated in full. The
other two, An. venezuelae Evans and An.
amazonicus Christophers are junior syn-
onyms of An. punctimacula Dyar and Knab
and An. mattogrossensis Lutz and Neiva re-
spectively. Since both are adequately de-
scribed in the literature, there is no need for
detailed redescriptions, though venezuelae
is illustrated. Anopheles maculipes and An.
mediopunctatus have clear priority and there
is no doubt as to their status. I have seen
the types of 4n. punctimacula and An. mat-
togrossensis and agree that 4n. venezuelae
and An. amazonicus are indeed junior syn-
onyms.
Anopheles mediopunctatus (Theobald)
(Figs. [A-E, 2A—D)
Theobald 1903, 3: 60-62 (as Cycloleppter-
on).
Diagnosis.—A yellow and brown to dark
brown species with predominantly yellow
palpi and 3 small dark spots on the scutum.
Wings with broad brown and pale yellowish
scales and 3 primary costal dark spots. Legs
412 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Anopheles mediopunctatus (Theobald)
HOLOTYPE
| -
'
2
Sa
&
x
ee ie
os Ah SS
=e Fonte SA ne
a Ee Se Oe ES
, i = a. were
= —% “> * eae, Tm = SS nern ee
<4 ea =
(i mi
, <
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‘ f . | = ¥
| \ fg 8
5 AS ~~
oe she =
| i aX, a xy
| ' S x
ait F #5; hy &
are Aries (Leak
Fig. 1. A-D, holotype Anopheles mediopunctatus (Theobald). A, legs, posterior view. B, habitus. C, head
lateral view. D, thorax, lateral view. E, venter of abdomen. All scale lines in mm.
VOLUME 90, NUMBER 4 413
oe
\/} KO
h \| / ji i}; \- | Fan\
V; Ye byl |
\Wy, 4
\ H\\\" W477 Z|
x
J right side right side
f/f J lateral aspect mesal aspect
if 1 ae
7
7 J
Anopheles mediopunctatus (Theobald)
HOLOTYPE
Fig. 2. A-D, holotype Anopheles mediopunctatus (Theobald) male genitalia. A, claspette, lateral view. B,
claspette, mesal view. C, genitalia before dissection. D, aedeagal leaflet. Scale line in mm.
414
speckled pale yellowish on brown, tarso-
mere 5 of all legs pale. Pleuron with a patch
of scales on the middle of the mesokatepi-
sternum and several scales on the upper
mesanepimeron. Abdomen with postero-
lateral brown and pale yellow scale patches
on terga II-VI.
Label data.—Red circle ““Holotype”’;
handwritten ‘““Cycloleppteron mediopunc-
tatus (Type male) (F.V.T.)”; handwritten
“Sao Paulo Dr. Lutz’; cardboard circle to
which specimen is attached by a small pin
has on its underside some nearly illegible
writing “Santos S---ya 14 VI 02 E---
Carvaye
Also affixed to the pin with the specimen
is a piece of thin glass attached to a piece
of cardboard. Originally this held the male
genitalia which is now mounted on a slide
with the following labels: Left label, hand-
written “male genitalia Cycloleppteron me-
diopunctatus Theobald,” printed “Holo-
type.”’ Right label, handwritten “Brazil: Sao
Paulo Dr. Lutz Santos ?Suanya 14.VI.02
ex carva see pinned collection.” Also a red
circle holotype label. This preparation was
remounted for study.
Also received from the British Museum
was a slide mount of a wing presumably in
Canada balsam. It also has a red circle ho-
lotype label but cannot be part of the aduli
holotype since both wings are present even
though part of one is glued to the cardboard
to which the specimen is pinned. This one
is probably the mount sent by Lutz to Theo-
bald as mentioned in the original descrip-
tion.
Condition of specimen.— Male. Overall
in very good condition, not rubbed or faded.
Left foretarsomere 5 missing. Left wing bro-
ken offat the presector dark spot, the broken
portion glued to the cardboard base. Gen-
italia dissected, slide mounted; abdominal
segments I-VI present. A small pin is in-
serted through sternum I, the tip emerging
posteriorly on the scutum. The genitalia were
mounted in a water soluble medium of un-
known composition. When the media was
dissolved the genitalia preved to be very
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
brittle and the weight of the small coverslip
caused much damage to the gonocoxites.
Further manipulation in order to see the
claspettes also resulted in some damage.
Published here are drawings before re-
mounting and interpretation of the struc-
tures revealed by remounting (Fig. 2A—D).
Description. — Wing length 4.2 mm; width
0.8 mm; proboscis 3.9 mm; Forefemur 2.0
mm.
Head: Integument of postgena, occiput,
vertex, interocular space, pedicels and clyp-
eus dark brown, with a light covering of
silvery pollinosity, more dense and appar-
ent at the vertex. Frontal tuft of long pale
yellowish white setae; tuft continuous onto
the vertex. Five or 6 slender pale yellowish
white appressed, ocular scales present. Ver-
tex and occiput with numerous erect pale
brown spatulate, truncate scales; 8 to 10 long,
pale brown ocular setae present on each side;
about 8 long pale brown postgenal setae and
a few long pale brown labial basal setae pres-
ent. Antennal pedicel with 6 to 7 scattered
slender pale yellowish scales on its outer
surface; 14 antennal flagellomeres present;
flagellomere | with 6 or 7 slender pale yel-
lowish white scales on its mesal and dorsal
surfaces and a few dark brown scales and
setae on its lateral surface; flagellomeres 2-
12 quite plumose, the setae pale yellowish
brown; flagellomeres 13 and 14 elongated,
13 about twice length of 14, with sparser
much shorter setae than remainder of an-
tenna. Palpi dark brown in ground color;
dark scales dark brown, pale scales pale yel-
low but darkening to nearly golden yellow
on palpomere 5; basal scales broad and erect,
the scales of palpomere 5 long slender and
appressed; setae at tip of palpomeres 4 and
5 mostly pale yellow but with some brown
setae mostly at the tip of palpomere 4. Pro-
boscis dark brown, covered with slender
mostly appressed dark brown scales and
short dark setae; scattered pale yellowish
brown scales intermixed on basal half; the
scales at the base broader, more numerous
and erect.
Thorax: Scutal integument with 3 prom-
VOLUME 90, NUMBER 4
inent dark brown spots, 2 on either side just
posterior to the ends of the prescutal sutures
and one at the posterior margin of the scu-
tum which continues onto the scutellum.
Scutum otherwise pale brown to brown in
ground color covered with silvery blue pol-
linosity and scattered slender pale yellowish
to pale brown setae; a few slender scales
present between the scutal angle and the
wing base; denser pale narrow scale-like se-
tae present anteriorly; small darker spots
which lack pollinosity are scattered over the
surface of the scutum corresponding to setal
insertions. Ground color of pleuron brown
to dark brown, the pale areas made so by a
covering of silvery pollinosity, the dark areas
by dark brown pollinosity; antepronotum
with a dense patch of dark brown spatulate
scales anteriorly and a few pale scales pos-
teriorly; antepronotal setae numerous, pale
yellowish brown; upper proepisternum with
3-4 pale setae, below and anterior to these
1-2 small pale scales present; pleural setae
and scales pale yellow and pale yellowish
white, respectively; mesokatepisternum with
upper intermixed scales and setae, a small
median dense patch of scales, and a few
setae below; upper mesanepimeron with a
patch of long setae and just below these 2
broad scales. Forecoxa with an upper an-
terior patch of small yellow scales, inter-
mixed with and continuing below these
scales are about |2 long dark setae; laterally
a small dense patch of white scales below
and a few scattered pale yellow scales above;
posteriorly is a dense patch of long dark
brown scales with a few dark setae inter-
mixed. Foretrochanter mostly with small
appressed yellow scales and a few short pale
setae, the posterior scales dark brown. Mid-
and hindcoxae and trochanters with white
scale patches except for a small patch of pale
yellow scales and pale yellow setae mesally
on the trochanters. Other leg segments as
figured, the dark areas dark brown, the light
areas pale yellow. Tarsomere 5 pale on all
legs. Wing scales mostly broad, the dark
scales mostly dark brown: with 3 main dark
costal spots underlain by dark integument;
415
white scales present on either side of the 3
main dark spots, remainder of wing a mix-
ture of white and pale yellowish scales; pale
yellowish scales predominate on veins R, . ;,
M,.2, M,;, M> and much of CuA. A slight
notch present where costa and subcosta in-
tersect. Humeral crossvein with dark scales
above and below. Halteres dark brown with
white scales on the dorsal aspect of the ped-
icel and around the dorsal margins of the
scooped out capitellum.
Abdomen: Mottled dark brown to pale
brown, with long and abundant, pale yel-
lowish brown setae. Posterolateral margins
of terga II-VI with small patches of broad
dark brown and pale yellowish scales; the
pale scales mostly dorsal to the dark scales:
ventrally with a scattering of quite broad
pale yellowish scales and paired patches of
dark brown scales apically on either side of
the midline of sterna III-V. Segments VII+
not present and the apical portion of ster-
num VI has been disturbed.
Male genitalia (Fig. 2A—D).—Fig. 2C is
the genitalia as observed before dissection.
The dorsal and ventral lobes of the clasp-
ettes were obscured and are illustrated sep-
arately (Fig. 2A, B). Dorsal lobe with 3 mod-
ified setae corresponding to the clubbed setae
of other anophelines; 2 of these are shorter
and rounded at the apex, the third is mod-
ified into a hook-like structure. Apex of ven-
tral lobes with strong sinuous setae which
appear to be in a symmetrical conforma-
tion; just below the apex are 2 smaller but
prominent setae on the mesoventral aspect;
numerous long strong spicules along the
dorsomesal aspect for most of the length of
the lobe. Lobes of tergum IX prominent,
slightly arched outward from each other.
With one pair of aedeagal leaflets, leaflets
with small aciculae on their inner margins
at the base. Gonocoxite with 2 primary se-
tae which may correspond to the parabasals
of other Anopheles (Anopheles), or one para-
basal and one internal seta; the most basal
seta originates on a prominently raised base;
a primary long seta apparently present sub-
apically on the dorsal aspect.
416
Discussion.— Anopheles mediopunctatus
is the oldest name in a group of what may
prove to be several closely related species.
The names presently considered as junior
synonyms of 4n. mediopunctatus are Cy-
clolepidopteron rockefelleri Peryassu, An.
costai Da Fonseca and Da Silva Ramos and
An. costalimai Coutinho. Anopheles bonnei
Da Fonseca and Da Silva Ramos is consid-
ered valid. I cannot speculate on the iden-
tities of the above nominal species without
further study and material. Anopheles me-
diopunctatus and the other or others in this
group all share complex male genitalic
structures which feature very long 9th tergal
lobes, highly modified dorsal and ventral
lobes and unusual placement of the para-
basal and internal spines.
One of the characters commonly used to
distinguish mediopunctatus 1s the presence
of white scales on the first sternum. This
character cannot be seen in the holotype
since the first sternum was destroyed by a
mounting pin.
Examination of approximately 40 male
terminalia from specimens which key to An.
mediopunctatus from many localities in
South America yielded none that are true
mediopunctatus. It now seems likely that
there are many misidentifications of this
species in the literature and in museums.
Anopheles maculipes (Theobald)
(Fig. 3A-E)
Theobald 1903, 3: 81-83 (as Arribalzagia).
Diagnosis.—A brown to dark brown
species with 3 faint dark spots on the scu-
tum, speckled legs and posterolateral ab-
dominal scale tufts. Wing with 3 distinct
costal dark spots and slender dark brown
and pale yellow scales.
Label data.—Red circle “‘Holotype”’;
handwritten ““Anopheles maculipes (Type)
Theobald” and “Sao Paulo Brazil Dr. Lutz.”
Condition of specimen.— Female. In fair
condition, both front legs missing, midtar-
somere 5 missing on left, hindtarsomere 5
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
missing on both sides, right hindtarsomere
4 missing. Abdomen flattened out and
twisted around but intact. Last 4 flagello-
meres of left antenna missing. Specimen is
mounted on a small pin through prosternal
area, emerging in the middle of the scutum;
pin is affixed to a cardboard circle.
Description. — Wing length 4.5 mm; width
1.5 mm; proboscis 2.3 mm; palp 2.19 mm
(e083 25 0'53373;.0:78>-45 0.452 552053):
abdomen approximately 3.35 mm.
Head: Occiput and clypeus with dark
brown integument covered with concolor-
ous pollinosity. Posterior and lateral vertex
with erect slender spatulate and mostly
truncate brown scales; frontal tuft with elon-
gate and some spatulate whitish-yellow
scales and setae; inner margin of eye at the
vertex with small recumbent white scales;
several pale scales on gena below where eyes
meet: postgenal setae brown; 5 to 6 brown
ocular setae per side. Clypeus as figured,
original description says of “peculiar form”
but what Theobald meant is not clear. An-
tennal pedicel dark brown with slender white
scales on dorsolateral third; flagellomere 1
with slender white scales, most on inner sur-
face. Palp clothed with numerous slender
brown erect and semierect spatulate scales
with basal rings of white scales at juncture
of palpomeres 2,3: 3,4 and 4,5; tip of 5 with
a few white scales; 3 with a few white scales
which suggest a pair of ill-defined median
patches. Proboscis with erect brown scales
at its base and recumbent brown scales and
short semierect setae along its length, evi-
dence of a few small pale scales just before
labellum.
Thorax: Thorax somewhat rubbed, integ-
ument yellowish brown to reddish brown,
covered with silvery pollinosity and sparse-
ly clothed with pale yellowish setae. Scutum
with 2 small dark spots just posterior to the
ends of the prescutal sutures and one dark
spot at the posterior margin of the scutum
which continues onto the scutellum; scutum
with scattered, mostly small spots which lack
pollinosity and correspond to the insertion
VOLUME 90, NUMBER 4 417
Anopheles maculipes (Theobald)
HOLOTYPE
Fig. 3. A-E, holotype Anopheles maculipes (Theobald). A, head, lateral view. B, legs, posterior view. C,
habitus. D, thorax, lateral view. E, venter of abdomen. All scale lines in mm.
418
of scattered setae. Antepronotum with a
dense tuft of brown scales anteriorly, the
remainder with approximately 25 long yel-
lowish to brown setae. Mesopostnotum with
a small median posterior dark streak. Pleu-
ron. Proepisternum with 3 long upper setae
and a slender white scale; 4 pale spiracular
setae present; mesokatepisternum with 6
long upper setae and 3 lower setae and 6
lower white scales; prealar knob with 10
long setae; upper mesanepimeron with 4 long
setae. Legs. Forecoxa with an anterior dense
patch of small brown scales and long brown
setae and an apical patch of white and brown
scales. Midcoxa with 2 outer and one inner
white scale patches. Hindcoxa with a single
white scale patch. Mid- and hindfemora and
tibiae brown scaled with speckling of yel-
lowish scale patches. Midtarsomere |
speckled, midtarsomere 2 with a single yel-
lowish spot, the remaining tarsomeres brown
except tip of midtarsomere 5 which is yel-
lowish; tips of hindtarsomeres 1-4 pale
scaled: hindtarsomere | with about 6-9 yel-
lowish white spots some of which form rings;
hindtarsomere 2 with 2-3 spots, one form-
ing a median ring on left side; hindtarso-
meres 3 and 4 brown except for broad white
apices as noted above. Wing scales brown,
dark brown on the presector, middle and
preapical dark marks of costa, the remain-
der yellowish except for some white patches
interspersed. No notch where costa and sub-
costa intersect. Humeral crossvein with
scales above and below. Halteres with a yel-
lowish stem, capitellum dark brown ven-
trally but mostly white-scaled dorsally ex-
cept for the dark bare center of the depressed
area.
Abdomen: Integument of abdomen dark
brown with paler dorsolateral areas; with
covering of yellowish-brown pollinosity.
Dorsum with numerous, sometimes long,
pale yellowish setae, without scales except
for a few pale brown scales posterolaterally
on terga II-VIII, most numerous on terga
VIII where they are narrower and not con-
centrated in patches as the other terga. Ven-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
trally only sterna IV—VIII plainly visible,
with sparse yellowish-brown setae and scat-
tered scales in an irregular pattern as fol-
lows: irregular lines of white scales midven-
trally and small lateral clumps on V-VII;
small patches of brown scales posterolater-
ally on IV-VI; a large tuft of brown scales
midapically on VII; sternum I bare.
Discussion.—This species was the first
member of the nominal taxa Arribalzagia
to be described, Anopheles mediopunctatus,
also an Arribalzagia, was described in the
same publication in a different genus. In his
description of the genus, Theobald (1903:
81) says it 1s closely related to old world
Myzorhynchus. Subsequent speculation
(Reid and Knight 1961) has also suggested
this. Comment on this relationship is be-
yond the scope of the present paper. Theo-
bald in characterizing Arribalzagia says ““No
scaly ventral apical tuft can be detected,”
but this apical tuft can easily be seen on the
type of An. maculipes described here.
Anopheles amazonicus Christophers
Christophers 1923: 71-78, Plate IV.
Anopheles amazonicus is a junior syn-
onym of An. mattogrossensis. The original
description is quite adequate, therefore only
notes and further comments will be pre-
sented here.
Label data.— Pink circle “Type female”;
red circle ‘‘Holotype’’; ‘“‘Holotype of
Anopheles amazonicus Christophers”’;
‘Anopheles (Myzorhynchella) nigra(?)
Theob.”; “A.A. Clark R. Amazon June
1915”: School of Trop. Med. Liverpool
University label ‘““Anopheles amazonicus
Christophers Amazon.”
Condition of specimen. — Female. In fair-
ly good condition except slightly rubbed.
Small pin through mesanepimeron. Right
antenna present only to 4th flagellomere.
About half the fringe scales of the right wing
are missing. Left midleg missing. Tarso-
meres 3-5 on right midleg missing. Right
hindleg missing. Left hindleg with all tar-
someres missing.
VOLUME 90, NUMBER 4 419
Anopheles venezuelae Evans
HOLOTYPE
/— 2—
Sana Ltt) Ney
Fig. 4. A-D, holotype Anopheles venezuelae. A, legs, posterior view. B, habitus. C, thorax, lateral view. D,
head, lateral view. All scale lines in mm.
420
Discussion.—The original description
agrees well with the holotype specimen with
a few small differences or additions as fol-
low. Palpi have a few white scales at junc-
tures of palpomeres 2,3; 3,4 and 4,5. An-
tennal flagellomere | with narrow pale scales
mostly on the inner aspect but some outer
scales also present, no dark scales as stated
in the original description. Upper middle of
mesanepimeron with 3 setae, the upper
mesanepimeron with a patch of long setae.
Abdominal sternum | with a few small pale
setae. The white scales mentioned in the
original description on the venter of the ab-
domen are actually yellowish. The median
tuft of scales on sternum VII has yellow
scales anteriorly and dark brown scales pos-
teriorly.
Anopheles amazonicus was synonymized
with An. mattogrossensis by Shannon (1933:
135) after comparison by Evans of several
specimens of presumed 4n. mattogrossensis
with the type of An. amazonicus. I have also
seen the holotype of An. mattogrossensis and
fully agree. Anopheles mattogrossensis has
several characters unique among the Arri-
balzagia: Upper middle of mesanepimeron
with setae; midventral pale scale patches on
the abdomen and small setae on sternum |.
In addition mattogrossensis lacks leg spec-
kling, no dark spots on the notum, and pos-
terolateral scale tufts, all typical of most oth-
er Arribalzagia.
Anopheles venezuelae Evans
(Fig. 4A-D)
Evans 1922: 213-222, Plate XI, in subgenus
Arribalzagia.
Anopheles venezuelae is a junior synonym
of An. punctimacula.
Label data.—School of Trop. Med., Liv-
erpool University label ““A. venezuelae Ev-
ans La Cabrero Estado Carabobo 1921 Dr.
M Nunez Tovar’; “Holotype of Anopheles
venezuelae Evans det. J. Chainey 1975.”
Condition of specimen.—Female. The
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
specimen is incomplete but the remaining
portions are in good condition. Missing:
Abdomen; right wing and part of left wing,
most of left wing is affixed to plastic base
into which is pinned the specimen; flagello-
meres 3-5 missing on both sides; palpo-
meres 2—5 missing on both sides though half
of 2 on left is present; left and right fore-
tarsomeres 2-5, left midtarsomeres 2-5,
right midtarsomeres 3-5, left and right
hindtarsomeres 2-5.
Discussion.—Anopheles venezuelae was
described from a single specimen in 1922.
When Evans received more material it be-
came apparent that the characters that she
used to distinguish it from An. punctimac-
ula were no more than normal variation. In
1923 Evans sunk An. venezuelae under An.
punctimacula. 1 have seen the type of An.
punctimacula and studied in great detail
material of all stages from throughout its
range and agree with her assessment.
ACKNOWLEDGMENTS
Special thanks to Taina Litwak for her
excellent illustrations and interpretation of
slide mounted structures, and to E. L. Pey-
ton for his valuable advice and for the dif-
ficult dissection of the An. mediopunctatus
genitalia. I also thank Bruce Townsend for
the loan of the material reviewed here and
Ronald Ward for critical review of the
manuscript and generous assistance in pub-
lication of the manuscript.
The views of the author do not purport
to reflect the position of the Department of
the Army or the Department of Defense.
LITERATURE CITED
Christophers, S. R. 1923. An Anopheles of the My-
zorhynchus group (Anopheles amazonicus sp. n.).
Ann. Trop. Med. Parasitol. 17: 71-78.
Evans, A. M. 1922. Notes on Culicidae in Venezuela,
with descriptions of new species. Part H. Ann.
Trop. Med. Parasitol. 16: 213-222.
. 1923. Notes on Culicidae in Venezuela, with
VOLUME 90, NUMBER 4
descriptions of new species. Part III. Ann. Trop.
Med. Parasitol. 17: 101-111.
Reid, J. A. and K. L. Knight. 1961. Classification
within the subgenus Anopheles (Diptera, Culici-
dae). Ann. Trop. Med. Parasitol. 55: 474-488.
421
Shannon, R. C. 1933. Anophelines of the Amazon
Valley. Proc. Entomol. Soc. Wash. 35: 117-133.
Theobald, F. V. 1903. A monograph of the Culicidae
or mosquitoes. Vol. 3, 359 pp., British Museum
(Natural History), London.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 422-427
GENETIC VARIATION AND SYSTEMATICS OF FOUR TAXA OF
NEOTROPICAL WALKING STICKS (PHASMATODEA: PHASMATIDAE)
RONALD A. VAN DEN BusscCHE, MICHAEL R. WILLIG,
RONALD K. CHESSER, AND
RosBertT B. WAIDE
(RAVDB, MRW, RKC) Department of Biological Sciences and The Museum, Texas
Tech University, Lubbock, Texas 79409; (MRW and RBW) Center for Energy and En-
vironment Research, GPO Box 3682, San Juan, Puerto Rico 00936.
Abstract. —Electrophoretically detectable genetic variation for six isozymes encoded by
seven loci was analyzed in four taxa of walking sticks (Phasmatidae) that occur in a
neotropical rainforest in eastern Puerto Rico. No phylogenetic analysis previously has
been conducted on any phasmatid. All seven loci exhibited variation among Diapherodes
achalus, Lamponius portoricensis, Pseudobacteria yersiniana, and an unnamed taxon
(species X). Coefficients of genetic distance between these four taxa ranged from 0.349 to
0.571. The UPGMA of Rogers’ genetic distance indicated considerable genic dissimilarity
among taxa (the two taxa which were least dissimilar connected at a value of 0.350). The
four taxa represent a holophyletic group for which an outgroup was not analyzed, this
situation, in conjunction with the short internode distance in the Fitch-Margoliash anal-
ysis, provides only limited resolution of the phylogenetic associations among the four
taxa.
Key Words:
bacteria.
The Phasmatidae, or walking sticks, are
primarily a tropical group of folivores that
occasionally have an economic impact on
human-manipulated systems (Campbell
1960, 1961, 1966, 1974, Campbell and
Hadlington 1967, Mananec 1966, 1967,
1968, Paine 1968). Most phasmatids are ac-
tive nocturnally, and remain inactive on the
surface of leaves and stems, or in leaf litter,
during the day. Even at night, walking sticks
are usually immobile and cryptic (Cott 1940,
Moxey 1972). Besides these few cursory ob-
servations, the natural history of the Phas-
matidae is poorly known. The best contem-
porary review of their biology was presented
by Bedford (1978); however, his review pri-
marily focused on Old World taxa. Al-
electrophoresis, isozymes, Puerto Rico, Diapherodes, Lamponius, Pseudo-
though many studies have been conducted
on the taxonomy of the Phasmatidae (see
Bedford 1978), a dearth of information is
available concerning their phylogenetic re-
lationships. Detailed studies in the New
World have lagged far behind those in the
Old World.
Moxey (1972), in a detailed but yet un-
published study concerning systematics of
walking sticks from the West Indies, rec-
ognized 54 Antillean species, distributed
into 16 genera. Many of these species are
restricted to one or only a few islands. Other
than the work by Moxey (1971, 1972), the
most recent research on phasmatids of the
West Indies focused on population dynam-
ics and natural history of Lamponius por-
VOLUME 90, NUMBER 4
toricensis from the Tabonuco rainforest of
Puerto Rico (Willig et al. 1986). We pres-
ently are conducting studies aimed at ex-
amining the genetic bases of food preference
and spatial distribution in L. portoricensis.
The only other published study on Puerto
Rican walking sticks examined the karyo-
types in a selected few species and evaluated
their response to low-level gamma irradia-
tion (Virkki 1970). No phylogenetic anal-
ysis has been attempted on the Phasmati-
dae.
Puerto Rico and nearby Mona Island are
inhabited by 11 species of walking sticks
representing six genera (Moxey 1972). Of
these 11 species, eight are endemic to Puerto
Rico, two are found only on Puerto Rico
and St. Thomas, and one is restricted to
Mona Island. Of the taxa that we examined
from eastern Puerto Rico, D. achalus, L.
portoricensis, and P. yersiniana have wide-
spread distributions throughout the moun-
tainous regions of Puerto Rico. The fourth
taxon is unnamed, and taxonomic work is
being pursued by Garrison and Willig (pers.
comm.). Although this unnamed taxon
shares some morphological characteristics
with the genus Lamponius, Moxey’s (1972)
morphological treatment of the West Indian
taxa was not based upon features of the male
genitalia (the primary characteristics used
in systematic studies of the Orthoptera) and
may represent spurious results. We do not
formally classify this taxon and refer to it
hereafter as species X. Species X occurs pri-
marily in the mossy dwarf-forests (above
1000 m) of the Caribbean National Forest,
located in the Luquillo Mountains on the
eastern part of the island. The purpose of
this project was to examine, using protein
electrophoresis, the phylogenetic and phe-
netic relationships of phasmatids that occur
in eastern Puerto Rico.
MATERIALS AND METHODS
Walking sticks were collected in the Ca-
ribbean National Forest (18°10'N, 65°
30'W), Puerto Rico, between 12 June and
423
2 August 1985. Specific localities of collec-
tion and sample sizes (N) for the four taxa
examined were: Diapherodes achalus, (a.)
km 10.6 on route 186 (N = 1), El Yunque
Quadrangle, Municipality of Naguabo; (b.)
km 13.5 on route 191 (N = 4), El Yunque
Quadrangle, Municipality of Naguabo;
species X, km 13.5 on route 191 (N = 13),
El Yunque Quadrangle, Municipality of Na-
guabo; Lamponius portoricensis, near route
180 (N = 66), El Verde Field Station, Mu-
nicipality of Rio Grande; Psuedobacteria
yersiniana, (a.) km 10.6 on route 186 (N =
13), El Yunque Quadrangle, Municipality
of Naguabo; (b.) km 13.5 on route 191 (N
= 5), El Yunque Quadrangle, Municipality
of Naguabo. After collection, all individuals
were transported to El Verde Field Station
and were identified to specific level. Each
specimen (minus abdomen) was placed in
a 1.5 ml Eppendorf tube and immediately
frozen; upon arrival at the Department of
Biological Sciences, Texas Tech University,
specimens were stored at — 70°C.
Prior to allozymic analysis, each individ-
ual was homogenized in a buffered solution
(pH = 6.8). The tissue homogenate was ana-
lyzed using standard horizontal starch-gel
electrophoretic techniques (Selander et al.
1971, Harris and Hopkinson 1977). The
following loci were examined: acid phos-
phatase (Ap); aldehyde oxidase (Ao); ester-
ase-1, -2, and -3 (Es-1, -2, -3); glucose de-
hydrogenase (Gdh); glucose phosphate
isomerase (Gpi); glutamate oxaloacetate
transaminase-! and -2 (Got-1, -2); leucine
amino peptidase-1 and -2 (Lap-1, -2); ma-
late dehydrogenase-1, -2, and -3 (Mdh-l,
-2, -3); nucleoside phosphorylase (Np): pep-
tidase-1 and -2 (Pep-B-1, -2); and phos-
phoglucomutase-1, -2, and -3 (Pgm-1, -2,
-3). Only loci with consistent banding pat-
terns (Ap, Es-1, Gdh, Lap-1, Mdh-1, -2,
Pgm-1) were used in the subsequent anal-
yses.
When multiple isozymes of a protein were
present, the locus that migrated the farthest
anodally was designated as “1,” and loci
424
that migrated progressively in the direction
of the cathode were given higher numerical
designations. For each locus, the most com-
mon allele was designated as “100” and oth-
er alleles were assigned numeric values ac-
cording to their mobility relative to the most
common allele.
Genetic distances between each pair of
taxa were calculated from allelic frequency
data (Nei 1972, Rogers 1972). Nei’s (1972)
and Rogers’ (1972) genetic distance values
were similar, therefore, only Rogers’ (1972)
genetic distance values were used in sub-
sequent analyses. Relationships among
species X, D. achalus, L. portoricensis, and
P. yersiniana were analyzed by genetic dis-
tances (Rogers 1972) and summarized in
the form of a distance dendrogram that was
obtained from a UPGMA (unweighted pair-
group method using arithmetic averages;
Sneath and Sokal 1973) clustering method.
Phyletic relationships also were summa-
rized in the form of an unrooted tree, pro-
duced by the Wagner parsimony analysis
(Farris 1970) using the WAGNER78 pack-
age and the Fitch-Margoliash analysis of the
distance matrix (Fitch and Margoliash
1967).
RESULTS
Allele frequencies of the seven polymor-
phic loci and their distribution within taxa
appear in Table |. For the Ap locus, in which
a total of seven alleles were detected, D.
achalus was polymorphic for the two slow-
est alleles (“*85” and 90’), species X was
fixed for the “95” allele, and L. portoricensis
was fixed for the “100” allele. Pseudobac-
teria yersiniana was extremely polymorphic
in possessing the five fastest alleles (95”
through 110°) at the Ap locus, thereby
sharing the “95” allele with species X and
the “100” allele with L. portoricensis. Only
three alleles were detected at the Es-1 locus.
Diapherodes achalus, species X, and P. yer-
siniana were each fixed for the “100” allele,
whereas L. portoricensis was polymorphic
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and possessed all three alleles. No fixed dif-
ferences or unique distribution ofalleles was
detected for either Gdh or Lap-1, with the
exception that P. yersiniana possessed a
greater number of alleles than any other tax-
on examined for these loci. For Mdh-1, D.
achalus and species X shared two of the six
alleles (“95 and ‘100’ alleles) and each
had approximately the same frequency in
each taxon. Lamponius portoricensis was
fixed at this locus for the “100” allele, which
also was detected in D. achalus and species
X. Again, P. yersiniana exhibited a high de-
gree of polymorphism by possessing all six
of the alleles for Mdh-1. For Mdh-2, only
D. achalus was fixed for an allele (*100”
allele), whereas the other three taxa were
each characterized by the presence of all
three alleles. Finally, for Pgm-1, only L. por-
toricensis was fixed for an allele (*100” al-
lele), whereas the other three taxa each ex-
hibited much polymorphism. Rogers’ (1972)
genetic distance values between D. achalus
and species X, D. achalus and L. portori-
censis, and D. achalus and P. yersiniana were
0.349, 0.524, and 0.475, respectively. Dis-
tance values between species X and L. por-
toricensis, species X and P. yersiniana, and
L. portoricensis and P. yersiniana were
0.455, 0.426, and 0.571, respectively. Ge-
netic distance relationships among the four
taxa of walking sticks are summarized in
the form of a distance dendrogram (Fig. 1).
Based on a phenetic analysis of allozymic
data, species X is more similar to D. achalus
than to L. portoricensis or P. yersiniana.
Both the Wagner and Fitch-Margoliash
analyses for phyletic relationships gave
identical tree topologies; therefore, only the
result of the Fitch-Margoliash analysis is
presented. Phyletic relationships, summa-
rized by the Fitch-Margoliash analysis were
generated from an unrooted tree that reflects
the actual observed genetic distance in the
length of the branches (Fig. 2). The average
value of heterozygosity (H) for these four
taxa is 0.060. Heterozygosity estimates for
each taxon are: H = 0.051 (species X), H =
VOLUME 90, NUMBER 4
Table 1. Allele frequencies of seven variable loci
for four taxa of walking sticks collected from the Ca-
ribbean National Forest (18°10’N, 65°30'W), Puerto
Rico. See text for locus abbreviations. The common
allele is designated as the “100” allele and additional
alleles are numbered according to the mobility of their
products relative to that of the common allele. Alleles
not listed in the table are: Ap-110, Es-1-95, Gdh-95,
Mdh-1-115, Mdh-2-115, and Pgm-1-120 (their fre-
quencies can be obtained by subtraction).
L. portori- P,
Locus Allele D. achalus Species X censis yersiniana
Ap 105. 0.000 0.000 0.000 0.267
100 0.000 0.000 1.000 0.233
98 0.000 0.000 0.000 0.367
95 0.000 1.000 0.000 0.100
90 =0.400 0.000 0.000 0.000
85. 0.600 0.000 0.000 0.000
Es-1 105 0.000 0.000 0.562 0.000
100 1.000 1.000 0.308 1.000
Gdh 115) 0.400 0.143 0.000 0.067
112. 0.000 0.000 0.000 0.133
110 0.000 0.000 0.000 0.667
105. 0.200 «(0.214 0.175 (0.067
100 0.400 0.643 0.635 0.067
Lap-1 110 0.400 0.000 0.000 0.056
105 0.600 0.286 0.063 0.000
100 0.000 0.500 0.813 0.167
95 0.000 0.215 0.125 0.778
Mdh-1 110. 0.000 0.077. 0.000 0.177
100 0.500 0.462 1.000 0.118
95 0.500 0.462 0.000 0.177
90 0.000 0.000 0.000 0.441
80 0.000 0.000 0.000 0.059
Mdh-2 110 0.000 0.357 0.273 0.177
100 1.000 0.571 0.561 0.235
95 0.000 0.071 0.167 0.529
Pgm 110. =©0.100 0.214 =—0.000 0.250
105. 0.100 0.286 0.000 0.056
100 0.800 0.393 1.000 0.583
98 0.000 0.036 0.000 0.000
95 0.000 0.000 0.000 0.028
85 0.000 0.000 0.000 0.056
75 0.000 0.071 0.000 0.000
0.057 (D. achalus), H = 0.044 (L. portori-
censis), and H = 0.087 (P. yersiniana).
DISCUSSION
Rogers’ genetic distance values indicate
that considerable genetic divergence has oc-
curred among the four taxa examined. Val-
ues range from 0.349 for the pairwise com-
0.56 0.49 0.42 0.35 0.28 0.21 0.14 0.07 0.00
ee
r = 0.856 D. achalus
species X
P. yersiniana
L. portoricensis
Fig. 1. Dendrogram that depicts the relationship of
Diapherodes achalus, Lamponius portoricensis, Pseu-
dobacteria yersiniana, and an unnamed taxon (species
X) derived from a UPGMA of Rogers’ genetic distance
coefficients.
parison of species X and D. achalus to 0.571
for the pairwise comparison of L. portori-
censis and P. yersiniana. Nevo (1978) re-
ported levels of heterozygosity for a variety
of invertebrates; however he did not report
data for the Phasmatidae. The heterozygos-
ity values in this study occur within the range
of those reported for closely related Or-
thoptera and other insects.
The limited research that has been re-
ported on stick-insects from Puerto Rico
has focused on taxonomy (Gray 1835, Bur-
meister 1838, Saussure 1868, Brunner and
Redtenbacher 1892, Rehn 1903, Rehn and
Hebard 1938, Wolcott 1923, 1936, 1941,
1948, Moxey 1971, 1972), systematics
(Moxey 1972), population dynamics and
natural history (Wolcott 1951, Willig et al.
1986), or effects of radiation (Virkki 1970).
Based upon distributional data, Moxey
(1972), suggested that the Antillean genera
considered herein (Diapherodes, Lampon-
ius, and Pseudobacteria) probably each
evolved from Central American stocks,
which subsequently invaded the West In-
dies from west to east. Moreover, he noted
that Lamponius and Diapherodes probably
evolved from a common stock. Our allo-
zymic data provide limited support to the
hypothesis that L. portoricensis and P. yer-
siniana may have evolved from different
ancestral stocks in that they are separated
by a genetic distance much greater than are
the other taxa in the UPGMA (Fig. 1).
426
D. achalus L. portoricensis
P. yersiniana
species X
Fig. 2. Unrooted tree generated by a Fitch-Mar-
goliash analysis based on Rogers’ genetic distances for
Diapherodes achalus, Lamponius portoricensis, Pseu-
dobacteria yersiniana and an unnamed taxon (species
X).
Nonetheless, the large genetic distance be-
tween all taxa in the UPGMA (Fig. 1) and
the short internode distance in the Fitch-
Margoliash tree (Fig. 2) fail to provide con-
clusive information about the systematic
relationships of these taxa or the proper af-
filiation of species X. Clearly, the phylo-
genetic tree should be viewed with caution
because it does not involve a monophyletic
group and no out group was used to root
the tree. Five possible phylogenies (we ex-
clude those with unresolved trichotomies if
rooting occurs at vertices) are congruent with
the Fitch-Margoliash topology (Fig. 2). In
three of them, species X is more closely
related to D. achalus than to any of the other
taxa. Alternatively, species X may be less
related to any of the other taxa than those
taxa are to each other, or L. portoricensis
and P. yersiniana, as a group, may be more
closely related to species X than any of those
three taxa are to D. achalus.
Future studies addressing phasmatid sys-
tematics in the West Indies should obtain
adequate samples of all taxa occurring
throughout the West Indies, and include an
outgroup by which one could root a phy-
logenetic tree. Thereafter, it would be pos-
sible to test a variety of hypotheses con-
cerning phasmatid systematics and thereby
facilitate the classification of species X as
well.
ACKNOWLEDGMENTS
We thank J. K. Jones, Jr., R. W. Garrison,
GaSe Spicer Ja‘. ‘Cokendolphers Re D:
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Bradley, G. R. Camilo, M. J. Hamilton, and
especially R. D. Owen for providing critical
reviews of an earlier draft of this manu-
script. This study was funded by the Office
of Health and Environment Research, De-
partment of Energy, and Oak Ridge Asso-
ciated Universities via a Faculty Partici-
pation grant to MRW. We also acknowledge
the Graduate School of Texas Tech Uni-
versity for support of MRW and RAVDB,
and State of Texas Organized Research Fund
for support to MRW.
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Burmeister, H. 1838. Handbuch der Entomologie.
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Campbell, K. G. 1960. Preliminary studies in pop-
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1974. Factors affecting the distribution and
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trees. Syst. Zool. 19: 172-189.
Fitch, W. M. and E. Margoliash. 1967. Construction
of phylogenetic trees. Science. 155: 279-284.
Gray, G. R. 1835. Synopsis of the species of insects
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belonging to the family of Phasmidae. London. 48
pp.
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of enzyme electrophoresis in human genetics.
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dymuria violescens on the growth of alpine ash.
Australian For. 30: 125-130.
1967. Mortality and diameter growth in
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lian For. 31: 221-223.
1968. Influence of defoliation by the phas-
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growth and pattern of growth rings in alpine ash.
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West Indies: Two new genera. Psyche 78(1/2): 67—
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. 1972. The stick-insects (Phasmatodea) of the
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published Ph.D. dissertation, Harvard Univ. 211
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lations: Patterns and theory. Theor. Pop. Biol. 13:
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control in Fiji of the coconut stick insect Graeffea
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island of Porto Rico. Trans. American Ent. Soc.
29(2): 129-136.
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and species of West Indian Mantidae and Phas-
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taxonomy. W. H. Freeman and Co., San Francis-
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A Study of Irradiation and Ecology at El Verde,
Puerto Rico. U.S. Atomic Energy Comm.
Willig, M. R., R. W. Garrison, and A. J. Bauman.
1986. Population dynamics and natural history
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99: 47-50.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 428-439
ANTENNAL SENSILLA AND SETAE OF EVAGETES PARVUS
(HYMENOPTERA: POMPILIDAE)
MarINA A. LANE, FRANK E. KURCZEWSKI, AND ROBERT B. HANNA
(MAL, FEK) Environmental and Forest Biology, S.U.N.Y. College of Environmental
Science and Forestry, Syracuse, New York 13210; (RBH) N. C. Brown Center for Ultra-
structure Studies, S.U.N.Y. College of Environmental Science and Forestry, Syracuse,
New York 13210.
Abstract.—The antennae of cleptoparasitic female Evagetes parvus were investigated
using scanning electron microscopy. Setae and sensilla placodea, corrugated conical sen-
silla, pit organs, sensilla campaniformia, sensilla trichodea A, B, C, D, sensilla basiconica
and sensilla spatulata were located, described and illustrated. The sensilla of E. parvus
most clearly resemble those of two other pompilids, Anoplius tenebrosus and A. viaticus;
however, spatulate sensilla were not found in either of these species. Two large zones on
the flagellum of FE. parvus are dominated by corrugated conical sensilla and placoid sensilla,
respectively. The former are found only on the flattened ventral surfaces of flagellomeres
2-10, whereas the latter surround these sensilla and occur also on flagellomere 1. The
corrugated conical sensilla in conjunction with the sensilla basiconica and sensilla spatulata
are probably used in locating the host (buried spider) because of their ventral position on
the antennal surface and their morphology.
Key Words:
Members of the genus Evagetes are clep-
toparasitic, Ovipositing on spiders which
have been captured by other genera of Pom-
pilidae. The most distinguishing character-
istic of the genus is the short antennae which
are thickened and somewhat flattened ven-
trally in the female (Evans 1950). Females
are usually observed walking on sandy sur-
faces while tapping the soil with their an-
tennae, or stalking nesting pompilid wasps.
Female Evagetes frequently enter nests being
provisioned by other pompilids either be-
fore or after closure. Such a wasp is able to
detect a buried spider by some clue, most
likely olfactory and/or tactile, whereupon
she unearths the paralyzed prey, destroys
the pompilid’s egg, and lays her own egg on
the prey (Evans and West Eberhard 1970).
cleptoparasite, chemoreception, mechanoreception, proprioception
The subsequent closure by a female Eva-
getes may be thorough or loose. Such clep-
toparasitic behavior has been noted in E.
parvus (Cresson) by Evans (1950) and Evans
and Yoshimoto (1962). Similar activities of
other species of Evagetes have been report-
ed by Richards and Hamm (1939) and Ev-
ans et al. (1953) and summarized by Krom-
bein (1979).
While searching for the buried paralyzed
spider, Evagetes females extend their an-
tennae outward in a stiff “V” and rapidly
tap the ground with the ventrally flattened
surfaces. The particular sensory sensilla used
in detecting the spider are probably located
on the ventral surfaces of the antennae. The
intent of this paper was to examine and
identify the antennal sensilla and setae of
VOLUME 90, NUMBER 4
female Evagetes parvus, using SEM, and to
attempt to determine which sensilla are used
to locate the buried prey.
METHODS AND MATERIALS
Female EF. parvus were collected from a
sand pit near Auburn, Cayuga County, New
York, during the summers of 1981 and 1982.
They were kept refrigerated in ventilated
glass vials until used. Preparation consisted
of excising either the antennae or heads, and
immersing these in methylene chloride for
two days to dissolve the waxy layer. Two
pairs of antennae were then mounted with
silver paint, and sputter-coated in a Tech-
nics sputter-coater with ionized gold-pal-
ladium alloy. Another specimen, kept in
methylene chloride, was mounted on a stud
with silver paint and coated with gold in a
Kinney SC2 high vacuum metal evaporator.
One specimen was air-dried in an attempt
to see if there were any differences in the
final appearances of the sensilla. This spec-
imen was then mounted on double sticky
tape, grounded with silver paint, and sput-
ter-coated with gold-palladium alloy. Spec-
imens were then viewed on an ETEC Auto-
scan scanning electron micropscope, at
accelerating voltages of 10 and 20 KV. There
were no differences found in antennal fea-
tures between this method and the first one
used.
The nomenclature used to describe the
various sensilla has been modified from
Agren (1977) and Alm and Kurczewski
(1982).
RESULTS
Female Evagetes parvus have filiform
antennae, ca. 2.6 mm long, which are com-
posed of scape, pedicel, and flagellum with
flagellomeres designated 1-10 proximally to
distally. The ventral surface is in contact
with the substrate during host-searching, is
flattened from flagellomere 2-10, and is
highly sensory. The scape, pedicel, and fla-
gellomere 1 are mainly setaceous. In de-
scribing the sensilla we refer to the antennae
429
Fig. 1.
Antenna of female Evagetes parvus, as viewed
dorsally, showing the curled, resting position typical of
many female Pompilidae.
being held horizontally in front of the wasp
as during host-searching. When the anten-
nae are curled back in the resting position
(Fig. 1), the medial and ventral surfaces are
actually facing outward away from the in-
sect. The sensilla found are described below
in order of their prominence and complex-
ity except for s. spatulata which are de-
scribed near the end because they resemble
s. basiconica.
SENSILLA
Sensilla placodea: A placoid sensillum is
elongate, convex and has a membranous fold
encircling its base (Figs. 2A, B). It is ori-
ented parallel to the long axis of the antenna,
and its dimensions are 5 x 15 um. Placoid
sensilla are abundant, extending from a small
area on flagellomere | along the sides and
over most of the dorsum of the remaining
flagellum. They are spaced evenly and num-
430
2'B
Fig. 2.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
A, Sensillum placodeum, flagellomere 8, sunken into cavity, with distal end projecting above antennal
surface. B, Sensillum placodeum, dorsal view, showing membranous fold around base, flagellomere 4.
Fig. 3.
A, Corrugated conical sensillum with subterminal furrow (arrow), flagellomere 3. B, Corrugated conical
sensillum showing exudate covering furrow (arrow), flagellomere 6.
ber ca. 17 in a column down the length of
a typical flagellomere.
Corrugated conical sensilla: Corrugated
conical sensilla are stout, truncate receptors
(Figs. 3A, B) which lean distally and are
cushioned against the cuticle by a thick
membrane. Numerous grooves extend from
base to apex. The tip may be indented (Fig.
3A) or covered with an exudate (Fig. 3B).
The sensillum is ca. 10 um long and 5 um
wide at the base. It is found abundantly only
on the flattened ventral surface of the fla-
gellum. These sensilla form a small triangle
on the underside of flagellomere 2, continue
along the entire ventral surface of the fla-
gellum, and end in a triangular area on the
underside of flagellomere 10.
Pit organs: These hollowed pits resemble
domes with holes in the centers (Figs. 4A,
B). A smooth peg may be visible in the bot-
VOLUME 90, NUMBER 4
Fig. 4.
surface, flagellomere 10. Irregular surface may be due to fixation.
Fig. 5.
A, Small pit organ, distal end, flagellomere 7. B, Large pit organ, revealing peg within pit, dorsal
A, Sensilla campaniformia (SC) and s. triichodea A (A) in asymmetrical sockets, flagellomere 1. B,
Broken sensillum trichodeum A, showing hollow center (arrow), flagellomere 5. p = pit organ.
tom of the pit (Fig. 4B). Two types of pit
organs are distinguished by the size of the
central aperture, one being ca. | wm and the
other, 3 um in diameter (Figs. 4A, B). Pit
organs are relatively few in number, varying
from 2 to 9 per flagellomere. They are found
in clusters on the medial and lateral surfaces
on the proximal half of the flagellum. To-
ward the distal end of the flagellum the clus-
ters are placed dorsally.
Sensilla campaniformia: This receptor has
a shallow circular depression with a central
papilla (Fig. 5A), is oval in shape, and has
a diameter of ca. 6 x 10 wm. Campaniform
sensilla occur only on the flagellum, their
greatest concentration being a group of 5-8
on the ventrolateral surface of flagellomere
1. They occur in small clusters at the middle
and distal ends of the remaining flagello-
meres.
Fig. 6.
(B) with heavy sculpturing, flagellomere 8. S = setae.
Figs i
a i
AN
Vie \ Nd A
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
A, Sensillum trichodeum B (B), flagellomere 8. p = pit organ S = setae. B, Sensillum trichodeum B
A, Medium-length sensillum trichodeum C (C), typical form, with sunken socket, flagellomere 3. B,
Long sensillum trichodeum C (C) with raised socket, ventral surface, flagellomere 1. C, Short sensillum tricho-
deum C (C) among corrugated conical sensilla, flagellomere 3.
Sensilla trichodea A: These hairs have
abrupt tips, are ca. 12 wm long (Fig. 5A),
and sit in asymmetrical sockets. The base
is large and bulbous and the thick shaft bends
acutely at about a 70° angle toward the distal
end of the antenna. The broken sensillum
in Fig. 5B shows that s. trichodea A are
hollow. Trichodea A occur in patches down
the center of the lateral surfaces of flagello-
meres 2-9 and are scattered on the medial
surfaces of flagellomeres 3-10.
Sensilla trichodea B: These slender hairs,
ca. 13 um long, taper gradually to a point,
and have no visible socket at the base (Fig.
6A). Trichodea B lie parallel to the antenna
and point distally. This sensillum is thinner
than surrounding setae, but is similar in
shape and sculpture to them. The sensillum
in Fig. 6B is only 9 um long and is scarce
on the dorsum of the antenna. The sensil-
lum shown in Fig. 6A is most dense in oc-
currence laterally on the flagellum and is
VOLUME 90, NUMBER 4
<<
Fig. 8.
with hollow double chamber (arrow), pedicel.
Fig. 9.
433
A, Group of sensilla trichodea D (D), proximal end of scape. B, Broken sensillum trichodeum D,
Sensillum basiconicum (SB), proximally set in socket, flagellomere 3.
Fig. 10. Sensillum spatulatum (SS) next to s. basiconicum (SB), flagellomere 6.
spaced evenly. Only one or two trichodea
B are seen on the dorsum of the scape and
pedicel.
Sensilla trichodea C: This slender, hair-
like sensillum (Figs. 7A, B, C) has a blunt
tip and 4—5 faint vertical furrows running
from base to tip. The flexible socket has a
circular, membranous collar which may be
depressed or elevated. This sensillum is
straight or curved slightly, 5-25 um long,
and stands nearly perpendicular to the an-
tennal surface. Sensilla trichodea C are lo-
cated on all of the flagellomeres: the longest
at the distal ends, the shortest on the ventral
surfaces, and medium-length ones in indis-
tinct, widely-spaced rings.
Sensilla trichodea D: Sensilla trichodea D
are similar to trichodea C, the difference
being a ring around the base of the hair in
trichodea D (Figs. 8A, B). The longest sen-
silla, 5O wm long, are located near the distal
ends of the scape and pedicel. The broken
434
trichodeum D in Fig. 8B shows a distinct
double chamber internally.
Sensilla basiconica: These pegs are ca. 6
um high, straight and stout, with a blunt,
almost flat, apex (Figs. 9, 10). There are 1 1-
12 vertical grooves on each sensillum. The
socket 1s large, round, and rarely depressed,
with the sensillum placed in the proximal
side of the socket. Sensilla basiconica are
distributed on the flattened ventral surface
of the flagellum.
Sensilla spatulata: This unusual fan-
shaped sensillum (Fig. 10) is oriented with
the concave face of the fan directed away
from the antennal surface. It has a large
round socket and is ca. 8 wm long. It is found
scattered widely on the ventral surface
among the corrugated conical sensilla.
Setae.—The setae are highly variable in
size and shape, commonly long and straight,
smooth or fluted, and not innervated (Fig.
11A). On the extreme dorsal surface of the
antenna they are short, thick, and deeply
grooved (Fig. 11B). On the distal half of the
flagellum the setae occur among the cor-
rugated conical sensilla and are broadly sa-
ber-shaped with deep grooves (Fig. 11C).
The setae at the antennal tip are relatively
long, broad apically, and longitudinally
grooved (Fig. 11D).
Sensillar zones and interrelationships. —
Two large zones on the flagellum are dom-
inated by corrugated conical sensilla and
placoid sensilla, respectively (Fig. 12). The
corrugated conical sensilla are found only
on the flattened ventral surfaces of flagello-
meres 2-10. The placoid sensilla surround
the corrugated conical sensilla, extending
from flagellomeres 1-10. Each zone has a
characteristic sensillar composition. Sensil-
la basiconica and s. spatulata are distributed
among the corrugated conical sensilla (Fig.
13). Pit organs, s. campaniformia, and s.
trichodea A are found among the placoid
sensilla (Fig. 14). The pit organs and s. cam-
paniformia are coincidentally dispersed in
two longitudinal bands which border the
sides of the corrugated conical sensillar zone.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
There are 2-4 of each type per band per
segment. On the distal half of the flagellum
the two bands gradually merge dorsally into
one broad band. In addition there are two
specialized sensory spots. There is a con-
centration of 5-8 campaniform sensilla on
the ventrolateral surface of flagellomere |
within the triangular patch of placoid sen-
silla. There is a group of 7-9 pit organs on
the medial surface of flagellomere 2.
The majority of s. trichodea A is concen-
trated along the border of the corrugated
conical sensillar zone with the remainder
widely dispersed on the dorsum of the fla-
gellum. Sensilla trichodea D are found in
small groups on the scape and pedicel,
whereas s. trichodea C are evenly distrib-
uted over the entire flagellum. The last third
of the terminal flagellomere is devoid of s.
placodea and corrugated conical sensilla, but
has an abundance of s. trichodea B and C.
DISCUSSION
In Hymenoptera, placoid sensilla are
abundant (Slifer 1970) and are thought gen-
erally to be olfactory organs (Schneider
1964). In female Evagetes parvus they are
distributed evenly and densely in a zone
directed dorsad and along both sides of most
of the flagellum. The sensilla placodea of
female E. parvus are nearly identical to those
of females of the pompilids Anoplius viati-
cus (L.) and A. tenebrosus (Cresson) (Wal-
ther 1979, Alm and Kurczewski 1982), are
smooth-walled, project from a sunken pit
above the antennal surface, and are mod-
erately elongate. In addition, s. placodea of
A. viaticus end in a point (Walther 1979).
Shapes of placoid sensilla vary within Hy-
menoptera. Compared to those of E. parvus,
placoid sensilla of Braconidae and Ichneu-
monidae are more elongate and often ex-
tend above the antennal surface at their dis-
tal ends. In Vespidae, placoid sensilla are
shorter, flatter on the surface (Callahan
1970), and similar to those of Pompilidae
in size and arrangement. Higher Apoidea
VOLUME 90, NUMBER 4
1118
Fig. 11.
10. B=s. trichodeum B; C = s. trichodeum C; SP = s. placodeum. B, Short, thick, spiralled seta (arrow), dorsum,
flagellomere 8. C, Grooved, saber-shaped seta (arrow), ventral surface, flagellomere 10. B = s. trichodeum B; C
= s. trichodeum C; CCS = corrugated conical sensillum. D, Thick, curved seta (arrow), tip of antenna. B = s.
trichodeum B; C = s. trichodeum C.
have numerous circular and flat placoid sen-
silla (Shifer and Sekhon 1960, Dietz and
Humphreys 1971, Agren 1977, 1978). Based
upon the presence of numerous small pores,
Norton and Vinson (1974) suggested that
elongate s. placodea of Ichneumonidae and
Braconidae are chemoreceptors. Kaissling
and Renner (1968) showed electrophysio-
logically that, in both sexes of Apis mellifera
L., this sensillum was stimulated by the
435
A, Long and straight setae, both smooth (single arrow) and fluted (double arrow) types, flagellomere
queen substance and the scent of Nasanov’s
gland.
The corrugated conical sensilla of E. par-
vus closely resemble those of Anoplius via-
ticus (described as s. basiconica P1) (Wal-
ther 1979) and 4. tenebrosus (Alm and
Kurczewski 1982). Callahan (1970) coined
the term “conical” for this type of sensillum
in the vespids Polistes metricus Say and P.
annularis (L.). Those of Polistes species have
436 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 12.
and corrugated conical sensilla (double arrow).
Fig. 13.
s. trichodea C (C). B = s. trichodea B.
Fig. 14.
trichodeum C (C).
a distinct terminal pore, while those of An-
oplius viaticus and A. tenebrosus (Walther
1979, Alm and Kurczewski 1982) and E.
parvus have a deep furrow which probably
indicates a terminal pore. Alm and Kurc-
zewski (1982) reported that Kaissling and
Klein (personal communication) found these
sensilla to be reminiscent of gustatory bris-
tles.
Medioventral surface of flagellomere 7, showing areas containing sensilla placodea (single arrow)
Zone dominated by corrugated conical sensilla (ccs), containing s. basiconica, s. spatulata (ss), and
Zone dominated by sensilla placodea (sp), containing s. campaniformia (sc), pit organs (p), and s.
Alm and Kurczewski (1982) noted that
these sensilla were found only on the an-
tennae of female Anoplius tenebrosus, and
Walther (1979) found the same for females
of A. viaticus. In E. parvus females the cor-
rugated conical sensilla may be used to de-
tect buried, paralyzed spiders. These sen-
silla are abundant on the ventral antennal
surface and their morphology suggests that
VOLUME 90, NUMBER 4
they are gustatory. The corrugated conical
sensilla may receive a chemical residue or
heavy odor molecule from buried prey or
from pompilid-manipulated sand particles,
and this provides the impetus for the wasp
to unearth the spider. The broad, flattened
apex provides a large surface area for che-
moreception. There are no long sensilla or
setae on the flattened ventral surface of the
antenna of E. parvus, thus allowing the cor-
rugated conical sensilla to contact the ground
surface without interference.
The pit organs of Evagetes parvus resem-
ble those of Anoplius viaticus and A. tene-
brosus (Walther 1979, Alm and Kurczewski
1982), Colletidae (Agren 1977), and An-
drenidae (Agren 1978). They appear as dif-
ferent-sized apertures through which the
largest reveals an internal peg. They are sim-
ilar to the s. ampullaceae and s. coeloconica
of Apis mellifera which Dietz and Hum-
phreys (1971) differentiated as “smaller” and
“larger” pores, respectively. Altner and
Prillinger (1980) reported that s. coeloco-
nica are associated with chemo-, thermo-,
or hygroreception. In both F. parvus and A.
mellifera (Esslen and Kaissling 1976) the pit
organs lie in a zone along the lateral and
medial surfaces of the antenna. In E. parvus
there are two different-sized and shaped ap-
ertures, indicating perhaps two functions.
Campaniform sensilla have a flexible
socket which, when distorted, exerts pres-
sure on an internal mechanoreceptor. They
therefore act as proprioceptors responding
to exocuticular stresses. These sensilla are
frequently concentrated near joints or on
structures subject to cuticular distortion
(McIver 1975). In E. parvus they are found
typically in groups of 1-4 at the middle and
distal parts of the flagellomeres in close as-
sociation with the pit organs located on the
medial and lateral surfaces of the flagellum,
at which points they would be effective as
proprioceptors. The antennal musculature
extends only to the first segment of the fla-
gellum, and this might explain the rather
high number located on flagellomere 1. Sen-
437
silla campaniformia on the antennae of Col-
letidae (Agren 1977) and Apis mellifera
(Dietz and Humphreys 1971, Esslen and
Kaissling 1976) have patterns of distribu-
tion similar to those of E. parvus, but struc-
turally have a more pronounced central
node. The campaniform sensilla of Anoplius
tenebrosus (Alm and Kurczewski 1982) are
similar in structure and distribution to those
of EF. parvus but Walther (1979) described
s. campaniformia of 4. viaticus to be dis-
tributed singly, not in groups.
The s. trichodeum A of Evagetes parvus
looks similar to the s. trichodeum A | of
Anoplius tenebrosus (Alm and Kurczewski
1982), trichodeum A of Prosopis communis
(Nylander) (Colletidae) (Agren 1977), and
the sicula-type sensilla of Odontomachus
ruginodis (Wheeler) (Formicidae) (Callahan
1975). They bend sharply over the antennal
surface, and are broadest along the axis per-
pendicular to the surface. Sensilla trichodea
A are distributed mainly on the lateral sur-
face of the flagellum in FE. parvus and A.
tenebrosus (Alm and Kurczewski 1982). The
trichodeum A sensillum of E. parvus has an
asymmetrical socket which appears to be
flexible in only one plane or direction. The
hole in the center of the broken sensillum
(Fig. 5b) indicates a chemoreceptor with a
sensory dendrite extending the length of the
hair-like sensillum. Although pores were not
seen on the s. trichodea A of FE. parvus the
sensillum, because of its location, comes into
contact with many different substances and
substrates and may function as a chemo-
receptor.
The thin, curved s. trichodea B are evenly
distributed around the flagellum of E. par-
vus. They are usually faintly grooved but
become thickened and deeply grooved on
the dorsum and among the thick and spi-
ralled setae. The s. trichodea B of FE. parvus
have no visible articulating membrane and,
therefore, are probably not mechanorecep-
tors.
The s. trichodea C of E. parvus appear
identical to the s. basiconica of Anoplius
438
tenebrosus (Alm and Kurczewski 1982) and
the s. chaetica of Apis mellifera (Whitehead
and Larson 1976). Because they extend
above the other sensilla, the long trichodea
C at the distal end of each flagellomere may
tactilely sense the adjacent segment.
There are few s. trichodea D in E. parvus
and these are found on the scape and ped-
icel. They resemble the s. trichodea C, ex-
cept that they have a ring of cuticle sur-
rounding the base, rather than a broad open
socket. The collar forms a narrow socket
which may or may not be flexible. The dou-
ble chamber observed in the broken sensil-
lum (Fig. 8b) indicates the presence of a
mechanoreceptor and a chemoreceptor, al-
though it would be difficult to explain why
a sensillum located only on the segments
closest to the head would be chemosensory.
The sensillum basiconicum of E. parvus
is stout, blunt, grooved longitudinally, and
set into a broad and flattened socket. The
basiconic pegs are interspersed among the
corrugated conical sensilla on the ventral
surface of the flagellum. Here they come
into contact with, or close to, surfaces or
objects and therefore may be mechano- and/
or chemoreceptive.
The sensillum spatulatum has not been
described in Hymenoptera. The term “‘spa-
tulata” was derived from its peculiar shape
and a similar sensillum described by Cal-
lahan (1975). This sensillum has a cylin-
drical base and the distal half fans out into
a lightly grooved, concave shell. The socket
is broad and flattened and nearly indistin-
guishable from the socket of a sensillum
basiconicum. The s. spatulata occur infre-
quently within the corrugated conical zone.
Based on the broad surface area, it is likely
that this sensillum is a chemoreceptor.
ACKNOWLEDGMENTS
We thank R. A. Norton, D. L. Dindal,
and W. A. Coté, Jr., all of SUNY-CESF, for
their constructive criticism of the manu-
script. We also thank A. C. Day and J. J.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
McKeon, N. C. Brown Center for Ultra-
structure Studies, SUNY-CESF, for their
assistance with SEM. K. V. Krombein,
Smithsonian Institution, introduced us to
Walther’s (1979) study of the antennal sen-
silla of aculeate Hymenoptera.
LITERATURE CITED
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(Hymenoptera: Apoidea). Int. J. Insect Morphol.
and Embryol. 6: 137-146.
1978. Flagellar sensilla of two species of An-
drena (Hymenoptera: Andrenidae). Int. J. Insect
Morphol. and Embryol. 7: 73-79.
Alm, S. R. and F. E. Kurezewski. 1982. Antennal
sensilla and setae of Anoplius tenebrosus (Cresson)
(Hymenoptera: Pompilidae). Proc. Entomol. Soc.
Wash. 84: 586-593.
Altner, H. and L. Prillinger. 1980. Ultrastructure of
invertebrate chemo-, thermo-, and hygroreceptors
and its functional significance. Int. Rev. Cytol. 67:
69-139.
Callahan, P. S. 1970. Insects and the radiation en-
vironment. Proc. Tall Timbers Conf. on Ecol. An-
imal Control by Habitat Management 2: 247-258.
1975. Insect antennae with special reference
to the mechanism of scent detection and the evo-
lution of the sensilla. Int. J. Insect Morphol. and
Embryol. 4: 381-430.
Dietz, A.and W. J. Humphreys. 1971. Scanning elec-
tron microscope studies of antennal receptors of
the worker honey bee, including sensilla campan-
iformia. Ann. Entomol. Soc. Amer. 64: 919-925.
Esslen, J. and K.-E. Kaissling. 1976. Zahl und verteil-
ung antennaler Sensillen bei der Honigbeine (Apis
mellifera L.). Zoomorphologie 83: 227-251.
Evans, H.E. 1950. A taxonomic study of the Nearctic
spider wasps belonging to the tribe Pompilini (Hy-
menoptera: Pompilidae). Part I. Trans. Amer.
Entomol. Soc. 75: 133-270.
Evans, H. E. and M. J. West Eberhard. 1970. The
wasps. University of Michigan Press, Ann Arbor.
265 pp.
Evans, H. E., C. S. Lin, and C. M. Yoshimoto. 1953.
A biological study of Anoplius apiculatus autum-
nalis (Banks) and its parasite, Evagetes mohave
(Banks) (Hymenoptera, Pompilidae). J. N.Y.
Entomol. Soc. 61: 61-78.
Evans, H. E. and C. M. Yoshimoto. 1962. The ecol-
ogy and nesting behavior of the Pompilidae (Hy-
menoptera) of the northeastern United States. Misc.
Pub. Entomol Soc. Amer. 3: 67-119.
Kaissling, K.-E. and M. Renner. 1968. Antennale
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Rezeptoren fiir Queen Substance und Sterzelduft
bei der Honigbiene. Z. Vgl. Physiol. 59: 357-361.
Krombein, K. V. 1979. Superfamily Pompiloidea,
pp. 1523-1571. Jn K. V. Krombein, P. D. Hurd,
Jr., D. R. Smith and B. D. Burks, Eds. 1979. Cat-
alog of Hymenoptera in America North of Mexico.
Vol. 2. Apocrita (Aculeata). Washington, DC.,
Smithsonian Instit. Press.
McIver, S. B. 1975. Structure of cuticular mechanore-
ceptors of arthropods. Ann. Rev. Entomol. 20:
381-397.
Norton, W. N. and S. B. Vinson. 1974. Antennal
sensilla of three parasitic Hymenoptera. Int. J. In-
sect Morphol. and Embryol. 3: 305-316.
Richards, O. W.and A.H. Hamm. 1939. The biology
of the British Pompilidae. Trans. Soc. Brit. Ento-
mol. 6: 51-114.
439
Schneider, D. 1964. Insect antennae. Ann. Rev. Ento-
mol. 9: 103-122.
Shier, E.H. 1970. The structure of arthropod chemo-
receptors. Ann. Rev. Entomol. 15: 121-142.
Slifer, E. H. and S. Sekhon. 1960. The fine structure
of the plate organs on the antenna of the honey
bee, Apis mellifera L. Experimental Cell Research
19: 410-414.
Walther, J. R. 1979. Vergleichende morphologische
Bertrachtung der antennalen Sensillenfelder ein-
iger ausgewahlter Aculeata (Insecta, Hymenop-
tera). Z. f. zool. Systematik u. Evolutionforschung
17: 30-56.
Whitehead, A. T. and J. R. Larson. 1976. Ultrastruc-
ture of the contact chemoreceptors of Apis mellif-
era L. (Hymenoptera: Apidae). Int. J. Insect Mor-
phol. and Embryol. 5: 301-315.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 440-457
REVISION OF THE GENUS LORITA BUSCK (LEPIDOPTERA: TORTRICIDAE:
COCHYLINI), WITH A DESCRIPTION OF A NEW SPECIES
MICHAEL G. POGUE
Department of Entomology, U.S. National Museum, NHB-127, Washington, D.C. 20560.
Abstract.—The genus Lorita Busck contains two species at present. The type species,
L. abornana Busck, is now considered a subjective junior synonym of Phalonia scarificata
Meyrick. A new species, L. baccharivora Pogue, is described. For this revision 154 adults,
3 larvae, and 4 pupae were studied. They are described and illustrated for both species,
and the egg is described and illustrated for L. baccharivora. Lorita scarificata occurs in
northeastern Brazil northward to French Guiana, northeastern Venezuela, Antigua, Puerto
Rico, Costa Rica, Mexico, southern California, Florida, and has been introduced to Ha-
waii. Lorita baccharivora occurs in the Florida peninsula, northeast coast of Texas, and
has been released in southeastern Queensland, Australia, for the biological control of sea
myrtle, Baccharis halimifolia L.
Key Words:
A revision of Lorita Busck was necessary
to correct nomenclature of the type species,
and to describe a new species being released
in Australia for biological control of the
weedy shrub, Baccharis halimifolia L. Lo-
rita was proposed by Busck (1939) to ac-
commodate a new species feeding on
dodder, Cuscuta californica Choisy, and
injurious to bell peppers, Capsicum an-
nuum L., being imported from Sinaloa and
Sonora, Mexico. Larvae bore into the stalks
and capsules of the latter, thus reducing their
commercial value (Busck 1939).
Saphenista, Thyraylia, and some Cochy-
lis are very similar in coloration and mac-
ulation, making it difficult to identify them
correctly without genital dissections. Dur-
ing the present study Phalonia scarificata
Meyrick was discovered to be a senior syn-
onym of Lorita abornana and the valid type
species of Lorita. Clarke (1963) illustrated
the lectotypes of galbanea Meyrick and
scarificata and placed them in Lorita. Based
biological control of weeds, Lorita baccharivora, L. scarificata, Baccharis
on these illustrations and examinations of
paratypes of both species, I concluded that
scarificata is a senior synonym of abornana
and galbanea belongs to the genus Saphe-
nista. Forbes (1931) described Saphenista
semistrigata from El Yunque, Puerto Rico,
but illustrated the male genitalia (pl. XLV,
fig. 26) of Lorita scarificata. The type of S.
semistrigata is a female and is not conspe-
cific or congeneric with L. scarificata.
Therefore, Saphenista semistrigata is a val-
id species, and the illustration by Forbes is
a misidentification.
For this revision 153 adults, 3 larvae, and
4 pupae were studied. Means and standard
deviations are given for some measure-
ments. The letter “‘n’’ denotes number of
specimens examined. The supraocular in-
dex equals height of head capsule above
compound eye divided by total height of
head capsule (from top of epicranium to tip
of subgenal process) (Kristensen and Niel-
sen 1979). Valval length was measured from
VOLUME 90, NUMBER 4
44]
Figs. 1-3. Adult moths: 1, Lorita scarificata (Meyrick), 4, forewing length 4.3 mm; 2, L. baccharivora, new
species, 6, holotype, forewing length 4.0 mm; 3, L. scarificata, 6, ventral projection of 6th abdominal sternite.
proximal tip of sacculus to apex of valva,
and maximum width was measured in a
vertical line from valval costa to ventral
edge of sacculus. Color names are followed
by a parenthetical number indicating colors
under the system of Smithe (1975, 1981).
Larval chaetotaxy follows Hinton (1946).
Lorita Busck
Figs. 1-47
Lorita Busck, 1939: 100.—Clarke, 1963:
20.—Razowski, 1977: 256.—Powell,
1983: 42.
Type-Species.—Phalonia scarificata
Meyrick 1917 (Fig. 1), senior synonym of
Lorita abornana Busck.
Adult (Figs. 1-2).— Forewing length 3.4—
5.4 mm.
Head (Fig. 4): Vestiture rough. Eye well
developed. Supraocular index 0.26-0.28.
Ocellus absent. Antenna filiform, scaled
dorsad, setose ventrad, 33-38 segments. La-
bial palpus porrect, scales of second seg-
ment with dorsal expansion greater than
ventral; third segment exposed, not covered
by second segment scaling, sensory setae
distributed along entire length of third seg-
ment; third segment 1.3 x first segment; sec-
ond segment 3.3 first segment, equal to
vertical eye diameter, length 5.2 width.
Maxillary palpus 1-segmented.
Thorax: Posterior crest present, but often
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 46. Head structure and wing venation: 4, L. scarificata (Meyrick), anterior view (scale = 0.5 mm); 5,
L. scarificata; 6, L. baccharivora, new species.
VOLUME 90, NUMBER 4
inconspicuous. Lateral scale tufts of meta-
notum hairlike. Prothoracic leg with epi-
physis 0.4-0.5 length of tibia. Mesothoracic
tibia with a single pair of unequal sized api-
cal spurs, longest 0.5—0.6 length of tibia.
Metathoracic tibia with 2 pairs of unequal
sized spurs; basal pair originating at 0.6 from
tibial base, with longest spur 0.4 length of
tibia; apical pair at subapex with longest
spur 0.4 length of tibia. Leg indices [femur :
tibia : (basitarsus)tarsus] for prothoracic leg
1:0.7:(0.5)1.6; mesothoracic leg 1:0.9:
(0.5)1.1; metathoracic leg 1:1.6:(0.8)1.4.
Forewing (Figs. 5-6): Male length 3.4—4.6
mm; length 2.5-3.1 x maximum width. Fe-
male length 3.8-5.4 mm; length 2.4—3.2 x
maximum width. Costa straight; apex
rounded; termen straight, oblique. Sc less
than 0.5 wing length. R1 originating beyond
middle of discal cell; R2 originating nearer
R3 than R1; R5 ending at termen. M3 and
CuA1 separate. CuA2 originating at 0.67
length of discal cell. CuP absent. Al+2
stalked at 0.4 total length.
Hindwing (Figs. 5-6): Male length 2.9-
4.0 mm; length 2.6-3.3 x maximum width.
Female length 2.14.4 mm; length 2.6-3.4 x
maximum width. Costa straight; apex pro-
duced; termen concave below apex. Sc+R1
less than 0.5 wing length. Rs and M1 stalked
at 0.67 length of M1. M3 and CuA1 con-
nate. CuA2 originating at 0.67 length of dis-
cal cell. Female with 3 frenular bristles.
Abdomen: Male with prominent ventral
projection of 6th sternite (Fig. 3); female
normal.
Male genitalia (Figs. 7-10): Uncus pres-
ent, thin, elongate, or divided. Gnathos ab-
sent. Socii attached to posterior margin of
tegumen. Tegumen well developed, trape-
zoidal. Transtilla a broad, well-developed
band with an elongate, stout, apically
rounded median projection. Valva length
1.7-2.0 maximum width; costa a thin
sclerotized band; sacculus a well defined
band appressed to ventral edge of valva.
Vinculum arms free. Aedoeagus large and
robust, 1.4-1.7= length of valva, bent at
443
apical 0.2 with a sharp ventrally produced
spine, cornuti numerous minute spinules on
vesica.
Female genitalia (Figs. 11-12): Papillae
anales setaceous, thin, apices broadly
rounded, joined by narrow connection.
Length of apophysis anterior 1.0-1.7 x
apophysis posterior. Sterigma well devel-
oped, rectangular, joined to ventral branch
of apophysis anterior. Ductus bursae short,
encircled by a series of elongate, narrow
sclerotizations. Corpus bursae covered with
numerous minute convolutions, signa con-
sisting of many spinules. Accessory bursa
present.
Discussion.—Lorita has a Nearctic dis-
tribution from Florida to California and
northern Mexico. Neotropical distribution
includes Mexico (Razowski 1986, Razowski
and Becker 1986), Costa Rica (Razowski
1986, Razowski and Becker 1986), Puerto
Rico (Forbes 1931), Antigua, British
Guiana, Venezuela, and Brazil (Razowski
1967, Razowski and Becker 1986). This is
the only cochyline genus known from the
Pacific Ocean islands; L. scarificata has been
introduced to Hawaii on the islands of Oahu
(Beardsley 1977) and Kauai (Mau 1979).
Males of Lorita can be distinguished from
those of other cochyline genera by the elon-
gate ventral process of the 6th sternum (Fig.
3). I have found Lorita mixed with speci-
mens of Spinipogon (Cochylini) in collec-
tions.
Lorita scarificata (Meyrick)
Figs. 1, 3-5, 7-8, 11, 13-25
Phalonia scarificata Meyrick, 1917: 3.
Saphenista semistrigata; Forbes, 1931: 355
[misidentification, pl. XLV, fig. 26, male
only].
Lorita abornana Busck, 1939: 101.—Com-
stock, 1939: 119.—Razowski, 1967:
202.—Beardsley, 1977: 391.—Mau, 1979:
4. NEW SYNONYM.
Lorita abornana chatka Busck, 1939: 101.
NEW SYNONYM.
444 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-12. Male and female genitalia: 7, L. scarificata (Meyrick), ventral view; 8, lateral view of aedoeagus;
9. L. baccharivora, new species, ventral view; 10, lateral view of aedoeagus; 11, L. scarificata, ventral view; 12,
L. baccharivora, ventral view (scale = 0.2 mm).
VOLUME 90, NUMBER 4
Lorita scarificata (Meyrick), Clarke 1963:
20.
Adult (Fig. 1).—A small moth with pale
horn (92) (color slightly less yellow than
cream (54)) ground color that reflects light,
giving a shiny appearance in fresh speci-
mens. An indistinct submedian band, a
smaller postmedian spot, and darker ter-
minal band mark the forewing. Coloration
and markings are variable throughout its
range.
Male: Forewing length 3.5-4.5 mm (Xx =
4.2 + 0.4 mm, n= 8).
Head (Fig. 4): Labial palpus pale horn
color; scales of middle segment expanded
dorsad, extending to middle of apical seg-
ment. Antennal scape pale horn, with a few
dusky brown scales around base, rest of an-
tenna pale horn with a few dusky brown
scales at base. Front and vertex pale horn.
Thorax: Mesonotum and tegula pale horn
with scattered darker scales along anterior
edge. Underside shining white; pro- and
mesothoracic legs with femur pale horn
having scattered dusky brown (19) scales
along posterior edge, tibia with 4 bands:
starting at apical end, bands 1| and 3 speck-
led pale horn and dusky brown, 2 and 4 pale
horn color, tarsi speckled pale horn and
dusky brown with pale horn apical rings;
metathoracic leg pale horn lightly scattered
with drab (27) scales.
Forewing (Figs. 1, 5): Length 2.6-3.0 x
maximum width. Ground color pale horn
with iridescence in fresh specimens; indis-
tinct submedian band clay (123B), indis-
tinctly bordered along proximal and distal
margins with a few dusky brown scales,
dusky brown scales more concentrated at
proxmial margin; a small indistinct clay-
colored postmedian spot just above tornus;
indistinct terminal band extending from
costa to termen clay-colored; small dusky-
brown scale patches evenly spaced along
costa from base to apex. Fringe yellow ocher
(123C) above and below. Underside dark
drab (119B) with distinct pale horn scale
445
patches evenly spaced along costa from base
to apex.
Hindwing (Figs. 1, 5): Length 2.9-3.9 mm
(Xx = 3.6 + 0.4mm, n = 8); length 2.9-3.3 x
maximum width. Costal fold atrophied; light
drab (119C). Underside white with light drab
scaling along costa, termen, and vein M2.
Fringe light drab with slightly darker basal
band above and below.
Abdomen: Pale horn above and below.
Male genitalia (Figs. 7-8): Uncus present,
thin, apex truncate. Socil attached apically
to tegumen, bases fused dorsally, basal 0.5
forming thumblike lobes that lack setae,
apical 0.5 bulbous and setose. Transtilla a
broad, well-developed band with an elon-
gate, stout, apically rounded median pro-
jection; median projection flanked by 2
broad, apically concave projections; these
lateral projections are 0.5 length of median
projection and extend to just below base of
socu. Valva length 1.8-2.0x maximum
width; costa forming a right angle at valval
base, continuing in a concave arc to round
apex; sacculus a thin well-defined band ap-
pressed to ventral edge of valva. Vinculum
arms free as denoted by median suture. Ae-
doeagus large, base round, apex truncate,
with ventrally produced spine, 1.4—-1.6 x
length of valva, length 3.1-3.6 x maximum
width; cornuti consisting of numerous mi-
nute spinules on vesica.
Female: Coloration and markings as in
male. Length of forewing 3.9-5.2 mm (X =
4.5 + 0.5 mm, n = 8); length 2.7-2.9 x
maximum width. Length of hindwing 3.2-
4.4 mm (xX = 3.8 + 0.5 mm, n = 8); length
2.9-3.2 x maximum width.
Female genitalia (Fig. 11): Papillae anales
setaceous, thin, slightly curved, apices
broadly rounded, joined by narrow connec-
tion. Length of apophysis anterior 1.0-1.7 x
apophysis posterior. Sterigma rectangular,
poorly developed, a median lightly sclero-
tized oval plate surrounding ostium. Col-
liculum well sclerotized, quadrate. Ductus
bursae distinct, basal 0.67 sclerotized. Cor-
pus bursae with fine convolutions, many
446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 13-19. L. scarificata (Meyrick), larval chaetotaxy: 13, lateral view of prothorax, mesothorax, and
abdominal segments 1, 6, 8, and 9; 14, dorsal view of head; 15, lateral view of head (scale = 0.2 mm); 16, right
mandible (scale = 0.1 mm); 17, labrum, dorsal view; 18, labrum, ventral view (scale = 0.1 mm); 19, dorsal
view of abdominal segments 8-10 (scale = 0.5 mm).
VOLUME 90, NUMBER 4
minute spinules. Accessory bursa originat-
ing ventrally, just below middle of ductus
bursae. Ductus seminalis from proximal half
of corpus bursae.
Larva (Figs. 13-19).—Length of last in-
star 4.8-6.4 mm (X = 5.6 + 1.1 mm, n =
2).
Head: Hypognathous, maximum width
0.6 mm. Color burnt sienna (132) to clay
(123B) in alcohol. Puncture Pb closer to P2
than Pl. Puncture Pa equidistant between
P2 and L1. Puncture Ga dorsal and anterior
to Gl. Six stemmata present with 2,3 and
4,5 contiguous. Mandible with 5 cusps, first
truncate. Labrum with 6 pairs of dorsal se-
tae, 3 pairs of ventral epipharyngeal setae.
Thorax: Prothoracic shield concolorous
with head. Prothorax with L group on same
pinaculum; mesothorax with L3 on sepa-
rate, large pinaculum. Prothorax with SV1
and SV2 on same pinaculum; mesothorax
with SV2 absent. Legs 3-segmented, single
tarsal claw.
Abdomen: SD2 minute, and present only
on segment 8. Ninth segment with D1 and
SD1 on same pinaculum; L group consists
of 2 setae, L] and L2. SV3 present only on
segment 6; SV1 and SV3 absent on segment
9. Prolegs on segments 3-6 and 10; crochets
on abdominal segments 3-6 uniordinal and
arranged in a complete circle; anal prolegs
with crochets uniordinal and arranged in a
semiellipse.
Pupa (Figs. 20-25). — Male length 5.6 mm
(n = 1); female length 5.7 mm (n = 1) (in
alcohol). Eyes clearly visible. Anterior group
of spines on abdominal segments 2-8 in male
and 2-6 in female consisting of 2 rows, the
anterior row having fewer and smaller spines
than second row. Segment 9 in male and
segments 8-9 in female consisting ofa single
row of slightly larger spines. Posterior row
of spines minute and on segments 2-8 in
male and 2-7 in female. Spiracles peg-
shaped. A single saggitate genital slit on an-
terior edge of segment 9, another similarly
shaped slit in middle of segment 10. Female
with 3 genital slits, one on anterior edge of
447
segment 8, a second on anterior edge of seg-
ment 9, and a third on a pad in middle of
segment 10. Cremaster with 4 hooks on an-
terior edge and a pair of lateral hooks from
spines on ventral side of segment 10; 3 pairs
of lateral hooks and one pair of caudal hooks
on edge of segment 10.
Types.—Lectotype (Phalonia scarificata
Meyrick), designated by Clarke (1963), 2,
“Bartica, British Guiana, Parish, .12.12.,”
genitalia slide No. 6284, in British Museum
(Natural History). Holotype (Lorita abor-
nana), °, U.S.A., California, Los Angeles
Co., El Segundo, emerged 11 July 1938, W.
D. Pierce, larva on Cuscuta californica, U.S.
National Museum type no. 53250, genitalia
slide USNM 23826, in U.S. National Mu-
seum, Smithsonian Institution, Washing-
ton, D.C.
Material examined.—BRITISH GUI-
ANA: Bartica, Parish .1.13, 1 2 genitalia
slide USNM 23726. “Mallali, Parish .3.13,”
1 3, genitalia slide USNM 23725. BRITISH
WEST INDIES: ANTIGUA: Flat Top
Point, 13 Apr. 1958, J. F. G. Clarke (Smith-
sonian-Bredin Exped.), 2 4; English Harbor,
20 Apr. 1958, J. F. G. Clarke (Smithsonian-
Bredin Exped.), 2 4; St. Anns Hill, 21 Apr.
1958, J. F. G. Clarke (Smithsonian-Bredin
Exped.), | 6. MEXICO: 30 Jan. 1942, in
bell peper fruit, Nogales No. 51068, Lot No.
42-1560, 1 2°. SINALOA: 12 Jan. 1939, in
bell pepper, Nogales No. 28970, Lot No.
39-1430, 1 4; 20 Jan. 1939, in bell pepper,
Nogales No. 28978, Lot No. 39-1464, 1 2;
19 Jan. 1939, in bell pepper, Nogales No.
28990, Lot No. 39-1807, 1 6; 24 Jan. 1939,
in bell pepper, Nogales No. 28994, Lot No.
39-1873, 1 6; March 1939, from bell pepper,
1 6; San Blas, iss[ued] (=emerged) 3 March
1939, Nogales No. 29358, on bell pepper,
1 6. SONORA: 21 Jan. 1939, in bell pepper,
Nogales No. 28971, Lot No. 39-1463, 1 2:
21 Jan. 1939, in bell pepper, Nogales No.
29098, Lot No. 39-2240, 1 6. PUERTO
RICO: Coamo Springs, 4 Apr. 1930, Cor-
nell Univ., Lot No. 795, Sub 15, | 2, gen-
italia slide USNM 23785. UNITED
448 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
22 25
Figs. 20-25. L. scarificata (Meyrick), pupal structure: 20, 6 ventral view; 21, 6 dorsal view (scale = 1.0 mm);
22, 4 ventral view of terminal segments (scale = 0.5 mm); 23, 2 ventral view; 24, 2 dorsal view (scale = 1.0
mm); 25, 2 ventral view of terminal segments (scale = 0.5 mm).
STATES: CALIFORNIA: Los Angeles Co.: italia slide USNM 23826 [holotype of Lor-
El Segundo, emdg. 11 July 1938, larva on ita abornana Busck]; emdg. (=emerged) 13
Cuscuta californica, W. D. Pierce, 1 2, gen- July 1938, 1 4, genitalia slide USNM 23856;
VOLUME 90, NUMBER 4
emdg. 14 July 1938, 1 9, genitalia slide
USNM 23856; emdg. 15 July 1938, 1 2,
wing slide USNM 23856, genitalia slide
USNM 23856. San Diego Co.: Del Mar, 12
Aug. 1959, R. A. Mackie Coll., 1 4; 19 Aug.
1959, 1 9; 26 Aug. 1959, 1 9, genitalia slide
MGP 637; 2 Sep. 1959, 1 9; Mission Bay,
10 June 1959, 1 6. FLORIDA: Collier Co.:
Chokoloskee, 19 4, genitalia slides USNM
23126, USNM 23163, 12 2, genitalia slide
USNM 23125. Dade Co.: Homestead, 22
Oct. 1959, D. O. Wolfenbarger, 1 2; 29 Oct.
1959, 1 6, SEM slide USNM 23465; 2 Nov.
1959, 3 2, genitalia slide USNM 23219; 4
Nov. 1959, | ¢. Escambia Co.: Pensacola,
23 Sep. 1961, Shirley Hills, | 2. Highlands
Co.: Archbold Biol. Sta., S. W. Frost, 1 4.
Manatee Co.: Bradenton, Gulf Coast Exp.
Sta., 24 March 1955, E. G. Kelsheimer, |
6. Monroe Co.: Key Largo Key, | Nov. 1964,
Mrs. Spencer Kemp, | 2, wing slide USNM
23130; 3 Jan. 1967, 1 9; 3 Jan. 1968, 2 2; 7
Jan. 1968, 2 4, genitalia slides USNM 23164,
USNM 23850; 22 Jan. 1968, 2 2; 3 Nov.
1967, 1 2; 24 Nov. 1967, 1 6; 19 Dec. 1967,
1 2; Key Largo, 19 Oct. 1964, Mrs. Spencer
Kemp, | ¢, 2 2. Sarasota Co.: Siesta Key, 3
Jan. 1960, C. P. Kimball, | 4; 11 Jan. 1960,
1 6; 14 Jan. 1960, 1 6; 13 March 1953, 1 6;
27 March 1960, | 2; 2 April 1964, 1 4; gen-
italia slide USNM 23131; 3 April 1960, 2
4; 13 April 1960, 1 4; 29 April 1960, 1 9; 3
May 1960, 1 4, 1 9; 4 May 1960, 2 4; 13
May 1960, | 4, wing slide USNM 23846;
21 Nov. 1953, 192 10) Dec: 19535 1 6219
Dec. 1959, 1 2, genitalia slide USNM 23218;
24 Dec. 1953, 1 4, genitalia slide USNM
23162. HAWAII: Oahu: Waianae, Dec.
1974, J. W. Beardsley collector, reared on
Chrysanthemum blossom, 2 4, genitalia slide
USNM 24208; Ewa, Oct. 1974, J. W.
Beardsley collector, light trap, | 4; Hono-
lulu, Mar. 1974, J. W. Beardsley collector,
light trap, 1 4, genitalia slide USNM 23751.
VENEZUELA: BOLIVAR: 5 km E. Tum-
eremo, 12 Feb. 1976, C. M. & O. S. Flint,
Jr., 1 2, genitalia slide USNM 25452; Cd.
Guayana, Rio Caroni, Parque Llovizna, 13
Feb. 1976, C. M. & O. S. Flint, Jr., 1 ¢.
449
Host.—Several hosts have been recorded
for L. scarificata. This species is a pest on
bell peppers with larvae boring into the stalks
and capsules, which may reduce their com-
mercial value (Busck 1939). Capsicum is
also native to the Neotropics and West In-
dies and has been cultivated since pre-Inca
times (Hill 1952). Comstock (1939) and
Busck (1939) also report dodder as a food
plant of L. scarificata in the sand dune area
of El Segundo, Los Angeles Co., California.
This species has also been reared from
Chrysanthemum (Compositae) blossoms in
Waianae, Oahu, Hawaii.
Distribution (Maps 1-—2).—Lorita scari-
ficata is widely distributed in the southern
Nearctic and Neotropical Regions and has
been introduced into Hawau. It has been
collected in California, Texas, Florida,
Mexico, Puerto Rico, Antigua, Venezuela;
Razowski (1967) reported it from Brazil.
Discussion.—This species is variable in
coloration. Darker forms have the forewing
markings dark drab (1 19B) with more dusky
drab scales along their margins, or the fore-
wings are covered with dusky brown scales
with barely any evidence of banding. These
darker forms are found within the same se-
ries as the lighter form described above.
Genitalia do not vary within the sexes.
Males of L. scarificata can be separated
from those of L. baccharivora by: 1) lack of
a hindwing costal fold, 2) hindwing vein Sc
longer, 3) uncus thin, elongate, 4) socius
fused to base of uncus, and 5) thumblike
lobes lack setae and are attached laterally
to tegumen. Female genitalia are differen-
tiated by: 1) sterigma less heavily sclero-
tized, with a median oval plate surrounding
ostium basally, 2) presence of a well-devel-
oped colliculum, and 3) ductus bursae more
slender and distinct from corpus bursae.
Lorita baccharivora, new species
Figs. 2, 6, 9-10, 12, 26-47
A small moth with forewing ground color
pale horn (92) having a darker median cos-
tal spot and terminal band (Fig. 2).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
450
*(satdads Mau ‘D4OALMDYIIDG “T
= 9[BURL {(YOUAIJA)) DIVIYIUDIS “T = 9[OIID) BOLVIDUTY [BNUID puke YLION UI MOT JO UONNQMISIq
NOILD3FO8d W3YV-1VNDO3 TWHILNNIZY Ly3a8WVv7
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saw
VOLUME 90, NUMBER 4
Map 2.
Male: Forewing length 3.7-4.5 mm (x =
4.0 + 0.3 mm, n = 8).
Head: Labial palpus pale horn, speckled
with dusky brown (19); scales of middle seg-
ment expanded dorsad, extending to middle
of apical segment. Antennal scape pale horn,
with a few dusky brown scales around base,
rest of antenna pale horn with a few dusky
brown scales at base. Front and vertex pale
horn scattered with dusky brown scales.
Thorax: Mesonotum and tegula varying
from pale horn with indistinct anterior, me-
dian, and posterior bands of dusky brown
to drab (27). Underside shining white; pro-
thoracic coxa and femur dusky brown with
scattered pale horn scales, tibia dusky brown
with a median band of pale horn, tarsi dusky
brown with apical ring of pale horn: meso-
thoracic femur pale horn with scattered
dusky brown scales, tibia and tarsi as in
prothoracic leg; metathoracic leg pale horn
with scattered dusky brown scales, tarsal
color lighter than in other legs.
°
200 400 600 800 1000km
La eS a ee
9 100 200 300 400 500 600 m,
Distribution of L. scarificata (Meyrick) in South America.
Forewing (Figs. 2, 6): Length 2.5-2.9 x
maximum width. Ground color pale horn
with median costal spot extending to an-
terior margin of discal cell, and dusky brown
terminal band extending from costa to ter-
men; remainder of wing heavily suffused
with dusky brown scales. Fringe drab. Un-
derside and fringe grayish horn (91).
Hindwing (Figs. 2, 6): Length 3.1-4.0 mm
(x = 3.1-4.0 + 0.3 mm, n = 8); length 2.8-
3.2 x maximum width. Costal fold present,
enclosing hair pencils; dirty white becoming
smoke gray (45) toward apex. Underside
dirty white with dusky brown striations in
costal half. Fringe dirty white with smoke
gray basal band above and below.
Abdomen: Dirty white above and below.
Male genitalia (Figs. 9-10): Uncus bifid.
Socius fused from basal 0.5 of uncus to mid-
dle of tegumen. Transtilla a well developed
band with a broad median projection ex-
tending to base of uncus. Valval length 1.7-
2.0 maximum width; costa sinuate; elon-
452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 26-30. L. baccharivora, new species, egg and larval structure: 26, dorsal view,
«150: 28, dorsal surface detail,
x 150; 27, ventral view,
«700: 29, foreleg, x 600; 30, crochets of abdominal segment 6, x 600.
VOLUME 90, NUMBER 4
453
Figs. 31-34.
maxilla,
gate spatulate scales arising from costal half
of valva and extending from base to apex;
apex sharply rounded; sacculus well devel-
oped, extending along entire ventral edge of
valva. Vinculum arms free. Aedoeagus | .5—
1.7 length of valva, length 2.7-3.3 x max-
imum width; cornuti numerous minute spi-
nules on vesica.
Female: Markings of forewings similar,
but cinnamon (123A) in color. Hindwing
coloration as in male. Length of forewing
4.0-4.5 mm (x% = 4.3 + 0.2 mm, n = 8);
length 2.6-2.9* maximum width. Length
L. baccharivora, new species, larval mouthpart structure: 31, ventral view, x 300; 32, tip of
< 2200; 33, labium, ventral view, x 1200; 34, tip of labial palp, x 6000.
By) as 0)
maximum
of hindwing 3.4-3.9 mm (x =
mm, n = 8); length 2.7-3.2 x
width.
Female genitalia (Fig. 12): Papillae anales
setaceous, elongate. Length of apophysis an-
terior 1.0Q—1.1 x apophysis posterior. Sterig-
ma well sclerotized, quadrate, minute spi-
nules medially, just posterior to ostium.
Colliculum absent. Ductus bursae short,
broad, gradually merging with corpus bur-
sae. Corpus bursae convoluted, covered with
many minute spinules at juncture of ductus
bursae. Accessory bursa present, from mid-
454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
s °
S
=
Figs. 35-41. L. baccharivora, new species, larval chaetotaxy: 35, lateral view of prothorax, mesothorax, and
abdominal segments 1, 6, 8, and 9; 36, dorsal view of head; 37, lateral view of head (scale = 0.1 mm); 38, right
mandible (scale = 0.1 mm); 39, labrum, dorsal view; 40, labrum, ventral view (scale = 0.1 mm); 41, dorsal
view of abdominal segments 8-10 (scale = 0.5 mm).
VOLUME 90, NUMBER 4
455
Figs. 42-47. L. baccharivora, new species, pupal structure: 42, 6 ventral view; 43, é dorsal view (scale = 1.0
mm); 44, 4 ventral view of terminal segments (scale = 0.5 mm); 45, 2 ventral view; 46, 2 dorsal view (scale =
1.0 mm); 47, ° ventral view of terminal segments (scale = 0.5 mm).
dle of ductus bursae. Ductus seminalis from
proximal end of corpus bursae.
Egg (Figs. 26-28).— Length 0.6 mm. Dor-
sal side sculptured with 4- and 5-point star
shapes; ventral side smooth. Laid singly on
leaves of host.
Larva (Figs. 29-41).—Maximum length
of last instar 7.3 mm, n= |.
Head: As in L. scarificata, except color
raw umber (223); puncture Pb closer to P2
than P1, but less so than in former species;
first cusp of mandible pointed.
456
Thorax: As in L. scarificata, except me-
sothorax with L3 on smaller pinaculum.
Abdomen: Asin L. scarificata, except SD2
absent from segment 8; SV1 present on seg-
ment 9.
Pupa (Figs. 42-47).—As in L. scarificata,
except anterior group of spines minute and
in a single row on abdominal segment 2;
anterior group of spines on abdominal seg-
ments 3-8 in male and 3-7 in females con-
sisting of 2 rows, the anterior row having
fewer and smaller spines than second row;
segment 10 in male and segments 9-10 in
female consisting of a single row of spines;
posterior row of spines minute and on seg-
ments 2-7 in both sexes. Male with a single
genital slit on a pad in middle of segment
9, another slit in middle of segment 10. Fe-
male with 3 genital slits, in middle of seg-
ment 9, a second on anterior edge of seg-
ment 10, anda third longer slit just posterior
to middle of segment 10. Cremaster with 4
hooks on anterior edge and a pair of lateral
hooks from spines on ventral side of seg-
ment 10; 3 pairs of lateral hooks and one
pair of caudal hooks on edge of segment 10.
Holotype.—é, genitalia slide USNM
23928, Lake Okeechobee, Okeechobee Co.,
Florida, 27°N, 81°W, September, 1984, leg.
D. Green, laboratory reared, Brisbane, Aus-
tralia, leg. W. A. Palmer. Type in the U.S.
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
Paratypes. UNITED STATES: FLOR-
IDA: 1967, W. Haseler, | 4, genitalia slide
MGP 923. Dade Co: Homestead, | April
1958, 1 6, 22 April 1959, 1 6, 8 February
1959, 1 2, 10 April 1959, 1 2, D. O. Wol-
fenbarger. Glades Co.: Lakeport, 26°59’N,
81°06’W, 5 October 1984, leg. D. Green, ex
larva, Baccharis halimifolia L., | 4, genitalia
slide USNM 23755, 1 2. Highlands Co.: Lake
Placid, Archb. Biol. St., 10-11-67, W. H.
Haseler, ex foliage Baccharis halimifolia, |
4, genitalia slide MGP 924, Parker Is., 26-
29 May 1964, R. W. Hodges, 2 6. Monroe
Co.: Tavernier, 25°00'N, 80°28’W, 9 May
1985, leg. W. A. Palmer, 85129-2-1, ex lar-
va, Baccharis halimifolia L., 2 8, 2 2, geni-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
talia slide USNM 23862. Okeechobee Co.:
Lake Okeechobee, 27°N, 81'W, Sep. 1984,
D. Green leg., laboratory reared Brisbane,
Australia, W. A. Palmer leg., 9 3, genitalia
slides USNM 23860, 23861, 23863, 4 9,
genitalia slide USNM 23859. TEXAS: Har-
ris Co.: Houston, 15 May 1979, 1 6, USNM
23930; 19 August 1966, A. & M. E. Blan-
chard, 2 4, genitalia slide USNM 23931.
Clear Lake City, Armand Bayou, 20 May
1986, Coll. P. E. Boldt, reared from leaves
of Baccharis halimifolia, | 6. Deposited in
USNM.
Host.—Baccharis halimifolia L., Com-
positae, sea myrtle.
Distribution (Map 1).—Known from
south central to southern Florida, and east-
ern Texas.
Discussion.— This species is smaller and
the males are darker than in L. scarificata.
Males are easily recognized by the promi-
nent costal fold on the hindwing, which is
atrophied in L. scarificata. Male genitalia of
L. baccharivora can be separated from those
of L. scarificata by: 1) uncus bifid, 2) valva
with sinuate costa, apex pointed, and elon-
gate spatulate scales arising from costal half,
3)a more developed sacculus, and 4) a wider
aedoeagus. Female genitalia can be differ-
entiated by: 1) papillae anales more elon-
gate, 2) sterigma more heavily sclerotized
and quadrate with a median patch of minute
spicules above ostium, and 3) colliculum
absent.
ACKNOWLEDGMENTS
I thank W. A. Palmer of the North Amer-
ican Field Station, Queensland Department
of Lands, Temple, Texas for the specimens
of Lorita baccharivora and J. K. Liebherr
of Cornell University for the loan of the
holotype of Saphenista semistrigata. For re-
viewing the manuscript, I thank J. F. G.
Clarke and D. R. Davis, of the National
Museum, Smithsonian Institution, Wash-
ington, D. C., R. W. Hodges, Systematic
Entomology Laboratory, BBII, % National
Museum of Natural History, Smithsonian
Institution, Washington, D. C., and W. E.
VOLUME 90, NUMBER 4
Miller, Department of Entomology, Uni-
versity of Minnesota, St. Paul, Minnesota.
LITERATURE CITED
Beardsley, J. W. 1977. Notes and exhibitions: Lorita
abornana Busck. Proc. Hawaiian Ent. Soc. 22: 391.
Busck, A. 1939. A generic review of the family Phal-
oniidae with descriptions of two new genera and
one new species. Bull. S. Calif. Acad. Sci. 38: 98—
HUT:
Clarke, J.F.G. 1963. Catalogue of the type specimens
of Microlepidoptera in the British Museum (Nat-
ural History) described by Edward Meyrick, Vol.
IV. London, 521 pp.
Comstock, J. A. 1939. Notes on the larva of Lorita
abornana Busck (Lepidopt.). Bull. S. Calif. Acad.
Sci. 38: 119.
Forbes, W. T. M. 1931. Supplementary report on the
Heterocera or moths of Porto Rico. J. Dep. Agric.
Porto Rico 4: 339-394.
Hill, A. F. 1952. Economic Botany. McGraw-Hill
Book Co., Inc. New York, 560 pp.
Hinton, H. E. 1946. On the homology and nomen-
clature of the setae of lepidopterous larvae, with
some notes on the phylogeny of the Lepidoptera.
Trans. Roy. Entomol. Soc. London, 97: 1-37.
Kristensen, N. P. and E. S. Nielsen. 1979. A new
subfamily of micropterigid moths from South
America. A contribution to the morphology and
phylogeny of the Micropterigidae, with a generic
457
catalogue of the family (Lepidoptera: Zeuglop-
tera). Steenstrupia 5: 69-147.
Mau, R. 1979. Notes and exhibitions: Lorita abor-
nana Busck. Proc. Hawaiian Ent. Soc. 23: 4.
Meyrick, E. 1917. Descriptions of South America
Micro-Lepidoptera. Trans. R. Entomol. Soc. Lond.
1917: 1-52.
Powell, J. A. 1983. In R. W. Hodges, et al., eds.,
Check List of the Lepidoptera of America North
of Mexico. London, 284 pp.
Razowski, J. 1967. South American Cochylidae
(Lepidoptera) from the collection of the British
Museum (Natural History). Acta Zool. Cracov. 12:
163-210.
1977. Catalogue of the generic names used
in Tortricidae (Lepidoptera). Acta Zool. Cracov.
32: 207-295.
1986. New and little known Neotropical Co-
chylidil (Lepidoptera, Tortricidae). Acta Zool.
Cracov. 29: 373-396.
Razowski, J. and V. O. Becker. 1986. Cochylidii
(Lepidoptera, Tortricidae) collected in Central
America and Mexico. Acta Zool. Cracov. 29: 441—
500.
Smithe, F. B. 1975. Naturalist’s Color Guide. The
American Museum of Natural History, New York,
New York, 8 color plates.
. 1981. Naturalist’s Color Guide, Part III. The
American Museum of Natural History, New York,
New York, 37 pp., 9 color plates.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988. pp. 458-461
THE HOST SPECIFICITY AND BIOLOGY OF
ARISTOTELIA IVAE BUSCK (GELECHIIDAE) AND
LORITA BACCHARIVORA POGUE (TORTRICIDAE), TWO
MICROLEPIDOPTERA SELECTED AS BIOLOGICAL CONTROL
AGENTS FOR BACCHARIS HALIMIFOLIA (ASTERACEAE) IN AUSTRALIA
G. DIATLOFF AND W. A. PALMER
(GD) Alan Fletcher Research Station, Queensland Department of Lands, 27 Magazine
St., Sherwood, Queensland 4075, Australia; (WAP) North American Field Station,
Queensland Department of Lands, 2801 Arrowhead Circle, Temple, Texas 76502.
Abstract. —The host specificities and biologies of two microlepidoptera, Aristotelia ivae
Busck (Gelechiidae) and Lorita baccharivora Pogue (Tortricidae) were determined prior
to their utilization for the biological control of the noxious weed Baccharis halimifolia.
Both species were multivoltine foliage feeders with generation times of approximately 6
weeks. Host specificity was assessed by the ovipositional preference and ability of larvae
to feed on 65 plant species. Aristotelia ivae oviposited all but 2 eggs on B. halimifolia and
larvae developed only on this plant. Lorita baccharivora oviposited all but two eggs on
B. halimifolia and larvae also only developed on this plant. It was concluded that both
species were host specific to Baccharis. Permission for their introduction into Australia
was obtained. Both species were released in Australia and establishment of A. ivae has
been confirmed.
Key Words:
Following its introduction into Queens-
land, Australia, before 1900, the North
American shrub Baccharis halimifolia L.
(Asteraceae: Astereae: Baccharineae) be-
came a serious weed in SE Queensland and
NE New South Wales by invading pastures
and land cleared for reforestation. The plant
was declared noxious in 1951 and subse-
quently a biological control program to find
and introduce suitable host specific insects
from the New World was implemented. This
program consisted, in essence, of intensive-
ly surveying appropriate areas, selecting
stenophagous species from available knowl-
edge, testing the host range of these species
experimentally and, if their host range were
limited to Baccharis, mass rearing and re-
leasing the insects in Australia.
Aristotelia, Lorita, Baccharis, biological control, host specificity
Two foliage feeding microlepidoptera Ar-
istotelia ivae Busck (Gelechiidae) and Lorita
baccharivora Pogue (Tortricidae) were found
infesting B. halimifolia along the eastern
seaboard of the United States (Palmer and
Bennett 1988). Literature reviews and
consultations with relevant experts indicat-
ed that they might be host specific to Bac-
charis. Furthermore their occurrence in
Florida was thought advantageous as this
state has a climatic similarity to Queens-
land.
The genus 4ristotelia Huebner contains
39 species none of which is considered an
agricultural pest (Arnett 1985). Busck (1904)
described both the moth and larva of A. ivae
from material collected at Palm Beach,
Florida and reared by H. Dyar in 1900. Two
VOLUME 90, NUMBER 4
other species have been recorded from Bac-
charis. Tilden (1951) described the life his-
tory of A. argentifera Busck which has B.
pilularis DC. and the closely related Eri-
cameria ericoides (Lessing) as hosts. Re-
cently we found a very similar undescribed
species (R. Hodges pers. com.) on B. pilu-
/aris that might also be host specific to Bac-
charis.
The genus Lorita Busck, recently revised
by Pogue (1988), contains two species, L.
baccharivora and L. scarificata (Meyrick).
L. scarificata is a widely distributed,
polyphagous species that is a pest of bell
peppers, Capsicum annuum L. (Solanaceae)
(Pogue 1988).
This paper describes the host specificity
studies undertaken to ascertain the host
range of these insects. Before permission
could be sought to introduce them into Aus-
tralia, it was necessary to demonstrate that
they were specific to Baccharis and that no
native or commercial plant species in Aus-
tralia would be endangered. In the course
of these studies the biologies of the insects
were observed and these are also reported.
BIOLOGY
The following descriptions are based on
our laboratory and field observations of the
various life stages.
Aristotelia ivae.—Eggs which are 25 mm
in length, oval in shape and greenish-white
with characteristic orange markings in col-
or, were Oviposited in leaf axils, furrows of
young stems or the midribs of leaves. Eclo-
sion occurred in 5 to 10 days and larvae
moved to young growing tips to feed be-
neath a loose silken webbing. Five larval
stadia were observed by the finding of cast
head capsules. The first four stadia were of
about 4 days duration. The final stadium
was 9 days including a prepupal period of
3 days. Pupation occurred in leaf litter sur-
rounding the plant and adult eclosion oc-
curred after 10 to 14 days. The life cycle
was completed in about 6 weeks.
459
Lorita baccharivora.— Eggs are round and
slightly flattened, whitish in color and trans-
lucent. They were oviposited along the mid-
rib on the upper surface of fully expanded
leaves where they were relatively easily dis-
cernible. Eclosion occurred in 10 to 20 days.
Neonate larvae were active and fed on near-
by leaves. After 2 to 3 days larvae moved
to growing tips where they fused young
leaves together with silk to form a tube. The
larva lived within this tube but left it to feed
on adjacent leaves. The leaves and the grow-
ing point in the tube usually died so that
further growth of the stem was arrested. Pu-
pation occurred within the tube and the life
cycle was completed in 4 to 6 weeks.
Hosts, DISTRIBUTION AND PHENOLOGY
Aristotelia ivae is found throughout much
of the range of B. halimifolia and has been
collected from Maryland (Kraft and Denno
1982) to Texas where it has also been found
on B. neglecta Britton (Palmer 1987). Ex-
cept for one series, all specimens have been
collected from Baccharis sp. Busck (1904)
gave Iva frutescens L. as the host for the
type series he described but we believe that
the host was misidentified (cf. Palmer and
Diatloff 1987).
A. ivae isa multivoltine species. Although
more abundant in spring and early summer,
it was found throughout most of the year in
Florida. Occasionally damage to the plant
was extensive with the leaves of the plant
having a characteristic skeletonised appear-
ance. Defoliation usually proved to be only
a temporary setback for the plant which i1n-
variably recovered. High rates (>50%) of
parasitism were observed and a species of
Apanteles sp. (Braconidae) was collected
from larvae.
Lorita baccharivora has been collected in
Florida and Texas (Pogue 1988). In Florida
we collected it from Gainesville to the Flor-
ida Keys and found it almost throughout
the year. While it is an abundant, wide-
spread species in Florida it is only rarely
encountered in Texas. Damage to the plant
460
Table 1. Plant species against which 4. ivae and L.
baccharivora were tested to obtain permission for their
introduction into Australia.
Apiaceae: Daucus carota L.; Pastinaca sativa L.
Anacardiaceae: Mangifera indica L.
Asteraceae: Baccharis halimifolia L.; Carthamus tinc-
torius L.; Chrysanthemum sp.; Dahlia sp.; Helian-
thus annuus L.; Lactuca sativa L.
Brassicaceae: Brassica oleraceae (L.) Alef.; Brassica
rapa L.
Bromeliaceae: Ananas comosus (L.) Merr.
Caricaceae: Carica papaya L.
Chenopodiaceae: Beta vulgaris L.
Convolvulaceae: Ipomoea hatatas (L.) Lam.
Cucurbitaceae: Cucumis melo L.; Cucumis sativus L.;
Curcubita maxima Duch.
Fabaceae: Arachis hypogaea L.; Centrosema pubescens
Benth.; Desmodium canum (Gmel.); Glycine wightii
(R. Grah. ex Wight & Arn.) Verde.; G/ycine max L.
Merr.; Medicago sativa L.; Phaseolus atropurpureus
DC.; Phaseolus vulgaris L.; Pisum sativum L.; Stizo-
lobium sp.; Stylosanthes gracilis, Trifolium repens
L.; Vigna catjang V.
Linaceae: Linum usitatissimum L.
Malvaceae: Gossypium hirsutum L.
Mimosaceae: Leucaena leucocephala (Lam.) de Wit.
Musaceae: Musa sapientum M.
Passifloraceae: Passiflora edulis Sims
Pinaceae: Pinus radiata D. Don.; Pinus taeda L.
Poaceae: Avena sativa L.; Digitaria decumbens Stent.;
Panicum maximum Jacq.; Paspalum dilatatum Poir.:
Pennisetum clandestinum Chiov.; Saccharum offi-
cinarum L.; Sorghum vulgare L.; Triticum aestivum
L.; Zea mays L.
Proteaceae: Macadamia integrifolia Maid & Betche
Rosaceae: Fragaria vesca L.; Malus sylvestris Mill.;
Prunus domestica L.; Prunus persica (L.) Batch.; Py-
rus communis L.; Rosa sp.
Rutaceae: Citrus limon (L.) Burm. F.; Citrus paradisi
Macfady.; Citrus reticulata Blanco; Citrus sinsensis
(L.)
Sapindaceae: Litchi chinensis Sonn.
Solanaceae: Capsicum annuum L.; Lycopersicum es-
culentum Miller; Nicotiana tabacum L.; Solanum
tuberosum L.
Vitaceae: Vitis vinifera L.
Zingiberaceae: Zingiber officinale Roscoe.
was significant when populations were high,
especially on plants under 2 m high. High
rates (>50%) of parasitism were observed
and Apanteles sp. (Braconidae) and Macro-
centrus sp. (Braconidae) emerged from lar-
vae. The only known host plant for this
species is B. halimifolia.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Host SPECIFICITY
The host specificity testing was conducted
at the Archbold Experimental Station at
Lake Placid, Florida in 1968. The same ex-
perimental design was used for both species
although testing was not done concurrently.
The ovipositional preference of the moths
and the food preference of larvae were tested
against a list of plants suggested by the Com-
monwealth Department of Health (Table 1).
Ovipositional preference was determined
by placing cuttings of young actively grow-
ing foliage from all of the test plants (Table
1) ina 40 x 36 x 30 cm glass-sided cage.
The cuttings were held in glass vials with
water. Twenty unsexed moths were placed
in the cage with a honey-water mixture.
When the B. halimifolia was obviously in-
fested, all the cuttings were carefully ob-
served and all eggs counted. Any infested
cuttings were kept for further observations
on the hatching and survival of larvae. The
experiments were replicated twice.
Food preference was determined by plac-
ing five field collected, partly grown larvae
on each of the 65 test plants, held in plastic
pots. The plants were carefully observed
daily for 10 days and any larvae noted. These
experiments were also replicated twice.
Aristotelia ivae.—Heavy oviposition oc-
curred on B. halimifolia within 3-5 days
and the resulting larvae developed rapidly.
Two eggs were found on one cutting of Lew-
caena leucocephala (Lam.) de Wit. but eggs
were not found on any other plant. These
two eggs hatched but the neonate larvae did
not feed and had disappeared by the next
day. When the field collected larvae were
placed on test plants they left all but the B.
halimifolia plants within 2 days. Some wan-
dered away and died while others moved to
the B. halimifolia plants and commenced
feeding. No feeding was attempted on any
plant other than B. halimifolia.
Lorita baccharivora.— Heavy oviposition
occurred on B. halimifolia cuttings within
3 to 7 days. Resulting larvae fed on the ex-
panded leaves for 2-3 days before moving
to the growing tips to form silken tubes.
VOLUME 90, NUMBER 4
Two eggs were also found on one cutting of
Prunus persica (L.) Batch but eggs were not
found on any other plant. These two eggs
hatched but the resulting larvae did not at-
tempt to feed. When field collected larvae
were placed on test plants they left all plants
other than B. halimifolia within 3 days.
Some wandered away and were lost but most
moved onto the B. halimifolia plants where
they occupied almost every available ter-
minal. Neither feeding nor fusing of leaves
to form a tube was attempted on any plant
other than B. halimifolia.
In both cases, the tests indicated that the
insects were sufficiently specific to release
in Australia. Both insects exhibited a strong
host finding mechanism in that virtually all
their eggs were laid on the Baccharis cut-
tings. This evidence is particularly signifi-
cant as, under cage conditions, many species
of moths oviposit on plants and objects that
would not be utilised under natural condi-
tions. The subsequent feeding tests con-
firmed the narrow host range indicated by
the oviposition tests. Larvae could survive
on only B. halimifolia of the 65 plant species
tested.
RELEASE IN AUSTRALIA
Permission was obtained to import the
insects into Australia. On arrival, both
species were required to be bred through
one generation in quarantine facilities to en-
sure that all parasites were eliminated. They
were then mass reared in non-quarantine
facilities to produce sufficient numbers for
field release.
Aristotelia ivae was introduced in south-
eastern Queensland in 1969 when approx-
imately 25,000 moths were released at five
locations. It readily established in this area:
at one site it spread out over 40 sq. k within
2 years of release. By 1974 it was found
throughout the range of B. halimifolia in
Australia. Although there have been local-
ized areas where high populations have
caused significant damage on smaller plants,
generally it has not occurred in sufficient
densities to control the weed.
461
Lorita baccharivora was imported into
Australia in 1969 but was not then suc-
cessfully reared. Following further impor-
tations in 1984, it was successfully mass
reared at the Alan Fletcher Research Station
and released at various locations in south-
eastern Queensland in 1986. Moths have
not yet been recovered from the field so
establishment has not been confirmed. Fur-
ther releases were made until the end of
1987.
ACKNOWLEDGMENTS
We thank R. W. Hodges of the Systematic
Entomology Laboratory, Agricultural Re-
search Service, USDA who identified A. ivae
and M. G. Pogue of the Department of
Entomology, National Museum of Natural
Histoyr, Smithsonian Institution who re-
viewed the genus Lorita and described L.
baccharivora.
LITERATURE CITED
Arnett, R. H., Jr. 1985. American Insects: A Hand-
book of the Insects of America North of Mexico.
Van Nostrand Reinhold Co., New York. 850 pp.
Busck, A. 1904. New species of moths of the super-
family Tineina from Florida. Proc. U.S. Nat. Mus.
233225.
Kraft, S. K. and R. F. Denno. 1982. Feeding re-
sponses of adapted and non-adapted insects to the
defensive properties of Baccharis halimifolia L.
(Compositae). Oecol. 52: 156-163.
Palmer, W. A. 1987. The phytophagous insect fauna
associated with Baccharis halimifola L. and B. ne-
glecta Britton in Texas, Louisiana and Northern
Mexico. Proc. Entomol. Soc. Wash. 89: 185-199.
Palmer, W. A.and F. D. Bennett. 1988. The phytoph-
agous insect fauna associated with Baccharis hal-
imifolia L. along the eastern seaboard of the United
States. Proc. Entomol. Soc. Wash. 90(2): 216.
Palmer, W. A. and G. Diatloff. 1987. Host specificity
and biology of Bucculatrix ivella Busck, a potential
biological control agent for Baccharis halimifolia
L. in Australia. J. Lepid. Soc. 41: 23-28.
Pogue, M. G. 1988. A revision of the genus Lorita
Busck (Lepidoptera: Tortricidae: Tortricinae:
Cochlyini), with a description of a new species.
Proc. Entomol. Soc. Wash. 90(4): 440.
Tilden, J. W. 1951. Microlepidoptera associated with
Baccharis pilularis. 11. Tortricidae, Phaloniidae,
Gelechiidae. Wasmann Journal of Biology 9: 239-
254.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 462-470
NESTING, MATING AND FORAGING HABITS OF MELISSODES
(MELISSODES) TEPIDA TEPIDA CRESSON IN IDAHO
(HYMENOPTERA: ANTHOPHORIDAE)
D. C. TRIPLETT AND A. R. GITTINS
Division of Entomology, University of Idaho, Moscow, Idaho 83843.
Abstract.— Field studies are reported on nest construction, mating, and female foraging
behavior of Melissodes tepida tepida Cresson, at two nesting sites in southwestern Idaho.
Females form loose nesting aggregations but construct and provision individual burrows
in the ground. Adult males exhibit protandry and their flight activity approaches territorial
behavior patterns. Melissodes tepida tepida is a polylectic subspecies visiting a number
of different host plants for pollen as well as nectar.
Key Words:
Studies of wild bee species, especially
those which evidence potential for popu-
lation enhancement and management, form
an integral part of a continuing research pro-
gram for pollination of seed crops in Idaho.
Melissodes tepida tepida Cresson, an aggre-
gate nester, frequently found in seed grow-
ing areas of southwestern Idaho, exhibits
characteristics which initially suggest po-
tential for management as a pollinator in
forage and hybrid vegetable seed produc-
tion. This paper records the first portion of
our detailed studies on the biology of this
subspecies.
The species is found nesting gregariously
in well-defined ‘“‘bee beds.” Like most oth-
ers of more than 90 species of Melissodes,
M. t. tepida biology has been unknown.
LaBerge (1956a, b) reclassified the /. tepida
complex into three subspecies, M. ¢. tepida
Cresson, M. t. timberlakei (Cockerell) and
M. t. yumensis LaBerge, and mapped the
geographic ranges of each. Recent papers
recording details on nesting habits of other
Melissodes species, and serving as back-
ground information for our studies, include
Hurd and Linsley (1959), Evans and Linsley
flower bee, native bee biology, protandry, groundnesting.
(1960), LaBerge (1961,1963), Butler et al.
(1962), Thorp and Chemsak (1964), Clem-
ent (1973), Batra and Schuster (1977),
Buchmann and Jones (1980), Parker et al.
(1981), Tepedino and Parker (1982), and
Bouseman (1987). Refer to Thorp and
Chemsak (1964), Clement (1973), and
Buchmann and Jones (1980) for list of ear-
lier recorded publications on Melissodes spp.
biology.
Linsley (1946) has presented the only pre-
viously recorded information on the biology
of the species.
Ours and other collecting records indicate
M. t. tepida is a polylectic group with pref-
erence for visiting flowers of the families
Euphorbiaceae, Lamiaceae, Fabaceae, and
the composite genus Gutierrezia.
NESTING SITES
Two nesting sites near Parma, Canyon
County, Idaho, were selected for field stud-
ies and as a source of laboratory material.
Site | was a 2 m x 30 m strip along the
southern edge of a farm road. Distichilis
stricta (Torr.) Rydg., a salt grass, provided
VOLUME 90, NUMBER 4
the only vegetative cover overlaying the en-
tire site in varying density. The soil is a
Moulton sandy loam saline with the top 5—
6 cm a mixture of litter and sandy loam,
and with moist sandy loam densely packed
beneath. D. stricta roots and rhizomes are
found matted to a depth of over 20 cm
throughout the site. Site 2, nearly 4 km from
site 1, was approximately 90 m square with
an evenly, densely distributed cover of D.
stricta. The site was otherwise markedly
similar to site | in soil characteristics.
METHODS AND MATERIALS
Twelve equal-sized plots delimited by
plastic stakes were established at each site.
Wind velocity was recorded continuously
during field observation periods using ane-
mometers set at elevations of 30 cm, | m
and 2 m. Humidity and temperature data
were also recorded. At site 1, soil temper-
atures were monitored at depths of 5, 10,
15, 20, 25 and 30 cm beneath the surface
using copper-constant thermocouples re-
cording directly through a Leeds and Nor-
thrup potentiometer.
Early field observations centered upon
determining emergence periods of . ¢. tep-
ida and associated insects. Metal screen
mesh emergence cages (1.5 x 2 x 1 m)
distributed randomly on the sites were used
to trap the emerging insects. Daily checks
were made, data recorded, and trapped
specimens were then released.
To facilitate tracking individual bee ac-
tivity, specimens were captured with nets,
dusted with a powdered, daylight, fluores-
cent dust (trade name Day-Glo) and re-
leased. Three distinctive colors were used:
neon-red, fire-orange, and arc-yellow. After
release, individual bees were easily ob-
served and the color identified at distances
up to 10 m. This pigmented powder was
also used to follow construction of burrows.
Selected burrows were tagged by placing
numbered plastic stakes near each entrance.
Each numbered burrow was then dusted
with Day-Glo. The burrows were marked
463
from three to six times at intervals during
the day using an aspirator bulb and directing
the powder into the burrow through a ta-
pered glass nozzle. Movements of earth in
the burrow and accompanying brushing ac-
tion by females distributed the powder
throughout the burrow. Their excavating
and burrow sealing activities incorporated
the dust into the plugging material of the
laterals and the cells walls. In addition, fe-
males returning from foraging were dusted
before entering a marked burrow. Using this
burrow-marking technique, 51 nests were
dusted and studied over the two-year study
period.
Each fall all marked burrows were exca-
vated using a standarized technique which
exposed the details of burrow construction.
The technique consisted of digging a trench,
with a 45 cm deep vertical face 30 cm from
the burrow entrance. Using a small knife,
brush, and air tubing, each burrow was ex-
posed step-by-step from the entrance to the
distal cells. A model of each burrow was
then constructed. Exposure of the burrow
construction was always complicated by the
maze of D. stricta roots.
All cells removed from excavated bur-
rows were placed in plastic cups, numbered,
stored in a styrofoam cooler, and taken to
the laboratory for detailed examination.
Over two hundred cells were removed and
used in the laboratory studies to supplement
observations and experiments conducted at
the nesting sites. These studies concentrated
primarily upon anatomical evaluations of
the life stages, determination of develop-
mental stadia reared under controlled tem-
peratures, and observations on larval feed-
ing. Results of the laboratory studies are
being presented in a subsequent paper.
Female activity at the nesting site was
charted continuously, including timing the
periods of foraging and nest construction.
Cylindrical screen cages (30 cm diameter)
were placed over individual burrows. As a
marked female emerged from the burrow or
approached the burrow subsequent to for-
aging, the cage was removed, allowing ac-
464
tivity of that individual to continue. Each
phase of insect activity was recorded. Stop
watches were used to time the activity in-
tervals.
Pollen was removed from a sample of cells
and from pollen loads on females at irreg-
ular intervals as the bees returned from for-
aging. Pollen slides were prepared for mi-
croscopic examination to determine the
pollen sources employing the MacCallum-
Goodpasture method which uses a gentian
dye for staining and consequently highlight-
ing individual pollen grains.
BIOLOGY
Adult emergence. — Following ecdysis
from the pupal case, teneral adults com-
monly remained in the cells for several days
before burrowing to the surface. During
emergence the bees chewed through the co-
coon and fecal cap, and burrowed vertically
through the soil leaving the old cell filled
with the shredded cocoon, fecal material,
and soil. Emerging bees remained in the
mouth of the exit burrow approximately 24
hours before beginning flight activity. MZ. t.
tepida exhibited protandry with males
emerging and establishing territorial flight
patterns over the nesting site approximately
seven days prior to emergence of the first
female bees. However, some overlap in the
range of emergence times for male and fe-
male bees did occur.
At the first nesting site, emergence began
at the west end of the site and extended to
the east end over approximately a two-week
period. Increase in plant cover on the site
west to east influenced soil temperature and
undoubtedly accounted for differences in
emergence times. Soil temperature records
taken at varying soil depths in the site sup-
ported this conclusion.
Male bees began actively establishing
flight patterns over the nesting sites in early
July. Excavation of nest samples at the time
of first male emergence exposed both de-
veloping pupae and female adults ready to
emerge.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
The normal life span of adult male bees
generally varied from 12 to 15 days (average
14 days). Two weeks after initial emergence,
a decline could be noted in the number of
male bees present at the sites, and by the
end of July few males were observed. The
normal life span of the female ranged from
18 to 23 days with an average of 20 days.
With the range in the emergence times, nest-
ing activity continued through early August.
Male flight activity.—The flight activity
of male M. t¢. tepida following emergence
exhibited a basic territorial behavior pat-
tern. The individual bee defended an area
14 cm to 30 cm in diameter from intrusion
by other male bees. A male would chase an
intruding bee for a distance of up to 3 m
from the defended area, and then return.
Male bees, captured and removed from
their territory and released elsewhere on the
site, would return to their original area of
activity usually within 30 minutes.
Mating.—Male aggressiveness was dis-
played in its mating behavior. As virgin fe-
males emerged from their burrows, they
were literally pounced upon by the males
either before they took flight or as they be-
gan to fly. Virgin females which attained
flight were knocked back down to the
ground. In many cases a newly emerged fe-
male accepted the male’s attempt to copu-
late without rejection. All mating took place
on the ground. The male mounted the fe-
male dorsally with his prothoracic legs
grasping the female around the mesothorax
and his mesothoracic and metathoracic legs
hooked around the female’s abdomen. Dur-
ing copulation the male displayed two dif-
ferent pulsating actions of his abdomen.
Commonly, first pulsations were very rapid
and short, lasting from five to ten seconds
in duration. Following this, pulsations be-
came slower and stronger, lasting approxi-
mately ten seconds. During these copula-
tory activities the antennae of the male came
in contact with the female’s antennae, but
no definite stroking patterns were noted. The
total period in copula ranged from 40 to 65
VOLUME 90, NUMBER 4
seconds with an average of 45 seconds. Fol-
lowing copulation the female departed and
began nesting activity.
Mated females remained attractive to all
males for a few days following copulation.
This suggests a probable sex attractant se-
creted by the female. During this period of
continued attraction, as females ap-
proached their burrows carrying pollen,
males would dart from an established flight
pattern, knock females to the ground and
attempt copulation. These copulatory at-
tempts were repulsed by the females who
freed themselves within seconds and re-
sumed nesting activity. Mated females be-
came less attractive to males with time and,
after approximately one week following
copulation, they were ignored by male bees
as they flew about the nesting site.
On two occasions during our extensive
nesting site observations, males were seen
attempting copulation with other males. In
each case contact between the two males
was terminated within a few seconds.
Sleeping behavior.—Males slept at the
nesting site, finding shelter under debris or
at the entrance of a female nesting burrow.
In a burrow, males slept with their heads
positioned outward. Females always slept
in the burrow. Excavation of burrows after
10:30 pm exposed inactive females in the
lower distal portion of the burrow.
Daily activity.—Although both soil tem-
peratures and air temperatures were record-
ed at nesting site 1, it was difficult to deter-
mine specific temperature thresholds for 7.
t. tepida activity. Generally, air tempera-
tures reached 21°C before full bee flight ac-
tivity was observed. At air temperatures
above 36°C activity was greatly reduced, al-
though a few individuals remained active
at temperatures of 38°C. Bee flight activity
was much reduced when wind velocities ex-
ceeded 24 km per hour. Normally male bee
activity began between 9:00 am and 9:30
am and continued throughout most of the
day. It gradually decreased following 1:00
pm and ceased by 5:30 pm. Female bee for-
465
aging activity began between 10:00 am and
10:30 am. Generally, foraging activity
reached a maximum by 11:00 am, de-
creased sharply between 1:00 pm and 2:00
pm, and continued to decline during the
afternoon as females spent more time in the
burrows. Foraging activity virtually ceased
by 6:30 pm. During the course of these stud-
ies two non-foraging females were observed
returning to the nesting site after 6:30 pm.
Nest construction.—Female ™. ¢. tepida
constructed and provisioned individual
burrows. Generally, a female constructed
only one burrow during the season, but some
females were observed excavating a second
or even a third burrow in the event of de-
struction or obstruction of the burrow en-
trance.
This subspecies showed a tendency to
construct entrances at the edge of rocks or
other ground debris, although some bees
burrowed in open areas covered only by the
salt grass vegetation. In areas supporting
large numbers of females, entrances were
frequently 2.5 cm to 12 cm apart, while in
less active areas distances between holes fre-
quently varied from 0.3 m to 2 m. When
numerous rocks were placed on the ground
in a central area of bee activity, small ag-
gregations of female bees, sometimes as
many as seven, began constructing burrows
at the edge of the rocks.
While initiation of nest construction at
the sites was noted at many differing hours
throughout the day, the majority of females
began nest construction during the morning
hours subsequent to 10:30 am.
Once a female had chosen an area for nest
construction, she landed and began scratch-
ing vigorously with the prothoracic legs,
kicking the soil behind the abdomen with
these and the mesothoracic legs. She quickly
penetrated the soil surface and removed the
dirt from the main shaft as she backed out
of the burrow. Tumulus formed by the re-
moval of the soil from the main shaft was
dissipated in a few days with weathering and
bee activity about the entrance. Below the
466 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
Section of a soil profile from site | exposing the main shaft and lateral of a M. ¢. tepida nest. Note
cell beneath lateral and partial outline of a second lateral and cell extending from first lateral.
first 5 cm of soil the ground became in-
creasingly more compacted. Here the fe-
male was seen to use a twisting action of the
head, loosening the soil particles with her
mandibles. Following completion of the first
cell, the female began excavation of a sec-
ond lateral, at an angle out from the main
shaft. Dirt excavated from this lateral was
repacked into the first lateral completely
plugging it to the main shaft.
Nest description (Fig. 1; Fig. 2A, B; Fig.
3A-C).—The open circular entrance (di-
ameter 7 mm) had no turret. The burrow
was formed at angles approaching 90° to soil
surface with the main shaft (diameter 6 mm)
extending to a depth of 7 to 13 cm. Laterals
branched out from the main shaft at differ-
ent levels at angles of 15° to 90° from the
main burrow. The distal end of each lateral
was curved and widened to 7 mm and ended
in a single, vertical cell. Depths of cells
ranged between 10 and 20 cm beneath the
soil surface at both nesting sites. The main
shaft and the laterals were smooth-walled
but lacked a lining as found in the cell. In
constructing a cell, the female first “‘roughed
out” the cell to a diameter of 1.4 cm, then
repacked the soil leaving a hard, smooth
crust lining the cell wall. The finished cy-
lindrical cell (diameter 7 mm, length 15 mm)
had a rounded base and tapered near the
top to about 6 mm in diameter. The spe-
cially constructed cell was found to be high-
ly resistant to water penetration.
Provisioning.—The female transported
unmoistened pollen on the scopal hairs of
the metathoracic legs. She approached her
nest, landed near the opening, and entered
the burrow immediately. In the burrow a
female must reverse position either at the
widened distal end of the lateral or within
the nest cell in order to deposit the pollen
load at the base of the cell. The female re-
moved the pollen load by rubbing the meta-
VOLUME 90, NUMBER 4
467
Fig..2;
A. Drawing of enlarged nest cell of MV. ¢. tepida to illustrate (a) plugged lateral, (b) spiraled cell cap,
(c) repacked and mixed cell wall, (d) egg, (e) pollen mass. Bar equals 0.5 cm. B. Illustration of generalized nest
of M. t. tepida. Bar equals 4 cm.
thoracic legs against the abdomen and uti-
lizing the lever action of the tibial spurs of
the opposing leg. Later, after the female had
completed stocking that cell, she mixed the
pollen mass with nectar and packed this lar-
val food into a semi-solid mixture, leaving
the upper surface slightly concave. These
pollen masses occupied about 30% of cell
volume in the lower portions of cells ex-
amined and averaged 4.5 grams in weight.
During embryogenesis the pollen-nectar
mixture fermented, forming a semi-liquid
mass which increased in volume.
Pollen samples extracted from cells or re-
468 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
b 4 aL Te
c
Fig. 3. Schematic sketches of three nests of M. ¢. tepida reconstructed after excavations during this study.
A.B.C. Lateral aspects of three nests. a.b.c. Dorsal aspects of same nests. Bar equals 4 cm; —O— represents
burrow entrances.
VOLUME 90, NUMBER 4
Sequence and duration
of pollen collecting trips
469
Time intervals at the nest
between pollen trips
2nrsic
1 hr
kk
I ee
*
10
a
{] (28 Se 44 5) "6 7 4
2 3 4 5 6 7
Interval Sequences
* Female returned without
pollen.
** Female remained in
burrow.
Fig. 4. Histogram illustrating a typical time sequence of foraging and at nest activities by an individual
female M. ft. tepida in stocking a single cell.
moved from foraging females were collected
from several species of Fabaceae (Medicago
sativa L., Melilotus officinalis (L.) Pall., Tri-
folium sp.), an Asteraceae species Gutier-
rezia sarothrea (Pursh) Britt. & Rusby., and
a Chenopodiaceae Atriplex sp., verifying that
female M. t. tepida are indeed polylectic.
Our records established that 5—6 foraging
trips by a female were required to provision
a nest cell. The time spent collecting pollen
in the field varied from 13 minutes to 35
minutes per trip with an average time of 15
minutes. The average period of time re-
quired for depositing a pollen load was ap-
proximately six minutes. As the female
neared completion of cell provisioning, she
often remained in the burrow for longer pe-
riods of time, mixing and packing the pol-
len-nectar mass or resting. A graphical pre-
sentation typical of the time sequence of
foraging and pollen deposition for a single
cell by an individual female bee is illus-
trated in Fig. 4. The authors did not observe
any variation in female foraging times as
correlated to plant species visited.
SUMMARY AND DISCUSSION
The extent of variability in the behavior
patterns among Melissodes species is note-
worthy. Nesting of M. ¢. tepida appears re-
stricted to alkali soils. This differs from re-
ports for other species in the genus, but
agrees with other species in that all nest in
soils with a sandy surface and moist com-
pacted soil below. Others have found species
generally nesting in bare soil areas while .
t. tepida was found on sites reasonably well-
covered by Distichilis stricta, an alkaline soil
indicator plant. Nesting is aggregated dis-
playing a tendency for concentration of bur-
rows under ground debris or along edges of
rocks or soil clods. Females were not seen
470
to share burrows or burrow entrances and
did not plug those entrances until the com-
pletion of the foraging period. Unlike some
species reported to construct single cell bur-
rows, M. t. tepida develops multicelled nests
and coats each cell with a waterproof lining
which, we suspect, from Batra and Hefetz
studies (1979), is of acetate material from
Dufour’s gland. They stock these cells about
one-third full with pollen mixed with some
nectar and compacted into a cylinder with
a concave surface.
Males begin to emerge from southwestern
Idaho nesting sites in early July and estab-
lish flight patterns ofa territorial-like nature
a week prior to female emergence. Females
begin emerging in mid-July and nesting ac-
tivity extends to about mid-August.
Our studies of the pollen sources showed
M. t. tepida to be polylectic, visiting a va-
riety of host plants for pollen including Atri-
plex sp., Gutierrezia sarothrea, Medicago
sativa, Melilotus officinalis, and Trifolium
spp. Studies of pollen samples revealed a
preference for Atriplex sp. Because of the
relatively small populations 1n this area and
the fact that the legumes comprise only 5%
to 10% of the pollen collected for cell pro-
visioning, the economic value of M7. t. tepida
as an important pollinator remains ques-
tionable. Nevertheless the fact these insects
form nesting aggregations in a specific nest-
ing medium and are relatively polylectic of-
fers potential for manipulation in hybrid
seed production, especially under large cage
management conditions. Further study and
experimentation with this potential is de-
sirable.
ACKNOWLEDGMENTS
The authors express their appreciation to
G. E. Bohart, Logan, Utah for his species
determinations, and to him, M. A. Brusven
and G. W. Bishop, University of Idaho, for
their advice on the study. We are also in-
debted to A. L. Steinhauer and F. G. Wood,
University of Maryland, and to J. B. John-
son and M. Rice, University of Idaho, for
their valuable assistance with this manu-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
script. Contribution No. 88721 from the
University of Idaho, Agricultural Experi-
ment Station.
LITERATURE CITED
Batra, S. W. T. and A. Hefetz. 1979. Chemistry of
the Cephalic and Dufour’s gland secretions of Mel-
issodes bees. Ann. Entomol. Soc. Am. 72(4): 514—
515:
Batra, S. W. T. and J. C. Schuster. 1977. Nests of
Centris, Melissodes and Colletes in Guatemala
(Hymenoptera: Apoidea). Biotropica 9(2): 135-
138.
Bouseman,J.K. 1987. Collection of Melissodes (Apo-
melissodes) apicata in Illinois (Hymenoptera:
Apoidea). J. Kans. Entomol. Soc. 60(2): 335-336.
Buchmann, S. L.andC. E. Jones. 1980. Observations
on the nesting biology of Melissodes persimilis
(Hymenoptera: Anthophoridae). Pan-Pac. Ento-
mol. 56(3): 200-206.
Butler, G. D., Jr., F. E. Todd, S. E. McGregor, and F.
G. Werner. 1962. Melissodes bees in Arizona
cotton fields. Arizona Univ. Agr. Exp. Sta. Tech.
Bull. 139: 1-11.
Clement, Stephen L. 1973. The nesting biology of
Melissodes (Eumelissodes) rustica Say, with a de-
scription of the larva (Hymenoptera: Anthro-
phoridae). J. Kans. Entomol. Soc. 46(4): 516-525.
Evans, H. E. and E. G. Linsley. 1960. Notes on a
sleeping aggregation of solitary bees and wasps.
Bull. So. Calif. Acad. Sci. 59: 30-37.
Hurd, P. D. and E. G. Linsley. 1959. Observations
on the nest site behavior of Melissodes composita
Tucker and its parasites, with notes on the com-
munal uses of nest entrances (Hymenoptera: Apoi-
dea). Entomol. News 70: 141-146.
LaBerge, W. E. 1956a. A revision of the bees of the
genus Melissodes in the North and Central Amer-
ica Part I. Univ. Kans. Sci. Bull. 37(2): 911-1194.
1956b. Ibid. Part IT. 38(1): 533-578.1957.
1961. Ibid. Part II. 42(5): 283-663.
. 1963. New species and records of little-known
species of Melissodes from North America. Univ.
Nebr. State Mus. Bull. 4: 227-242.
Linsley, E. G. 1946. Insect pollinators of alfalfa in
California. J. Econ. Entomol. 39: 18-29.
Parker, F. D., V. J. Tepedino, and G. E. Bohart. 1981.
Notes of the biology of a common sunflower bee
Melissodes agilis. J. New York Entomol. Soc. 89(1):
43-52.
Tepedino, V. J. and F. D. Parker. 1982. Interspecific
differences in the relative importance of pollen and
nectar to bee species foraging on sunflowers. En-
viron. Entomol. 1 1(1); 246-250.
Thorp, R. W. and J. A. Chemsak. 1964. Biological
observations on Melissodes (Eumelissodes) palli-
disignata. Pan-Pac. Entomol. 40: 75-83.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 471-473
TAXONOMIC NOTES ON RHYSSALINI AND RHYSIPOLINI
(HYMENOPTERA: BRACONIDAE) WITH FIRST NEARCTIC
RECORDS OF THREE GENERA
JAMES B. WHITFIELD
Department of Entomology, The Ohio State University, Columbus, Ohio 43210.
Abstract.—The braconid genera Dolopsidea Hincks, Neurocrassus Snoflak and Rhyssalus
Haliday are reported from the New World Nearctic Region for the first time. Notes are
provided to facilitate identification of these genera with reference to the key to Nearctic
braconid genera of Marsh et al. (1987), and with respect to other genera in their respective
tribes. Figures are provided but formal species descriptions await revisions of the re-
spective genera.
Key Words:
In recent sorting of undetermined roga-
dine and hormiine Braconidae from my own
collection, I have come across several closely
related genera that have not previously been
reported from North America, and were not
included in the recent manual of Nearctic
braconid genera (Marsh et al. 1987). A re-
classification of the subfamilies containing
these genera 1s now in preparation by the
author; the discovery of these genera in the
North American fauna is being published
now to facilitate the discovery of additional
material of these groups, and to update the
generic keys.
Dolopsidea Hincks, 1944
One female specimen of this genus, very
similar to the Palearctic D. indagator (Hal-
iday), was collected by malaise trap in the
Hocking Hills, Hocking Co., Ohio, by the
author, 3-9 June 1987.
The genus has often been confused, even
in Britain and Europe, with Rhyssalus Hal-
iday (see, e.g. Stelfox 1951) and Oncophanes
Foerster. It will run in the key of Marsh et
al. (1987) to Oncophanes (couplet 181) with
some difficulty but differs from Oncophanes
distribution, Dolopsidea, Neurocrassus, Rhyssalus
in having the medial lobe of the mesoscu-
tum sharply and strongly raised above the
lateral lobes, and in the metasomal tergites
2 and 3 being enlarged and highly polished,
and not sharply delineated from the epi-
pleura by creases. In addition, the ovipos-
itor and sheaths are usually at least as long
as the metasoma. In the mesoscutal struc-
ture and metasomal sculpturing Do/opsidea
also differs from Rhyssalus Haliday (see be-
low), which, in addition to being much
smaller, has a relatively unraised medial
mesoscutal lobe and normal-sized and
weakly sclerotized second and third meta-
somal tergites. All three genera have a dis-
tinctive propodeal carination pattern (Fig.
4), and have the spiracles of metasomal ter-
ga 2 and 3 positioned lateroventrally, below
the lateral edges of the dorsal faces of the
terga. The fore wing venation of the Ohio
specimen is shown in Fig. 1.
Neurocrassus Snoflak, 1945
A single male specimen was recovered
from the same malaise trap as the above
genus, 5 September 1987, at the same site.
It appears to be a rare genus even 1n Europe.
472 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-5. 1-3: Fore wings of 1, Dolopsidea sp., 2, Hocking Co., Ohio; 2, Neurocrassus sp., 3, same locality;
3, Rhyssalus sp., 2, Old Chelsea, Quebec. 4-5: Propodea of 4, Dolopsidea sp. (same specimen as above); 5,
Neurocrassus sp. (also same as above). Scale line = 0.5 mm (1-3); 0.2 mm (4-5).
VOLUME 90, NUMBER 4
The obvious distinguishing feature of the
genus is its peculiar, partially swollen wing
venation (Fig. 2 and see also Snoflak 1945
and Tobias 1986). Otherwise the genus fits
within the Rhysipolini sensu Belokobyl’ski
(1984), near Cantharoctonus Viereck, No-
serus Foerster and Pseudavga Tobias, al-
though it possesses a mosaic of characters
from these genera (c.f. Whitfield & van Ach-
terberg 1987). It will run in the key of Marsh
et al. (1987) with some difficulty, due to
equivocal hind wing characters, to couplet
180, containing Cantharoctonus and Rhy-
sipolis Foerster. It agrees best with Can-
tharoctonus but has a less broad transverse
groove at the base of the propodeum, as well
as the conspicuous wing features. The pro-
podeal carination 1s shown in Fig. 5.
Rhyssalus Haliday, 1833
From earlier collections I discovered a
female from Old Chelsea, Quebec, collected
on 18 July 1987. I have seen several other
specimens in the Canadian National Col-
lection, all from eastern Canada (Ontario
and Quebec), and probably conspecific.
This genus will run to the same point in
the key of Marsh et al. (1987) as Dolopsidea
(see above). I have given characters that will
separate these two genera from each other
and the other genera in the key. The fore
wing venation is shown in Fig. 3. Rhyssalus
also shares many characters with Pseudo-
bathystomus Belokobyl’skii (1987), which
so far has not been discovered in the Nearc-
tic fauna. Rhyssalus is distinctive in having
metasomal terga 2 and 3 mostly weakly
sclerotized and not enlarged relative to the
succeeding terga; some species also possess
clavate hind tibiae. This group of genera,
the Rhyssalini s.s., is much in need of re-
vision at the generic and specific levels.
ACKNOWLEDGMENTS
I appreciate the funding of the CanaColl
Foundation in supporting my studies of
473
these genera and related groups at the Ca-
nadian National Collection of Insects, Ot-
tawa. A fellowship awarded by the North
Atlantic Treaty Organization supported
earlier studies at the British Museum (Nat-
ural History) and the Rijksmuseum van Na-
tuurlijke Historie, Leiden, that clarified the
identities of these genera. I would also like
to thank Paul M. Marsh (U.S. National Mu-
seum, Washington) for loaning the Stelfox
collection of exothecine Braconidae to me,
which proved critical in interpreting some
earlier papers on these wasps. Norman F.
Johnson and Sydney A. Cameron provided
useful comments on a draft of this manu-
script.
LITERATURE CITED
Belokobyl’ski, S. A. 1984. [Division of the tribe Ex-
othecini s.1. (Hymenoptera: Braconidae) into two
with description of a new genus and subgenus.]
Zool. Zhurn. 63: 1019-1026. [In Russian.]
. 1987. Anew braconid genus of the supertribe
Exothecidil (Hymenoptera: Braconidae). Ento-
mol. Rev. 66: 116-120. [Ong. Russian version
1986.]
Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987.
An identification manual for the North American
genera of the family Braconidae (Hymenoptera).
Mem. Entomol. Soc. Wash. 13. 98 pp.
Snoflak, J. 1945. Neurocrassus gen. n. tesari sp. n.
Folia Entomol. 8: 25-27.
Stelfox, A. W. 1951. On the identity of Rhyssalus
indagator Haliday with Dolops hastifer and D. acu-
leator Marshall (Hym., Braconidae). Entomol.
Month. Mag. 87: 90-93.
Tobias, V. I. 1986. [Identification keys to the insects
of the European portion of the USSR. Vol. III, 27:
Hymenoptera. Parts 4, 5. Braconidae.] Akademia
Nauk USSR. Nauka Press. [In Russian.]
Whitfield, J. B. and C. van Achterberg. 1987. Clar-
ification of the taxonomic status of the genera Can-
tharoctonus Viereck, Noserus Foerster and Pseu-
davga Tobias (Hymenoptera: Braconidae). Syst.
Entomol. 12: 509-518.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 474-479
NOTES ON THE BIOLOGY AND IMMATURE STAGES OF
POECILOGRAPHA DECORA (LOEW)
(DIPTERA: SCIOMYZIDAE)
JEFFREY K. BARNES
Biological Survey, New York State Museum, The State Education Department, Albany,
New York 12230.
Abstract. —The puparium and third-instar cephalopharyngeal skeleton of the enigmatic
sciomyzid fly, Poecilographa decora (Loew), are described, and the distribution is mapped.
Notes on life history are presented, as well as ecological and taxonomic predictions based
upon morphology of the immature stages.
Key Words:
While working as a consultant entomol-
ogist with the Arctic Health Research Cen-
ter of the U.S. Public Health Service at An-
chorage, Alaska, during the summers of
1950-1952, the late Professor Clifford O.
Berg discovered that larvae of Sciomyzidae
prey upon mollusks. Following that discov-
ery, his own research and that of many stu-
dents and associates turned mainly to the
careful elucidation of life histories of the
sciomyzids of the world. About one third
of the approximately 600 described species
in this cosmopolitan family have been
reared, and the larvae of all are obligate
predators or parasitoids of various mol-
lusks. Most species feed on freshwater or
terrestrial, nonoperculate snails, but some
attack snail eggs, operculate snails, salt
marsh snails, slugs, or sphaeriid clams. Berg
and Knutson (1978) reviewed the biology
and systematics of the Sciomyzidae.
Nearly all students of Nearctic Sciomy-
zidae have attempted to rear the striking
and distinctive species, Poecilographa dec-
ora (Loew), the sole member of its genus.
However, no researcher has ever reared it
through the entire life cycle, and the liter-
ature on this species is largely restricted to
Diptera, Sciomyzidae, Poecilographa, immature stages
basic taxonomy and Johannsen’s (1935)
brief description of a puparium. My recent
discovery of three puparia in the field has
prompted the writing of this paper, which
I hope will stimulate renewed interest in
elucidating the life history of this mysteri-
Ous species.
BIOLOGY
Puparia were collected at Black Creek
Swamp, on Koontz Rd., Voorheesville, New
York (42°39'57’N, 73°58'05”W). They were
found among moist litter under a thin can-
opy of Ulmus rubra Muhlenberg and Frax-
inus pennsylvanica Marshall in an unflood-
ed area where adults had been collected
previously. The dominant low vegetation is
Aster simplex Willdenow and Onoclea sen-
sibilis Linnaeus. The locality is frequently
flooded during the spring and after heavy
rains, and it is surrounded by a creek and a
Typha and Sparganium marsh.
A puparium collected June 2, 1983, yield-
ed an adult female on June 15, and one
collected on June 16, 1983, yielded a female
on June 27. The third puparium, collected
June 30, 1983, yielded no adult and upon
close inspection was found to have a 0.7
VOLUME 90, NUMBER 4
mm diameter circular hole in the integu-
ment, probably an emergence hole ofa para-
sitoid. The single puparium that Johannsen
(1935) reported upon “... was found in a
bog in woods near Ithaca, New York, on
June 2” (p. 48), and it yielded an adult on
June 17. Johannsen’s puparium is not in the
Cornell University Insect Collection with
the adult it yielded, and it probably has been
lost. The others are deposited in the New
York State Museum.
On July 6, 1981, one male and three fe-
male P. decora adults were collected at Black
Creek Swamp and placed ina 5.0 x 8.5 cm
clear plastic vial fitted with a screen cap and
containing a layer of moist cotton, forest
litter, a resting stick, and an artificial diet
for the adults consisting of honey, brewer’s
yeast, and dehydrated milk. They were held
in an incubator at 20°C under a LD 16:8
lighting schedule. They mated and laid eggs
readily and frequently. The eggs, which are
creamy white, 0.83-0.91 mm long, and
striate, were placed in scattered, unorga-
nized groups, usually along the edge of a
piece of drying litter in the bottom of the
vial. On July 15, the eggs laid over the pre-
vious nine days were harvested from the
breeding vial, and it was found that none
had hatched. The eggs were placed on moist
cotton, and four of them hatched on July
29. These four had been kept especially
moist—so that the chorion actually ap-
peared wet: others had dried somewhat from
evaporation. The larvae were placed in a
dish of water, but no matter how carefully
they were manipulated, it was impossible
to make them float, even though micro-
scopic examination reveals that they pos-
sess short, interspiracular, hairlike process-
es or “float hairs.”’ Other researchers have
found that first-instar larvae of P. decora
float readily, with the posterior end at the
surface film (B. A. Foote, pers. comm.).
The eggs laid July 15-31 were submerged
in water for about 2 hours on July 31. None
had hatched up to that date, but by August
3, 15 had hatched; by August 6, 10 more;
475
by August 10, 36 more; and by August 19,
the remaining 16 had hatched. Very few
more eggs were laid, and all adults soon
died.
Attempts were made to rear first-instar
larvae on various gastropods, including Gy-
raulus sp. (Planorbidae), Lymnaea sp.
(Lymnaeidae), Oxy/oma sp. (Succineidae),
and an unidentified land snail (Discidae:
Discus sp.?), collected at Black Creek
Swamp. No feeding was observed, and all
larvae perished. Failure also resulted from
attempts to rear first-instar larvae on living
Biomphalaria glabrata (Say) and Helisoma
trivolvis (Say) (Planorbidae); living and
freshly killed Deroceras laeve (Miller) (Li-
macidae); living Haplotrema concavum
(Say) (Haplotrematidae); living Lymnaea
palustris (Miller) [= Stagnicola elodes(Say));
living Oxyloma decampi (Tryson) [= O. re-
tusa (I. Lea)] and eggs of Oxyloma sp.; living
Physella gyrina (Say) (Physidae); juveniles
and eggs of Stenotrema hirsutum (Say)
(Polygyridae); and living and dead Ventri-
dens demissus (Binney), and living Zoni-
toides arboreus (Say) and Z. nitidus (Miiller)
(Zonitidae) (B. A. Foote, pers. comm.).
Records of 210 adult male and 254 adult
female museum specimens reveal that P.
decora has a distribution typical of many
nearctic sciomyzids. Specimens have been
collected from central Saskatchewan east to
New Brunswick, south to Virginia, and west
to Colorado (Fig. 1). Adults are first seen in
late May, they peak in numbers in July, and
specimens are rarely taken after mid Au-
gust, although one male from Dickinson
County, Michigan, was collected on Sep-
tember 22, 1982. Therefore, it seems likely
the species is univoltine.
DESCRIPTION
Puparium (Fig. 2): Length, 4.8-5.4 mm;
greatest width, 1.7—2.0 mm; greatest height,
1.7-1.9 mm. Unicolorous reddish brown,
but segments 2—4 and 12 somewhat darker
than remainder. Integument opaque, with
finely pebbled sculpturing, especially dor-
476 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
da
rie wp {A
/
Collecting sites for Poecilographa decora.
Fig. 1.
sally and laterally; densely wrinkled on seg-
ments 2-4 and 12. Puparium elongate, sub-
cylindrical, dorsally convex and ventrally
nearly straight in profile. Primary and sec-
ondary integumentary folds faint; 2 second-
ary folds dorsally, 3 ventrally. Segment 1
invaginated. Segments 2—4 strongly tapered
Ay ne Oe ae byes of
“yy RE a
anteriorly. First 2 apparent segments (seg-
ments 2 and 3) dorsoventrally flattened, not
upturned, distinctly narrower than succeed-
ing segments. Anterior spiracles projecting
from anterolateral angles of segment 2, dark
brown, subcircular, bearing about 8 dis-
tinct, marginal papillae. Distinct tubercles
VOLUME 90, NUMBER 4 477
Figs. 2-4. Poecilographa decora. 2, Puparium, cephalic caps separated. 3, Posterior spiracular disc; psp =
posterior spiracle. 4, Cephalopharyngeal skeleton; eps = epipharyngeal sclerite; hps = hypopharyngeal sclerite;
mhk = mouthhook; psb = parastomal bar; tps = tentoropharyngeal sclerite; va = ventral arch; ab/cd = indentation
index.
478
and creeping welts absent from segments 2—
11. Segments 11 and 12 tapered posteriorly.
Segment 12 distinctly ventral in position,
not upturned, truncate, with mid-dorsal in-
dentation. Anus invaginated.
Posterior spiracular disc (Fig. 3) strongly
indented posteroventrally, bearing 4 pairs
of short, wrinkled lobes (indistinguishable
in | specimen) and 2 dorsomedial spiracular
plates. Dorsolateral and lateral lobes short-
est, about as long as diameter of spiracular
plates; ventrolateral and ventral lobes long-
er, 0.35—0.40 mm. Spiracular plates subcir-
cular, at apices of 2 dark brown to black
spiracular tubes; tubes shorter than diam-
eter of plates, longer on lateral surface than
on mesial; each spiracular plate with 3 elon-
gate-oval, diverging, yellow spiracular slits
and a mesial, subcircular, black spiracular
scar; plates lacking well-developed, inter-
spiracular, hairlike processes (float hairs).
Cephalopharyngeal skeleton (Fig. 4)
brown, 0.53 mm long; indentation index
(ab/cd) 44. Paired mandibles not fused, with
decurved mouthhooks; 4-5 decurved, light-
ly pigmented accessory teeth anteroven-
trally; and 2 small windows posterior to ac-
cessory teeth. Ventral arch convex below,
with 14 small teeth on anterior margin,
deeply emarginate posteromesially. Epi-
pharyngeal sclerite fused to anterior ends of
parastomal bars. Posterior ends of para-
stomal bars fused to tentoropharyngeal
sclerites. Hypopharyngeal sclerite H-shaped,
not fused to mandibles or tentoropharyn-
geal sclerites; anterior emargination about
1.5 x length of posterior emargination. Lig-
ulate sclerite anterior to hypopharyngeal
sclerite, small, V-shaped. Paired tentoro-
pharyngeal sclerites not fused, lacking dor-
sal bridge and fenestrations; ventral cornua
distinctly shorter than dorsal cornua.
DISCUSSION
Data from this preliminary study indicate
that Poecilographa decora is a typical mem-
ber of the subfamily Sciomyzinae, tribe Te-
tanocerini, and that the third-instar larva
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and puparium are terrestrial. Morphologi-
cal characteristics that support placement
of this species in the Tetanocerini include
the striate egg chorion, lack of well-devel-
oped ventral spinule patches on the larval
integument, presence of accessory teeth on
the mandibles, lack of a dorsal bridge be-
tween the tentoropharyngeal sclerites, lack
of a window in the dorsal cornu, lack of
tentoropharyngeal-hypopharyngeal fusion,
and a third-instar indentation index of less
than 50. The facts that this species can pu-
pate and oviposit in the absence of a host
also support placement in the Tetanocerini.
Knutson (1966), Knutson et al. (1970), and
Boyes et al. (1969, 1972) provide good sum-
maries of morphological and behavioral dif-
ferences between the Tetanocerini and Scio-
myzini.
The lack of well-developed interspiracu-
lar float hairs on the puparium, the fact that
the posterior end is not upturned (thus not
allowing the spiracles to contact atmospher-
ic air if the puparium were floating), and the
microhabitat of the puparia discussed here
indicate that the immature stages of P. dec-
ora are terrestrial. However, most reared
species of Tetanocerini are aquatic preda-
tors; their larvae live in water, floating just
beneath the surface film, and they attack
snails effectively there as well as on moist
shores or floating vegetation. All reared
species of Sciomyzini are parasitoids in ter-
restrial or semi-aquatic situations, but this
type of feeding behavior is also seen in a
few species of Tetanocerini and the only
reared species of Salticellinae. The tetano-
cerine terrestrial parasitoids are usually host
specific at the species, genus or family level.
As they mature, the larvae become quick-
killing predators and eventually leave the
shell of their last victim to pupate in soil or
litter. A few Tetanocerini feed on decaying
snails as well as fresh prey (Berg and Knut-
son, 1978). These observations suggest han-
dling methods and potential hosts to be used
in future attempts to rear Poecilographa
decora.
VOLUME 90, NUMBER 4
ACKNOWLEDGMENTS
Distributional data or pinned adults for
examination were provided by: D. Azuma,
The Academy of Natural Sciences, Phila-
delphia; H. J. Teskey, Agriculture Canada;
D. Grimaldi, American Museum of Natural
History; P. H. Arnaud, California Academy
of Sciences; B. C. Kondratieff, Colorado
State University; K. R. Valley, Common-
wealth of Pennsylvania, Department of Ag-
riculture; J. K. Liebherr, Cornell Univer-
sity; C. Salvino, Field Museum of Natural
History; S. L. Heydon, Illinois Natural His-
tory Survey; R. E. Lewis, Iowa State Uni-
versity; B. A. Foote, Kent State University;
C.-c. Hsuing, Macdonald College, McGill
University; R. L. Fischer, Michigan State
University; C. Vogt, Museum of Compar-
ative Zoology, Harvard University; E. W.
Balsbaugh, North Dakota State University;
C. A. Triplehorn, Ohio State University; G.
B. Wiggins, Royal Ontario Museum; L.
Knutson, United States National Museum;
R. E. Orth, University of California, Riv-
erside; S. Marshall, University of Guelph;
F. Merickel, University of Idaho; J. K. Gel-
haus, University of Kansas; M. F. O’Brien,
University of Michigan; P. J. Clausen, Uni-
versity of Minnesota; M. Coulloudon, Uni-
versity of Montreal; D. S. Chandler, Uni-
versity of New Hampshire; S. Krauth,
479
University of Wisconsin; and M. Kosztar-
ab, Virginia Polytechnic Institute and State
University. B. A. Foote, L. Knutson, and
K. R. Valley reviewed the manuscript and
provided helpful comments. B. A. Foote
kindly granted permission to incorporate
material from his notes on rearing efforts.
Contribution number 560 of the New
York State Science Service.
LITERATURE CITED
Berg, C. O. and L. Knutson. 1978. Biology and sys-
tematics of the Sclomyzidae. Ann. Rev. Entomol.
23: 239-258.
Boyes, J. W., L. V. Knutson, K. Y. Jan, and C. O. Berg.
1969. Cytotaxonomic studies of Sciomyzidae
(Diptera: Acalyptratae). Trans. Amer. Microsc. Soc.
88: 331-356.
Boyes, J. W., L. V. Knutson, and J. M. van Brink.
1972. Further cytotaxonomic studies of Scio-
myzidae, with description of a new species, Di-
chetophora boyesi Steyskal (Diptera: Acalyptra-
tae). Genetica 43: 334-365.
Johannsen, O. A. 1935. Aquatic Diptera. Part II.
Orthorrhapha—Brachycera and Cyclorrhapha.
Cornell Univ. Agric. Exp. Sta. Mem. 177. 62 pp.
Knutson, L. V. 1966. Biology and immature stages
of malacophagous flies: Antichaeta analis, A. atri-
seta, A. brevipennis, and A. obliviosa. Trans. Amer.
Entomol. Soc. 92: 67-101.
Knutson, L. V., J. W. Stephenson, and C. O. Berg.
1970. Biosystematic studies of Salticella fasciata
(Meigen), a snail-killing fly (Diptera: Sciomyzi-
dae). Trans. Roy. Entomol. Soc. Lond. 122: 81-
100.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 480-483
GENERIC REASSIGNMENTS OF NORTH AMERICAN SPECIES
CURRENTLY ASSIGNED TO THE GENUS SERICOTHRIPS HALIDAY
(THYSANOPTERA: THRIPIDAE)
SuEO NAKAHARA
Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, Belts-
ville, Maryland 20705.
Abstract.—Although the concepts of Sericothrips and related genera were revised in
1973, only 14 of the 51 Sericothrips species heretofore reported from North America (i.e.
Panama northward and the Caribbean Islands) have been reviewed and reassigned to the
proper genera thus far. In order to conform to current concepts, the North American
species were reviewed and assigned to Hydatothrips, Neohydatothrips or retained in Ser-
icothrips, and | species was treated as a nomen dubium. The current scientific names,
distributions, and literature citations for 2 Hydatothrips spp., 47 Neohydatothrips spp.
(including 3 spp. recently described from Mexico) and 3 Sericothrips spp. known to occur
in North America are presented here.
Key Words:
America, new combinations
Ina preliminary revision of the genus Ser-
icothrips Haliday and related genera, Bhatti
(1973: 403) revalidated Hydatothrips Karny
and transferred most of the Sericothrips
species he examined to Hydatothrips and
Neohydatothrips John. Although 51 species
of Sericothrips were reported by Jacot-Guil-
larmod (1971: 359), Johansen (1979: 169)
and Sakimura (1986: 356) from North
America (i.e. from Panama northward and
the Caribbean Islands), only 14 species have
been reevaluated thus far for their proper
generic assignments. Bhatti (1973: 405)
transferred 1 species to Hydatothrips, 7
species to Neohydatothrips and retained 2
species in Sericothrips; Johansen (1983: 107)
transferred 2 species to Neohydatothrips; and
Sakimura in Cho et al. (1987: 507) trans-
ferred | species to Neohydatothrips. O’Neill
(1972: 276) treated S. campestris Hood
(1939: 556) as a junior synonym of N. flor-
idanus (Watson). The remaining species are
Thysanoptera, Thripidae, Sericothrips, Hydatothrips, Neohydatothrips, North
brought into conformity with current con-
cepts here by reassigning | species to Hy-
datothrips, 34 species to Neohydatothrips,
retaining | species in Sericothrips, and treat-
ing | species as a nomen dubium. The ge-
neric assignments are based on the types
except for NV. desmodianus (Stannard), which
is based on identified type material, and N.
muirandai (Johansen), which is based on Jo-
hansen’s opinion (pers. comm., 1987). The
examined specimens are deposited in the
Thysanoptera collection of the U.S. Na-
tional Museum of Natural History (USNM)
located at Beltsville, Maryland.
According to Jacot-Guillarmod (1971:
395), the type depository of Sericothrips tri-
fasciatus (Ashmead) is the USNM, but I
have not been able to find the types. This
species, originally described as Thrips tri-
fasciatus (Ashmead 1894: 27), was assigned
to Sericothrips by Hood (1957: 53) ina foot-
note. He also stated: ““Watson suggested
VOLUME 90, NUMBER 4
(Bull. 168, Fla. Agr. Exp. Sta., p. 44, 1923)
that this might be Franklinothrips vespifor-
mis, but scarcely a word or phrase of Ash-
mead’s description could be applied to that
species.” Hood did not give his reason for
this assignment. Ashmead’s brief descrip-
tion is inadequate for distinguishing the
species. Because the types of trifasciatus
cannot be found, this species is here con-
sidered a nomen dubium.
The following list treats 2 Hydatothrips
spp., 47 Neohydatothrips spp. (including 3
spp. described by Johansen (1983: 107)) and
3 Sericothrips spp. for North America. The
literature citation for the original descrip-
tion is given for each species. References for
species assigned by Bhatti (1973), Johansen
(1983) and Sakimura (1987) are also cited.
The distribution records are based on the
work of Bailey (1957: 195), Beshear (1973:
11; 1979:211), Chiasson (1986: 45), Hunt-
singer et al. (1982: 48), Jacot-Guillarmod
(1971: 359), Johansen (1979: 169: 1983:
107), Sakimura (1985: 30; 1986: 356), Stan-
nard (1968: 345) and material in the USNM
collection. For the United States (US), the
states in postal abbreviations are given.
Hydatothrips Karny
sternalis (Hood) 1935: 148; Bhatti 1973:
405. Dist. Panama.
tricinctus (Hood) 1928: 231. New Combi-
nation. Dist. Brazil, Dominica, Guade-
loupe, Jamaica, Martinique, Trinidad.
Neohydatothrips John
albus (Jones) 1912: 6. New Combination.
Dist. US (CA).
andrei (J.C. Crawford) 1943: 39. New Com-
bination. Dist. US (VA).
annulipes (Hood) 1927a: 211; Bhatti 1973:
405. Dist. US (GA, IA, IL, NJ, NY, VA).
apicalis (Hood) 1927b: 137. New Combi-
nation. Dist. Canada (Alberta), US (IA,
IN, ND).
aztecus Johansen 1983: 113. Dist. Mexico.
baileyi (Hood) 1957: 53. New Combination.
Dist. US (CA).
481
baptisiae (Hood) 1916: 113; Bhatti 1973:
405. Dist. US (GA, IL, MD, NJ, NY,
VA).
basilaris (Hood) 1941: 139. New Combi-
nation. Dist. Cuba.
beachae (Hood) 1927b: 133. New Combi-
nation. Dist. Canada (Alberta), US (IA,
IL, ND).
burungae (Hood) 1935: 150; Johansen 1983:
113. Dist. Jamaica, Mexico, Panama.
catenatus (Hood) 1957: 51. New Combi-
nation. Dist. US (AZ).
chrysothamni (Hood) 1936: 85. New Com-
bination. Dist. US (CA, NV, OR).
collaris (Hood) 1936: 91. New Combina-
tion. Dist. US (AZ, NM).
ctenogastris (Hood) 1936: 93. New Com-
bination. Dist. US (AZ, TX).
desertorum (Hood) 1957: 52. New Combi-
nation. Dist. US (NM).
desmodianus (Stannard) 1968: 351. New
Combination. Dist. US (GA, IL, NJ).
ephedrae (Hood) 1957: 51. New Combi-
nation. Dist. US (AZ, NM).
flavicollis (Hood) 1954: 204. New Combi-
nation. Dist. Brazil, Jamaica.
floridanus (Watson) 1918: 53; Bhatti 1973:
405. Dist. US (FL, GA, IL, MD, MO, TN,
TX, VA).
fraxinicola (Hood) 1940: 545. New Com-
bination. Dist. US (NY).
geminus (Hood) 1935: 146. New Combi-
nation. Dist. Jamaica, Mexico, Panama,
Puerto Rico.
gracilipes (Hood) 1924a: 149; Sakimura in
Cho et al. 1987: 507. Dist. Jamaica, Mex-
ico NSH, 1X):
interruptus (Hood) 1927b: 136. New Com-
bination. Dist. US (GA, IA, IL, MD, NJ).
inversus (Hood) 1928: 232. New Combi-
nation. Dist. Dominica, Jamaica, Pana-
ma, Trinidad.
langei (Moulton) 1929: 230. New Combi-
nation. Dist. US (IL, WI).
mimosae (Hood) 1955: 134. New Combi-
nation. Dist. Costa Rica.
mirandai (Johansen) 1979: 169. New Com-
bination. Dist. Mexico.
482
moultoni (Jones) 1912: 7; Bhatti 1973: 405.
Dist sUS (GA, UT):
nubilipennis (Hood) 1924b: 312. New Com-
bination. Dist. US (IA, IL, ND, MD, NY,
PA, VA).
opuntiae (Hood) 1936: 88. New Combina-
tion. Dist. US (AZ, CA, NM).
pedicellatus (Hood) 1927b: 131. New Com-
bination. Dist. US (IL, NJ, TX).
portoricensis (Morgan) 1925: 3; Bhatti 1973:
405. Dist. Brazil, Cuba, Guadeloupe, Ja-
maica, Panama, Puerto Rico, St. Lucia,
Trinidad.
pseudoannulipes Johansen 1983: 109. Dist.
Mexico.
pulchellus (Hood) 1908: 363. New Combi-
nation. Dist. US (IL, MD).
rapoporti Johansen 1983: 110. Dist. Mexi-
co.
sambuci (Hood) 1924b: 313. New Combi-
nation. Dist. Canada (Ontario), US (DC,
IA, IL, MD, NJ, NY).
sensillis (Hood) 1936: 95. New Combina-
tion. Dist. US (AZ).
setosus (Hood) 1927b: 135. New Combi-
nation. Dist. US (AZ, TX).
signifer (Priesner) 1932: 172: Johansen 1983:
115. Dist. Mexico.
spiritus (Hood) 1927b: 138. New Combi-
nation. Dist. US (AZ).
tibialis (Priesner) 1924: 528: Bhatti 1973:
405. Dist. Mexico.
tiliae (Hood) 1931: 151. New Combination.
Dist. US (IA, IL, IN, ND, NY).
tissoti (Watson) 1937: 4. New Combination.
Dist. US (FL).
variabilis (Beach) 1896: 220; Bhatti 1973:
405. Dist. Canada (British Columbia),
Mexico, US (AL, AR, AZ, CA, DE, GA,
IA, KS, LA, IL, IN, MA, MD, MO, NC,
NJ; OK SC] TNiGEXeU TE VA):
vicenarius (Hood) 1955: 133. New Com-
bination. Dist. US (TX).
williamsi (Hood) 1928: 230. New Combi-
nation. Dist. Mexico, St. Croix.
zebra (Hood) 1940: 543. New Combina-
tion. Dist. US (NY).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Sericothrips Haliday
cingulatus Hinds 1902: 141: Bhatti 1973:
405. Dist. Canada (Alberta, Manitoba),
US (AR, GA, IA, IL, LA, MA, MD, MS,
NE, NY, TN, TX, VA).
pubescens Hood 1957: 50. New Combina-
tion. Dist. US (NY).
smithi Stannard 1951: 129; Bhatti 1973: 405.
Dist. US (GA, IL, NC, SC).
ACKNOWLEDGMENTS
I thank R. M. Johansen, Universidad Na-
cional, Autonoma de Mexico for lending
material for this study and providing infor-
mation on Mexican species, and the follow-
ing colleagues for their reviews of the manu-
script and useful suggestions: L. A. Mound,
British Museum (Nat. Hist.), London: R. J.
Beshear, University of Georgia, Experi-
ment; H. A. Denmark, Florida Dept. of Ag-
riculture and Consumer Services, Gaines-
ville; and R. D. Gordonand A. L. Norrbom,
Systematic Entomology Laboratory, Wash-
ington, D.C.
LITERATURE CITED
Ashmead, W.H. 1894. Notes on cotton insects found
in Mississippi. Ins. Life 7(1): 25-27.
Bailey, S. F. 1957. The thrips of California, Part I:
Suborder Terebrantia. Bull. California Ins. Surv.
4(5): 143-220.
Beach, A. M. 1896. Contributions to a knowledge of
the Thripidae of Iowa. Proc. Iowa Acad. Sci. for
1895 3: 214-228.
Beshear, R. J. 1973. The thrips of Georgia, suborder
Terebrantia. Univ. Georgia, Coll. Agric. Exp. Sta.
Res. Bull. 122, 26 pp.
. 1979. Additional records of thrips in Georgia
(Thysanoptera: Terebrantia). J. Georgia Entomol.
Soc. 14(3): 209-211.
Bhatti, J. S. 1973. A preliminary revision of Seri-
cothrips Haliday, sensu lat., and related genera,
with a revised concept of the tribe Sericothripini
(Thysanoptera: Thripidae). Oriental Insects 7(3):
403-449.
Chaisson, H. 1986. A synopsis of the Thysanoptera
(thrips) of Canada. McGill Univ., Lyman Ento-
mol. Mus. Res. Lab. Mem. No. 17, 153 pp.
Cho, J. J., W. C. Mitchell, R. F. L. Mau, and K. Sak-
imura. 1987. Epidemiology of tomato spotted
VOLUME 90, NUMBER 4
wilt virus disease on crisphead lettuce in Hawai.
Plant Disease 71: 505-508.
Crawford, J. C. 1943. A new Sericothrips on elm
(Thysanoptera: Thripidae). Proc. Entomol. Soc.
Wash. 45(2): 39-41.
Hinds, W. E. 1902. Contribution to a monograph of
the insects of the order Thysanoptera inhabiting
North America. Proc. U.S. Nat. Mus. 26: 79-242.
Hood, J. D. 1908. New genera and species of Illinois
Thysanoptera. Bull. Illinois St. Lab. Nat. Hist. 8(2):
361-379.
1916. Descriptions of new Thysanoptera.
Proc. Biol. Soc. Wash. 29: 109-123.
1924a. A new Sericothrips (Thysanoptera)
injurious to cotton. Can. Entomol. 56(6): 149-150.
1924b. New Thysanoptera from the United
States. Entomol. News 35(9): 312-317.
. 1927a. A contribution toward the knowledge
of New York Thysanoptera, with descriptions of
new genera and species. II]. Entomol. Americana
7(4): 209-245.
1927b. New Thysanoptera from the United
States. J. New York Entomol. Soc. 35: 123-142.
1928. New Neotropical Thysanoptera col-
lected by C. B. Williams. II. Psyche 34(6): 230-
246.
1931. Notes on New York Thysanoptera,
with descriptions of new genera and species. III.
Bull. Brooklyn Entomol. Soc. 26(4): 151-170.
1935. Eleven new Thripidae (Thysanoptera)
from Panama. J. New York Entomol. Soc. 43:
143-171.
1936. Nine new Thysanoptera from the
United States. J. New York Entomol. Soc. 44: 81-
100.
1939. New North American Thysanoptera,
principally from Texas. Rev. Entomol. 10(3): 550-
619.
1940. A century of new American Thysan-
optera. I. Rev. Entomol. 11(1-2): 540-583.
. 1941. Acentury ofnew American Thysanop-
tera. II. Rev. Entomol. 12(1-2): 139-243.
1954. Brasilian Thysanoptera. V. Proc. Biol.
Soc. Wash. 67: 195-214.
1955. New American Terebrantian Thysa-
noptera. J. New York Entomol. Soc. 62(3): 129-
138.
1957. Fifteen new Thysanoptera from the
United States. Proc. Biol. Soc. Wash. 70: 49-60.
483
Huntsinger, D. M., R. L. Post, and E. U. Balsbaugh
Jr. 1982. North Dakota Terebrantia (Thysanop-
tera). North Dakota Insects, Schafer-Post Series
No. 14, 102 pp.
Jacot-Guillarmod, C. F. 1971. Catalogue of the Thy-
sanoptera of the world, Part 2. Ann. Cape Prov.
Mus. (Nat. Hist.) 7(2): 217-515.
Johansen, R. M. 1979. Seis nuevos thisanopteros
(Terebrantia: Heterothripidae; Thripidae), de
Chiapas, Mexico. Ann. Inst. Biol. Univ. Nal. Au-
ton. Mexico 50 (Zool. Ser. 1): 159-178.
. 1983. Nuevos thrips (Insecta: Thysanoptera;
Terebrantia, Thripidae: Thripinae), de la Sierra
Madre Oriental y del eje Volcanico Transversal,
de Mexico. Ann. Inst. Biol. Univ. Nal. Auton.
Mexico 53 (Zool. Ser. 1): 91-132.
Jones, P.R. 1912. Some new California and Georgia
Thysanoptera. Misc. Paper, U.S. Dept. Agric. Bur.
Entomol. Tech. Ser. 23(1): 1-24.
Morgan, A. C. 1925. A new genus, a new subgenus
and seven new species of Thysanoptera from Porto
Rico. Florida Entomol. 46: 1-55.
Moulton, D. 1929. Contribution to our knowledge
of American Thysanoptera. Bull. Brooklyn Ento-
mol. Soc. 24(4): 224-244.
O'Neill, K. 1972. Mycterothrips Trybom, a review of
the North American species (Thysanoptera:
Thnpidae). Proc. Entomol. Soc. Wash. 74(3): 275-
282.
Priesner, H. 1924. Neue Thysanopteren. Sitzungsb.
Akad. Wiss. Wien (1) 133(10): [S27]-542.
1932. Neue Thysanopteren aus Mexico, ge-
sammelt von Prof. Dr. A. Dampf. Wiener Ento-
mol. Zeitung 49(3): [170]-185.
Sakimura, K. 1985. Notes and exhibitions. Some new
thrips record for Hawaii. Proc. Hawaiian Ento-
mol. Soc. 25: 30.
. 1986. Thrips in and around the coconut plan-
tations in Jamaica, with a few taxonomical notes
(Thysanoptera). Florida Entomol. 69(2): 348-363.
Stannard, L. J. 1951. Anew Sericothrips from Ilinois
(Thysanoptera: Thripidae). Kansas Entomol. Soc.
24(4): 129-130.
1968. The thrips, or Thysanoptera, of [hi-
nois. Bull. Illinois Nat. Hist. Surv. 29(4): 215-552.
Watson, J.R. 1918. Thysanoptera of Florida. Florida
Buggist 1(4): 53-55.
1937. Sericothrips with an unusual habitat.
Florida Entomol. 20(1): 3-4.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 484-494
REDESCRIPTION OF THE HOLOTYPE OF CULEX (CULEX) PEUS SPEISER
AND TAXONOMY OF CULEX (CULEX) STIGMATOSOMA DYAR
AND THRIAMBUS DYAR (DIPTERA: CULICIDAE)
Daniel Strickman
Walter Reed Biosystematics Unit, Department of Entomology, Walter Reed Army
Institute of Research. Mailing address: Museum Support Center, Smithsonian Institution,
Washington, D.C. 20560.
Abstract. —Examination and redescription of the holotype Culex (Culex) peus Speiser
1904 (substitute name for affinis Adams 1903) established that it is conspecific with Cx.
thriambus Dyar 1921. Therefore, Cx. thriambus is synonymized under peus; Culex stig-
matosoma Dyar 1907 is resurrected from synonymy; and Cx. eumimetes Dyar and Knab
1908 is transferred to synonymy under stigmatosoma. This paper also clarifies identifi-
cation of these species in the literature.
Key Words:
description
While studying the Culex (Culex) of Cen-
tral America, I noticed several differences
between descriptions (Adams 1903, Stone
1958) of the holotype of Culex peus Speiser
1904 (= affinis Adams 1903) and other spec-
imens and descriptions of the species. As a
result, I undertook a study with the purpose
of identifying the holotype. Since the ho-
lotype is a damaged female adult from Ar-
izona, I examined female adult specimens
of Cx. (standard abbreviation for Culex,
Reinert 1975) peus and the similar species,
Cx. thriambus Dyar 1921, with the objec-
tive of finding diagnostic characters still
present on the holotype of Cx. peus. Ex-
amination of specimens was limited to the
United States in order to assure that the
specimens belonged to one of the two species
involved and not to a possibly undescribed
form from Mexico or Central America.
The study established that the holotype
of Cx. peus is conspecific with Cx. thriam-
bus. As a result, Cx. thriambus is made a
synonym of Cx. peus. Furthermore, Cx.
stigmatosoma Dyar 1907 is resurrected from
Culex, affinis, eumimetes, peus, stigmatosoma, thriambus, taxonomy, re-
synonymy with Cx. peus. Culex eumimetes
Dyar and Knab 1908 is transferred from
synonymy under Cx. peus to synonymy un-
der Cx. stigmatosoma. Specimens identified
as Cx. peus since Stone (1958) are actually
Cx. stigmatosoma, and specimens identi-
fied as Cx. thriambus are Cx. peus.
The holotype of Cx. peus is redescribed
in this paper in much greater detail than by
Adams (1903) or Stone (1958). This de-
tailed description was considered necessary
because of the central role of the holotype
in the nomenclature of the species involved.
The redescription documents characters that
are not currently known to be significant,
but which could conceivably influence fu-
ture taxonomic decisions. Since the holo-
type is already damaged, redescription helps
assure that any future deterioration will not
result in permanent loss of characters.
METHODS
Evaluation of color on the holotype of Cx.
affinis was based on comparison of the spec-
imen to color samples of the four-color
VOLUME 90, NUMBER 4
printing process. The color samples (Kuep-
pers 1982) present mixtures of black (B),
cyan (C), magenta (M), and yellow (Y) in
all combinations of three of the colors against
a white page at 10% intervals (i.e. percent-
age coverage of the page with minute dots
used in color printing). A particular color is
designated as a combination of the per-
centage of each of the color inks (e.g.
B,o>M9Coo is a sky blue color). Each color
sample was viewed surrounded by a gray
mat under unfiltered tungsten light. For col-
or evaluation, the specimen was also viewed
under unfiltered tungsten light set at 5 volts.
Unfortunately, the same color may appear
as more than one combination of inks and,
the human eye is much more sensitive to
certain color ranges, such as light yellow,
than the 10% intervals can identify. There-
fore, light yellow scales have been called
“yellowish” in the description. Neverthe-
less, the system is useful because it provides
an objective reference to color and a mea-
surement of color that is reproducible on a
printed page.
Abbreviations and notations require some
explanation. The symbols “*4” and “2” rep-
resent adults of the respective sex. The sym-
bol “6G” is male genitalia. Fourth instar
larva is represented by “L”’ and pupa 1s rep-
resented by “P.” An asterisk indicates that
the stage was illustrated in the cited paper.
Where possible, collection or specimen
numbers were reported to allow location of
the exact specimen examined. All speci-
mens are in the U.S. National Museum
(USNM) unless otherwise noted (UAz =
University of Arizona, SEM = Snow En-
tomological Museum, University of Kan-
sas). Morphological nomenclature and ab-
breviations were taken from Harbach and
Knight (1980).
TAXONOMY
Culex peus Speiser
Culex peus Speiser, 1904: 148, replacement
name for affinis Adams.
Culex affinis Adams, 1903: 25, Oak Creek
485
Canyon, Arizona, USA, 2°, SEM; Coquil-
lett 1904: 261, synonymized under Cx.
tarsalis,; Theobald 1907: 394, synonymy
questioned.
Culex thriambus Dyar, 1921: 33, Kerrville,
Texas, USA, 6, USNM. New Synonymy;
Dyar 1928: 368, synonymized under stig-
matosoma; Edwards 1932: 206, listed as
var. of stigmatosoma; Galindo and Kel-
ley 1943: 87, resurrected.
Additional descriptions.— Cited as Cx.
peus: Stone 1958 (2). Cited as Cx. thriam-
bus: Dyar 1921 (6, 2, 6G, L). Dyar 1922 (8,
3, L); Galindo and Kelley 1943 (2, 4G, L);
Freeborn and Brookman 1943 (¢, L):; Free-
born and Bohart 1951 (2, 6G, L); Breland
1957 (L); Martinez Palacios 1952 (4, 2, 8G*,
L); Usinger et al. 1952 (2, L); Bohart and
Washino 1957 (2nd and 3rd instar L); Car-
penter and LaCasse 1955 (9*, 6, 6G*, L*);
Dodge 1963 (L); Nielsen and Linam 1963
(2, L); Myers 1964 (L); Forattini 1965 (8,
6G, L); Chapman 1966 (2, 6G, L); Cova Gar-
cia et al. 1966a (2, 6G*); Cova Garcia et al.
1966b (L*); Dodge 1966 (Ist instar L); Mu-
kherjee et al. 1966 (chromosomes* of L);
Bram 1967 (9, 8G*, L); Nielsen 1968 (2, 4G,
L); McDonald et al. 1973 (2); Bohart and
Washino 1978 (2*, L*); Darsie and Ward
1981 (9*, L*): Clark-Gil and Darsie 1983 (2,
L).
Material examined (all adult females). —
Arizona: Coconino Co.: Oak Creek Canyon,
holotype, F. H. Snow. Cochise Co.: St. Da-
vid, 24 Sep 1953, C. S. Richards, 2 2. Mar-
icopa Co.: Wickenburg, 29 Jun 1953, W.
W. Wirth. Pima Co.: Lake Sabino Canyon,
20 Oct 1962, J. Burger coll. no. 349, 5 9; 17
Nov 1962, coll. no. 353; 17 Nov 1962, coll.
no. 357, 2 2; 10 Mar 1963, coll. no. 373, 3
2; 20 Apr 1963, coll. no. 378; 26 May 1963,
coll. no. 383, 2 2; 28 Jun 1963, coll. no. 388;
17 Oct 1963, coll. no. 410, 3 2. Pinal Co.:
Boyce-Thompson Arboretum, 3 mi. S. of
Superior, 7 Jul 1963, J. Burger coll. no. 390,
9 ¢. Santa Cruz Co.: Madera Canyon, 21-
26 Aug 1954, W. A. McDonald coll. no.
486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Characters of various populations of Culex stigmatosoma and Culex peus, including the type
specimens, specimens from the states of type localities, and all specimens examined from the United States.
Percentages are followed by 95% confidence limits (CL) (Rohlf and Sokal 1969) and means are followed by
standard deviations (SD).
Proboscis Band Complete
Palpi with White Scales
Ratio of Basal Band
HT-5 with Dark Band to Length of HT-2
Population % (CL) (n) 0 (CL) (n) % (CL) (n) Mean + SD (n)
affinis type yes? no no* 0.06
Arizona peus 24 (10-41) (38) 3 (0-16) (37) 30 (17-52) (33) 0.09 + .02 (38)
Texas peus 10 (0-44) (10) QO (0-29) (11) 22 (3-62) (9) 0.08 + .02 (10)
California peus 15S (6-34) (34) 0 (0-11) (34) 42 (25-62) (33) 0.08 + .02 (33)
All peus 18 (10-29) (82) 1 (0-8) (82) 35 (24-47) (75) 0.08> + .02 (81)
stigmatosoma type yes yes yes 0.15
Arizona stigmatosoma 100 (71-100) (11) 100 (71-100) (11) 100 (65-100) (9) O:S- = -02;014)
California stigmatosoma 98 (93-100) (127) 100 (97-100) (126) 100 (97-100) (127) 0.13 + .02 (128)
All stigmatosoma 98 (93-100) (139) 100 (97-100) (138) 100 (97-100) (137) 0.13 + .02 (140)
* Destroyed on type specimen, extracted from original description (Adams 1903).
> Significantly different at the 95% level in a t-test of group means.
133, 8 2. California: Inyo Co.: China Ranch,
29-30 Oct 1955, Blodget and McDonald
coll. no. 177, 10 2. Riverside Co.: 1 mi. S.
Hurkey Creek Campground, San Jacinto
Mts., 9 Apr 1962, C. L. Hogue coll. no. 233,
5 9. San Bernardino Co.: Saratoga Springs,
Death Valley, 29 Oct 1955, Blodget and
McDonald coll. no. 176, 8 2. San Diego Co.:
Jamacha Junction, 11 Jul 1954, Belkin and
McDonald coll. no. 124, 10 ¢. San Luis
Obispo Co.: San Luis Obispo, 22 Aug 1948,
W. W. Wirth. Shasta Co.: US 299 and Trin-
ity Center Rd., 8 Sep 1950, J. N. Belkin
coll. no. 68. Texas: Bexar Co.: San Antonio,
14 Jul 1942, E. S. Ross, 2 2. Kerr Co.: Kerr-
ville, 20 Aug 1920, H. G. Dyar coll. no. Y2,
coll. no. Y4, coll. no. Y7 (type no. 23926),
coll. no. Ya 5 2. Travis Co.: Austin, 24 Oct,
A. L. Melander.
Diagnosis. —The adult female of Cx. peus
may be distinguished from Cx. stigmato-
soma on the basis of characters presented
in Table 1. In contrast to Cx. stigmatosoma,
the proboscis band of Cx. peus is usually
incomplete; the palpi lack broad, opaque
white scales; the dark band in the middle
of hindtarsomere 5 is usually absent making
this tarsomere completely white; and the
width of the basal light band on hindtar-
somere 2 is usually less than 0.10 of length
of hindtarsomere 2.
Culex peus sometimes has a complete
proboscis band and hindtarsomere 5 with a
dark band, character states usually associ-
ated with stigmatosoma. Only three ex-
amples out of 81 specimens displayed both
these characters. Of these, two (Inyo Co.,
San Diego Co., CA) were associated with
exuviae definitely identified as Cx. peus. The
third (Riverside Co., CA) was from a col-
lection of more typical adult females of Cx.
peus. Conversely, three female adults of Cx.
stigmatosoma displayed an incomplete pro-
boscis band, a character typical of Cx. peus.
Two of these specimens were from collec-
tions of stigmatosoma. The third was the
sole individual in a collection (Huntington
Beach, Orange Co., CA).
Although the proboscis band does not
provide complete separation of the species,
it is an important character (Table 1). As in
other members of the subgenus, light scales
on the middle of the proboscis form a band
that is either complete or incomplete dor-
sally. Bands with a narrow dark dorsal line
less than one scale wide were considered
complete. The bands seem to be the result
of a separation of the two sides of the labial
sheath, exposing the dark, unscaled stylets
within. The majority (82%) of Cx. peus had
an incomplete proboscis band restricted to
the ventral and lateral portions of the la-
VOLUME 90, NUMBER 4
bium. Although the proportion of speci-
mens with a complete proboscis band var-
ied from 10% in Texas to 24% in Arizona,
the differences between regions were not sig-
nificant at the 95% level, as judged by con-
fidence limits. Almost all Cx. stigmatosoma
specimens had complete proboscis bands
with a greater density of whiter scales than
in Cx. peus. There was no consistent differ-
ence between the two species in the length
of the band.
The presence or absence of white scales
on the palpi is a consistent difference be-
tween the species (Table 1). Every specimen
of Cx. stigmatosoma examined had at least
several large, opaque, white scales on the
dorsal and mesal sides of the apex of the
palpi. Often, the scales formed large, dis-
tinctive patches. Most Cx. peus lacked large,
opaque white scales on the palpi. The palpi
were either entirely dark scaled, or had small,
light, pearly scales on some of the surfaces.
Only one specimen had opaque white scales;
five scales were on one palpus.
Hindtarsomeres 1—4 of both species were
ornamented with white basal and apical
bands. Hindtarsomere 5 (HT-5) always fol-
lowed this same pattern in Cx. stigmato-
soma, displaying a distinct dark band in the
middle of the tarsomere. Culex peus varied
in this character, with 35% of those exam-
ined having HT-5 with a dark band and the
remaining specimens with HT-5 all white
(Table 1). As in the case of the proboscis
band, the proportion of peus with the dark
HT-5 band varied among populations, but
not significantly so. Generally, the dark-
scaled portion of HT-5 was not as distinct
In peus as in Cx. stigmatosoma.
Another character useful for separating
the two species was the ratio of the width
of the basal band to tarsomere length on
HT-2 (Table 1). The mean ratio was 0.13
for Cx. stigmatosoma and 0.08 for peus.
There were no significant differences be-
tween populations within each species. Fig-
ure 12 presents the data as frequency dis-
tributions, showing that the central value of
487
the ratio is different for the two species,
though the distributions overlap.
Remarks.—The lectotype of Cx. thriam-
bus and associated specimens conform to
the description of Cx. peus given above. The
lectotype was selected by Stone and Knight
(1957) from three syntypes designated by
Dyar. One female has the same accession
number (USNM Type No. 23926) and label
information as the type, including Dyar’s
code “Y7.’ Seven other females were col-
lected on the same date by Dyar in Kerr-
ville, but have different code numbers. Since
it is not clear what Dyar intended by his
code numbers (A. Stone, personal com-
munication; search of Smithsonian Ar-
chives failed to find relevant notes or let-
ters), the eight females may have come from
the same collection despite the application
of four different code numbers, lending con-
fidence to the assumption that the females
are the same species as the lectotype male.
One of the specimens has a dark band on
HT-5, six have HT-5 all white, and one
lacks HT-5 on both hindlegs (code number
Y7, USNM Type No. 23926). All of the
females lack white scales on the palpi and
have incomplete proboscis bands (one has
no proboscis). The mean value for the ratio
of the length of the light basal band to the
length of HT-2 for seven of the specimens
is 0.075 with a range of 0.065 to 0.087.
Redescription of holotype (Figs. 1-11).—
Condition of specimen: Specimen damaged.
On head, proboscis missing up to clypeus
except for short segment of single internal
stylet. Front of head collapsed horizontally
so that vertex overlies pedicels. Scales ob-
viously missing from parts of vertex, though
pattern and color of scales still discernible.
Antennae broken, all flagellomeres beyond
pedicels missing. Damage to thorax caused
by original pinning. No. | insect pin pierces
thorax, obscuring center of scutum and low-
er right pleuron. Area posterior to pin gen-
erally less rubbed of scales and setae than
area anterior to pin. Missing portions of legs:
left foretrochanter, forefemur, foretibia, and
488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
eee
De See ~
SOS
1 Tae PDI
Figs. 1-11. Culex peus Speiser, holotype. All scale bars are in millimeters. 1. Anterior side of hindleg (HT-4,
5 missing). 2. Posterior side of hindleg (HT-4, 5 missing). 3. Anterior side of midleg. 4. Posterior side of midleg.
5. Anterior side of foreleg. 6. Posterior side of foreleg. 7. Lateral view of head. 8. Dorsal view of head and
thorax. 9. Dorsal view of abdomen. 10. Lateral view of thorax with small piece of abdominal tergite I. 11.
Dorsal view of wing.
VOLUME 90, NUMBER 4
UK|
AI
Z
489
-type of affinis
Z
[__]-stigmatosoma
©
©
-type of
© stigmatosoma
LiL
Interval Midpoints of Ratio of
Basal Band to Length of Hindtarsomere - 2
Z
Z
Fig. 12.
Frequency distribution of the ratio of the width of the white basal band on hindtarsomere 2 to the
length of hindtarsomere 2 on female Culex peus (n = 81) and stigmatosoma (n = 140) from the United States.
foretarsus; left midfemur, midtibia, and
midtarsus; and left and right hindtarso-
meres 4 and 5. Portions of abdomen rubbed,
but scale patterns visible on all segments.
Gravid condition of abdomen has stretched
it in such a way that sternites hidden by
paper of label, to which abdomen is glued.
Head: Palpomere 3 clothed in dark brown
(Boy Y7o.Mo,) scales, broader on dorsal side
than on ventral. Integument of each pedicel
dark mesally. Scape about same color as
lighter parts of pedicel. White decumbent
scales of vertex and occiput narrow, flat,
curved, and end in fine point. Broad scales
on ocular suture and on postgena either
truncate or rounded at tip. Erect, furcate
scales on occiput and lateral potions of ver-
tex. Integument of vertex medium brown
(B,Y so>Mg.). Postocciput darkly pigmented
on lateral edge and coronal suture.
Dorsum of thorax: Anterior promontory
with 16 narrow, curved, flat, pointed white
scales. Lateral scutal fossal scales white,
similar to scales on anterior promontory but
broader. Median scutal fossal scales yellow-
ish, slightly curved, and uniformly wide
along length. Supraalar scales and lateral
prescutellar scales white, narrow, flat,
curved, and pointed. All undamaged setae
alike. Integument dark brown (Boy Y 79>M go)
with darker brown (Boo>M4 C,,) acrostichal
area. Scutellar scales narrow, curved, white,
and pointed; lateral scutellar scale groups (7
scales on left, 5 scales on right) smaller than
median scutellar scale group (greater than
30 scales). Each lateral scutellar lobe with
sockets for 5 large setae arranged in row of
4 ventrally and one dorsally. Insertions of
6 median scutellar setae in same plane. In-
tegument of scutellum lighter than that of
prescutellum.
Pleuron: Integument of pleural sclerites
490
light brown (BooY7o>M4.) or dark brown
(Boo ¥ 79Mgo). Positions and shapes of setae
as illustrated (Fig. 10). Antepronotum uni-
formly light brown; | 1 broad, truncate, white
scales on ventral portion. Postpronotum
with narrow, curved, flat, pointed scales
grouped on dorsal half of sclerite; most scales
yellowish, dorsal few white; integument
other than ventroposterior portion dark
brown. Proepisternum light brown with
white area just ventrad of setae. On mesan-
episternum, postspiracular area with 8
broad, rounded, white scales; integument
dark on post- and subspiracular areas, light
on hypostigmal area. Mesokatepisternum
with 21 broad, rounded, white scales; in-
tegument dark on most of ventral half and
central portion of prealar knob; edges of
prealar knob and of ventral half of meso-
katepisternum light. Paratergite apparently
lacks scales and setae (surface partially ob-
scured by shrinkage of pleuron). Basalare,
pleural wing process, and subalare with pale
integuments. Two groups of broad, round-
ed, white scales on mesanepimeron; |2 scales
in upper group, |1 scales in lower group.
Integument dark in center, light on edges.
Integument of mesokatepimeron, metepi-
sternum, and metameron pale; mesomeron
and mesotrochantin dark.
Wings: Scales of costa, subcosta, radius,
and radius-one broad and either rounded or
truncate; some broad scales paler than other
scales. Clear membrane between veins mi-
nutely stippled. Knob of haltere clothed in
minute pale scales; integument darker on
knob than on stem.
Legs: Pale scales probably discolored with
age (white on recently collected specimens
of Cx. peus); others dark brown (By)MooCo0)-
Femora, tibiae, and tarsi illustrated (Figs.
1-6). Forecoxa with 14 small, round, white
scales dorsally and inconspicuous scales
ventrally colored like integument; on left
side, ventral scales arranged in row below
white scales, followed ventrally by loosely
scattered scales; on right side, ventral group
of scales more densely arranged than on left
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
side. Foretrochanter with a few scattered
small, light-colored scales on ectal surface;
integument light except for darkening at
apical margin. Midcoxa with 5 broad, white
scales on middle of anterior surface. Tro-
chanter with six light, broad scales on mesal
surface; integument of posterior apical mar-
gin darkly pigmented. Hindcoxa has scat-
tered broad white scales on ectal surface.
Hindtrochanter with scattered light scales
on mesal and ventral sides; integument
darkened apically on mesal and ventral
sides.
Abdomen: Pattern of white and dark scales
as illustrated (Fig. 9). Integument appears
to darken posteriorly on each segment.
Culex stigmatosoma Dyar
Culex stigmatosoma Dyar, 1907: 123, Pas-
adena, California, USA, 2, USNM; Stone
1958: 236, synonymized under peus.
Culex eumimetes Dyar and Knab, 1908: 61,
Orizaba, Mexico, 6, USNM. New Syn-
onymy.
Additional descriptions.—Cited as Cx.
stigmatosoma: Dyar 1907 (2, L). Howard et
al. 1912, 1915 (2, 6G*, L*); Dyar and Knab
1917 (8G, L); Dyar 1922 (2); Freeborn 1926
(2, 6, 3G*, L); Dyar 1928 (in part peus: 2, 3,
4G*, L*); Aguilar 1931 (4G, L); Martini 1935
(2); Ripstein 1935 (9*, 6, 6G*, L*); Aitken
1942 (Aitken’s identification tentative: 2, L);
Galindo and Kelley 1943 (2, 6G, L); Rees
1943 (2, 6G, L); Freeborn and Brookman
1943 (2, L); Matheson 1944 (2, 4G, L); Pierce
et al. 1945 (2); Martinez Palacios 1950 (in
part peus: 6G*); Freeborn and Bohart 1951
(2, 6G, L*); Usinger et al. 1952 (2, L); Mar-
tinez Palacios 1952 (&G*); Stage et al. 1952
(2, 6G*, L); Lane 1953 (in part peus: 2, 6G*,
L*); Carpenter and LaCasse 1955 (9*, 4, 6G*,
L*); Breland 1957 (L*); Bohart and Wash-
ino 1957 (2nd and 3rd instars L*). Cited as
Cx. eumimetes: Howard et al. 1912, 1915
(2, , 6G*, L*); Dyar 1918 (6, 8G, L). Cited
as Cx. peus: Dodge 1963 (L); Myers 1964
(L*); Forattini 1965 (2, 8G*, L*); Cova Gar-
VOLUME 90, NUMBER 4
cia et al. 1966a (2, 6G*); Cova Garcia et al.
1966b (L*); Chapman 1966 (2, 6G, L); Bram
1967 (2, 6G*, L); Gjullin and Eddy 1972 (2,
8G*, L*); McDonald et al. 1973 (F); Bohart
and Washino 1978 (2, L*); Darsie and Ward
NOST(e%s IS*):
Material examined (all adult females). —
California: Los Angeles Co.: Pasadena, ho-
lotype, 21 May 1906, Dyar and Caudell coll.
no. C78. Arizona: Cochise Co.: Douglas, 23
Aug 1939, T. K. Ryan, 2 2; Lowell, 2 Aug
1939, T. K. Ryan, 3 2; Tombstone, | Sep
1939, T. K. Ryan. Pima Co.: Lake Sabino
Canyon, 17 Aug 1963, J. Burger coll. no.
399, 2 2: Tucson, Jul 1920 (UAz):; Tucson,
9 Feb 1941, R. A. Flock (UAz). Santa Cruz
Co.: 2 mi. W. of Patagonia, 24 Aug 1954,
W. A. McDonald. California: Alameda Co.:
Oakland, I. McCracken: 24 Jul 1903, 6 2;
26 Aug 1903, 4 2. Clear Lake Co.: Rocky
Point, 9 Nov 1947, H. P. Chandler. Contra
Costa Co.: Richmond, 3 Oct 1947, W. W.
Wirth. Humboldt Co.: Fortuna, 13 Aug
1948, W. W. Wirth. Kings Co.: Hanford, 8
Jul 1947, W. W. Wirth. Los Angeles Co.:
Bixby, 25 Jul 1949; Chilao Flat, San Gabriel
Mts., 18 Aug 1955, C. L. Hogue, 5 2; Mal-
ibu, 17 Sep 1952; Malibu Beach, 30 Nov
1963, T. J. Zavortink coll. no. 487, 9 2;
Malibu Beach, 17 Dec 1963, T. J. Zavortink
coll. no. 488; Pasadena, 21 May 1906, Dyar
and Caudell coll. no. C78, 11 2; Reseda, 25
May 1955. Marin Co.: Ft. Barry, 20 Sep
1957, Carpenter et al., 8 2; Lucas Valley, 10
Sep 1957, Carpenter et al., 9 2. Mariposa
Co.: Mariposa Co., 20 May 1960, A. R.
Barr, 5 2. Merced Co.: Snelling, R. M. Bo-
hart. Monterey Co.: Monterey, 10 Aug 1945.
Orange Co.: Alyso Canyon, 10 Oct 1952, J.
N. Belkin coll. no. 91, 16 2; Buelia Park, 22
Jul 1949, 6 2; Buena Park, 6 Jun 1949, 3 9;
Buena Park, 22 Jul 1949, 2 2; Huntington
Beach, 17 Jul 1949; Irvine Park, 24 Jun
1949; Laguna Beach, 4 Jun 1949; Orange
Co., 23 Jul 1950; San Juan Capistrano, 29
Jul 1949; Santa Ana, 2 Jun 1949; Santa Ana,
22 Jul 1949, 2 2. San Diego Co.: San Diego,
H. G. Dyar: 10-18 Apr 1916, coll. no. C, 4
49]
9; 17 Apr 1916, coll. no. C7, 2 2; 17 Apr
1916, coll. no. ABC, 2 2; 5 May 1916. Santa
Clara Co.: Mt. View, 15 Jul 1903, I.
McCracken; Stanford, I. McCracken: 26
May 1903, 5 9; 27 May 1903, 3 2; 8 Jul
1903; 10 Jul 1903; Stanford, 15 Jul 1961,
A. L. Melander. Solano Co.: Vacaville, 4
Jul 1949, R. M. Bohart. Tulare Co.: Coffee
Canyon, Tulare River, 29 Jul 1947, W. W.
Wirth. Ventura Co.: Lake Sherman, |7 Sep
1952, J. N. Belkin. Oregon: Curry Co.: Har-
bor, 8 Oct 1944, W. W. Yates.
Diagnosis.—See diagnosis for Cx. peus.
Remarks.—The holotype (Stone and
Knight 1957) of Cx. stigmatosoma is a fe-
male adult from Pasadena, California, col-
lected by H. G. Dyar in 1906 and part of a
long series of reared specimens. The type is
in excellent condition and conforms com-
pletely to the diagnosis of Cx. stigmatoso-
ma. Larvae and male genitalia from the same
collection fit descriptions of these stages in
recent literature.
The lectotype of Cx. eumimetes 1s a male
selected by Stone and Knight (1957) from
a series of 10 originally collected by Knab
in 1908 in Orizaba, Mexico. The genitalia
of the lectotype are not mounted, but the
appearance of the specimen is consistent
with other male Cx. stigmatosoma. The
mounted genitalia from one of the other 10
specimens in the original series (no. 437.2)
is definitely that of Cx. stigmatosoma based
on the presence of seta d on the subapical
lobe of the gonocoxite (Bram 1967).
DISCUSSION
Examination of specimens from the
United States showed that adult females of
Cx. peus and Cx. stigmatosoma are usually
distinguishable by the proboscis band (usu-
ally incomplete in Cx. peus, complete in Cx.
stigmatosoma), white scales on the palpi
(absent in Cx. peus, present in Cx. Stig-
matosoma), a dark band in the middle of
hindtarsomere 5 (usually absent in Cx. pews,
present in Cx. stigmatosoma), and the width
of the basal band on hindtarsomere 2 (nar-
492
rower in Cx. peus, wider in Cx. stigmato-
soma). The only constant character for sep-
arating the female adults of the species was
presence or absence of white scales on the
palpi. The other three characters were use-
ful, however, because very few individuals
had more than one of the other 3 character
states from the opposite species.
The holotype of Cx. peus (= affinis) was
more similar to material formerly desig-
nated Cx. thriambus and dissimilar to the
holotype of Cx. stigmatosoma. Adams
(1903) described hindtarsomere 5 on Cx.
affinis as all white, a character state present
in the majority of Cx. peus (formerly
thriambus) specimens and never present in
Cx. stigmatosoma. Also, the holotype of Cx.
peus lacks the white scales on the palpus
always associated with Cx. stigmatosoma
and never associated with Cx. peus. Finally,
the width of the basal band on hindtarso-
mere 2 1s within the range of Cx. peus and
outside the range of Cx. stigmatosoma (Fig.
12). Adams (1903) implied that the pro-
boscis band of the holotype of Cx. peus was
complete, a condition more typical of Cx.
stigmatosoma, but commonly present in in-
dividuals of Cx. peus.
Previous descriptions mentioned some of
the characters used to identify the holotype
of Cx. peus. The descriptions attributed a
complete proboscis band to Cx. stigmato-
soma and, with only two exceptions (Dyar
1921, Martinez Palacios 1950), an incom-
plete proboscis band to Cx. peus. Since Cx.
peus often has a complete proboscis band
(Table 1), the use of this character to sep-
arate Cx. stigmatosoma and Cx. peus has
probably led to misidentifications. The
presence of light and dark bands on the
hindtarsus has also been treated in past de-
scriptions and these descriptions agreed with
the findings presented here, though none
quantified either the proportion of Cx. peus
with HT-5 all white or the width of the light
hindtarsal bands. Some (Freeborn 1926,
Ripstein 1935, Carpenter and LaCasse 1955,
Bram 1967, and McDonald et al. 1973) de-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
scribed the white scales on the palpi of fe-
male adult Cx. stigmatosoma, contrasting
them with the lack of white scales on the
palpi of Cx. peus. Significantly, McDonald
et al. (1973) made this distinction between
the species in the state where the holotype
of Cx. peus was collected, lending support
to the importance of this character in the
type locality.
Geographic distribution of the species in
Arizona and Utah supports the nomencla-
torial changes made in this paper. Culex
stigmatosoma is restricted to the southern
part of Arizona in Yuma, Pima, Pinal, Santa
Cruz, and Cochise counties (McDonald et
al. 1973), well south of the type locality of
Cx. peus in Oak Creek Canyon, Coconino
County. Records of Cx. stigmatosoma in
Utah (Dyar 1928) were apparently false, as
the species has never been collected in the
state despite extensive collecting (L. T. Niel-
sen, personal communication) and Dyar’s
original specimens are lost. Culex peus, on
the other hand, occurs throughout much of
Arizona, extending north through Coconino
County (McDonald et al. 1973) all the way
to Washington County in southern Utah
(Nielsen and Linam 1963).
ACKNOWLEDGMENTS
The University of Arizona and the Uni-
versity of Kansas for lending specimens;
Smithsonian Archives for searching for H.
G. Dyar’s notes and letters; M. Sanderson
for visiting the type locality of Cx. peus; E.
L. Peyton, B. A. Harrison, and L. T. Nielsen
for helpful comments on taxonomy; R. A.
Ward and R. C. Wilkerson for helpful com-
ments on the manuscript; and Taina Litwak
for preparing the figures. Opinions and as-
sertions contained herein are the private
views of the author and are not to be con-
strued as official, nor as reflecting the views
of the supporting agencies.
LITERATURE CITED
Adams, C. F. 1903. Dipterological contributions.
Kansas University Science Bulletin 2: 21-47.
VOLUME 90, NUMBER 4
Aguilar, S. G. 1931. Claves para identificar mosqui-
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PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 495-500
WORKER SIZE AND PIRACY IN FORAGING ANTS
J. F. CARROLL
Livestock Insects Laboratory, Agricultural Environmental Quality Institute, Agricul-
tural Research Service, U.S. Department of Agriculture, Rm. 120, Bldg. 307, BARC-East,
Beltsville, Maryland 20705.
Abstract. — Foraging ants (1.8 to 12 mm long) vied for small (<3.2 mg, <1.5 mm) cheese
baits of two sizes. Small ants succeeded in gathering only the smaller baits. Although small
ants found the larger baits first ca. 50% of the time, they always lost the large baits to
solitary foragers of larger species. Small baits were generally gathered by the first species
to find them with the smallest (<2 mm long) ants successful in ca. 80% of the cases.
Recruitment was of no consequence in the success of these encounters. Control of baits
was exchanged only to species of ants of equal, or, more often, larger size. The largest
species, Camponotus pennsylvanicus, never foraged the small baits.
The niche of the small (<2 mm long) species of ants appeared to be defined in part by
consistently unsuccessful confrontations with individual foragers of larger species for
discrete food particles of a certain size range (i.e. too large or cumbersome for a solitary
forager of a small species to carry easily, but easily carried by individuals of a larger
species). The pirating of food items by solitary foragers may be an important part of the
foraging repertoire of many medium-sized and large ants.
Key Words:
Paratrechina
Ants exploit a variety of food resources
(e.g. honeydew, seeds, and living and dead
invertebrates) and, according to Carroll and
Janzen (1973), forage primarily for partic-
ulate and widely scattered food items. If
small enough, items are garnered by solitary
foragers, while recruitment is important in
foraging for items too large or cumbersome
for an individual ant to carry by itself. The
size of food items taken by various insect
species has been found to be related to the
overall size of the insect or the dimensions
of its food gathering organs (Hespenheide
1973, Wilson 1975). Such size-match rela-
tionships are known among the Formicidae
(Davidson 1977a, b, Bernstein 1979, Wil-
son 1978), but may not be universal in the
family (Rissing and Pollock 1984).
baits, Camponotus, Aphaenogaster, Myrmica, Leptothorax, Tapinoma,
Larger ants can take food particles of a
greater size range than small species, re-
sulting in overlap of dietary resources (Chew
and DeVita 1980). Direct interspecific com-
petition by foragers for large baits has been
well documented (e.g. Levins et al. 1973).
Field observations of single foragers of large
ant species wresting food items (primarily
dead arthropods) from groups of several
workers of small species prompted this in-
vestigation of the frequency of this sort of
competition.
MATERIALS AND METHODS
Observations were made at two sets of
bait stations at Beltsville, Maryland. One
set of 10 bait observation sites was on an
infrequently used sandy road through an
496
upland Virginia pine, Pinus virginiana, and
mixed xerophilous oak woods, while the
second set of 10 stations was along a path
through a more mesic woods of mixed hard-
woods (mostly oaks). Mosses, grasses, herbs
and sapling trees and shrubs grew between
the wheel ruts, but there was little litter on
the sandy road. The path was more shaded
and leaf litter plentiful. Bait stations were a
minimum of 10 m apart. Studies were con-
ducted May-September, 1200 to 1800 h
EDT.
Cheese cubes of two sizes (X = 3.2 + 0.21
mg [n = 10] ca. 1 x 1 x 1 mm and x =
0.52 + 0.20 mg [n = 10], ca. 0.5 x 0.5 x
.50 mm) served as baits. At each station a
large cube was dropped without respect to
locations of foraging ants, but so that I could
observe it. I observed the ensuing bait-re-
lated ant activity until the bait was carried
into an ant nest. This procedure was re-
peated, using the smaller baits. Entrances of
some ant nests were hidden beneath leaf
litter. In such cases, I waited ca. 3 min after
an ant with a bait disappeared in the litter,
and then I brushed away the litter to find
the nest entrance. The distances ants carried
baits were measured. I recorded the species
of ants involved in the fates of the cubes
and the type and sequence of the activities.
These bait drops were made in June and
July between 1215 EDT and 1715 EDT with
temperatures of 24 to 33°C.
Additional random drops, elsewhere along
the road and in the woods brought the total
number of bait stations to 31 for small baits
and 29 for large baits on the road and 20
for small baits and 32 for large baits in the
woods. To further verify the patterns of for-
aging success observed with the bait drops,
additional baits were placed in the paths of
foragers of species more commonly in-
volved in the random drops. All the addi-
tional drops were between 1215 and 1800
EDT at 24-33°C, but over a longer period,
May-September. While ants may exhibit
species-specific patterns in their daily for-
aging periods, the cast of characters, ob-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
served during the daily and late May-early
September time frames of this study, never-
theless remained remarkably constant.
Prenolepis imparis (Say), a dominant species
in cooler seasons was commonly seen at the
sites during those times, but was never in-
volved in the observations reported here.
The ant species were classified according
to size as Class I (<4 mm long), II (>4 and
<8 mm), III (>8 mm). The ants were mea-
sured in an extended position from the frons
to the tip of the gaster. Samples of each
species were collected for identification. The
species composition of the ant fauna of the
wooded and road sites was similar with the
pertinent exceptions that Aphaenogaster
treatae (Forel) was strictly limited to the
road, and the 4. rudis (Emery) to a lesser
degree to the woods, and that the Class III
Camponotus pennsylvanicus (DeGeer) was
also more prevalent in the woods.
The frequencies with which ant species
were first to find baits and frequencies of
successfully removing baits were analyzed
by Chi square contingency tables. Ant spec-
imens were identified by D. R. Smith, the
U.S. Department of Agriculture Systematic
Entomology Laboratory, U.S. National
Museum of Natural History, Washington,
Dic:
RESULTS
Small baits both on the road and the path
were never found first by the largest ants
(Class III), whereas, the smallest species
(Class I) were first to find both large and
small baits significantly more often than the
other size classes (P < 0.05) (Table 1). There
was no significant difference (P > 0.05) in
the frequency with which Classes II-III
found large or small baits (Table 2). On only
2 of 20 occasions Class III species were the
first to find the large baits.
Although Class I species were first at baits
for >50% of random drops, they never suc-
ceeded in gathering a larger bait, nor did
they remove fragments visible to the naked
eye from the larger baits. In every instance
VOLUME 90, NUMBER 4
Table 1.
to foraging ants.
Subfamily Species Size Class
Myrmicinae
Myrmica pinetorum (Wheeler) II
M. emeryana (Forel) II
Aphaenogaster rudis (Emery) II
A. treatae (Forel) II
A. sp. A Il
Pheidole bicarinata vinelandica (Forel) I
P. pilifera (Roger) I
Leptothorax curvispinosus Mayr I
Dolichoderinae
Tapinoma sessile (Say) I
Formicinae
Paratrechina parvula (Mayr) I
Lasius alienus (Foerster) I
Camponotus pennsylvanicus (DeGeer) Ill
Formica pallidefulva nitidiventris Emery II
F. subsericea Say Il
497
The size classes of ant species which were first to find cheese baits randomly dropped with respect
First to Find
Small Bait Large Bait
Road Woods Total Road Woods Total
2 1 3 2 1 3
6 5 11 fl 8 15
2 0 2 1 1 2
0 0 0 a 0 4
(0) 1 l 1 2 3
1 0 1 0) 0 0
1 0 1 0 0 0
3 2 5 2 3 5
3} 1 4 1 2
12 9 21 10 11 21
1 1 2 1 0 1
0 0 0 0 2 2
0 0 0) 0 1 1
o 0 0 a
31 20 51 29 32 61
* Small bait ca. 0.5 mg, ca. 0.5 mm}; large bait ca. 3.2 mg, ca. 1.5 mm’.
> Class I <4 mm long, Class II >4 and <8 mm long, Class III >8 mm long.
solitary foragers of larger species pirated the
baits from Class I ants, even when as many
as 5 of the smaller ants were present. Re-
cruitment of co-workers was of no conse-
quence in the final outcomes of these trials.
Occasionally a Class II forager relinquished
a large bait to a solitary large forager of
another species.
On the other hand, the ant to initially
discover a small bait, usually successfully
carried the bait to its nest. Class I species
Paratrechina parvula (Mayr) successfully
foraged all nine small baits it was first to
find on the road and 7 of 8 of the small baits
in the woods, while Lepotothorax curvispi-
nosus Mayr garnered | of 2 small baits on
the road and 2 of 3 in the woods. Tapinoma
sessile (Say) relinquished the only small bait
it found to P. parvula. Class I species were
always successful in garnering the smaller
baits when they were first to find them. For
baits on the sandy road or on leaf litter along
the path, the foraging success results were
similar. Myrmica emeryana (Forel) (Class
II) foragers were first to the large baits in
just 3 of 20 random drops, but garnered
them 10 times.
Class I species P. parvula, L. curvispino-
sus, and T. sessile relinquished large baits
when they were first to find them in all of
20, 5 and 2 instances respectively. Whereas
Class II species M. emeryana, Aphaenogas-
ter treatae and A. rudis successfully foraged
large baits on 7, 4 and 4 instances respec-
tively. In about half the observations, three
or four species of ants were involved in the
fate of large baits (Table 3). Typically, when
a forager of a small species (e.g. P. melan-
deri) was the first ant to find a large bait, it
palpated and tugged at the bait. Unable to
move the bait, the ant would repeatedly
move | to 3 cm from the bait, only to return
in a few seconds and repeat the palpating
and tugging. In a few instances, the ant would
498
Table 2. Fates of two sizes of cheese baits foraged
by three size-defined class worker ants of various species.
No. Times No. Times
Bait Size Class First to Successfully
Size* Stations” of Ants Find Bait Foraged Bait
Small Road I 7A 7A
II 3A 3A
Ill OB OB
Woods I 6A 6A
II 4A 4A
Ill OB OB
Road and I 3A 13A
Woods II 7A 7A
Ill OB OB
Large Road I 6A OA
Il 4A 9B
Ill OB 1A
Woods I 4A OA
II SA 8B
Ill 1B 2A
Road and I OA OA
Woods Ul 9A 17B
Ill 1B 3A
«Small bait ca. 0.5 mg, ca. 0.5 mm*, large bait ca.
3.2 mg, ca. 1.5 mm?.
» Ten bait drops each on road and in woods.
© Class I <4 mm long, Class I] >4 mm and <8 mm
long, Class III >8 mm long.
“Numbers in the same column, pertaining to the
source bait size and station, and followed by the same
letter are not significantly different (Chi square contin-
gency tables, P < 0.05).
leave and not return and fewer still were
instances of recruitment of nestmates. How-
ever, a forager of one of the larger species
generally arrived while the small forager was
alone at the bait. The larger ant wandered
to within about | cm of the bait before it
turned abruptly toward the bait. The inter-
loper often seized the bait, or when neces-
sary, tore it from the grip of the smaller ant,
and carried it directly nestward. Other times
the larger ant nipped the smaller one, par-
ticularly if the latter accidentally or aggres-
sively interfered with the larger ant seizing
the bait. Such brief attacks drove away the
smaller species even when two or three of
them were at the bait; nor did the smaller
ants pursue the interloper as it carried the
cube nestward.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 3. Sequence of control of large baits by ant
species and speed and distance baits were carried to
ants’ nests.*
Time
Distance Bait Speed
Bait Car- Bait
Sequence of Species Carned med Carned
Station Controlling Bait” (cm)© — (min)* (cm/min)<
Road PP, MP, PRAD 91958:0 11.4
Road PP, US, PP;TS; 127 5:0 25.4
PPS, ME
Road AT 36 «(1.0 35.6
Road ME 122 4.0 30.5
Road TS, MP, TS, LC, 61 1.5 40.7
TS, MP, MP, AT
Road AT 229) 220.0 wll4'3
Road ME 168 8.0 21.0
Road LC, MP, ME 86 3.0 28.8
Road PP, ME 122 7.0 17.4
Road PP, AT 130 0.8 155.4
Woods CP,* PP, ME 196 4.5 43.3
Woods’. KP:¢ EC).cPe L128) 16:55 sall?3s5
Woods LC, ME O71 es 64.4
Woods LC, PP,s ME Tulyer2eS 28.5
Woods PP, ME 48 3.0 16.1
Woods PP, AR 46 1.0 45.7
Woods AS 30,5:75 D3
Woods FS, ME, CP 549 1.25 438.9
Woods AR, CP,; AR 104 1.5 69.4
Woods ME DO 2225 46.7
* Baits ca. 3.2 mg, ca. 1.5 mm’.
» Abbreviations for ant species: AR = Aphaenogaster
rudis, AS = A. sp., AT = A. treatae, CC = Camponotus
castaneus, CP = C. pennsylvanicus, FP = Formica p.
nitidiventris, FS = F. subsericea, LC = Leptothorax
curvispinosus, ME = Myrmica emeryana, MP = M.
pinetorum, TS = Tapinoma sessile, PP = Paratrechina
parvula.
© By the last ant that took control of the bait and
brought the bait to its nest.
4 Abandoned bait.
© Attacked ant at bait, but did not gain control.
DISCUSSION
The 0.5-3.2 mg, 0.5-—1.0 mm range ap-
peared to be the lower limit of food particle
size at which relative forager size/strength
operated as a significant factor in the out-
come of competitive foraging for this as-
semblage of ant species and type of food.
Success in foraging for items smaller than
this threshold size appeared to be more de-
pendent on a forager’s ability to find a food
item before competitors. Unwieldiness of
food items (arthropods with appendages in-
VOLUME 90, NUMBER 4
tact) probably effectively creates the same
sort of strength related barrier to foraging
ants. Studies with larger baits (e.g. Levins
et al. 1973, Lynch et al. 1980) suggest that
there is a food item size threshold above
which the largest species must resort to re-
cruitment to efficiently exploit the item (Os-
ter and Wilson 1978). Such large food items
are therefore subject to multi-species use
before one colony can dominate or later
through pilfering by small species.
For large food items multi-species use 1s
likely and small items are more apt to be
gathered by individual small ants rather than
very large foragers. However, there may not
be a smooth ant size/bait size usage gradient
between the extremes, because small ants
may derive little or no food material from
particles that are too large for their individ-
ual foragers to carry easily, yet which are
readily pirated by solitary foragers of com-
mon large species.
In terms of energetics, it seems inefficient
for a small ant to consistently compete un-
successfully for a resource (1.e. food items
like the large cheese baits). One explanation
may be that the minimal quantity a worker
manages to remove in her infrabuccal pock-
et before the food item is lost isa worthwhile
payload. Also, food items of the dimensions
of the large cheese baits may be scarce in
natural conditions and thereby represent an
abnormal situation. However, baits of both
sizes did approximate the sizes of many
small invertebrates, which might die of a
variety of causes (e.g. drowned by a down
pour). Certainly the relative size distribu-
tion of available foods is of utmost impor-
tance in the natural environment (Wilson
1975).
The pirating of food items from rather
ubiquitous smaller ants by solitary foragers
may be an important behavior in some larg-
er species (e.g. Aphaenogaster spp., Myr-
mica emeryana). In this study, A. treatae
successfully foraged more large baits (6) by
seizing them from smaller ants, than by
being the first to find them (4), while /.
499
emeryana pirated 8 large baits and garnered
12 which they found first. Foragers of reg-
ularly interloping species may have been
aided in detecting baits by the activity of
the smaller ants already at the baits.
According to Oster and Wilson (1978),
the main disadvantage to reliance on re-
cruitment is the time it consumes. In this
study recruitment occurred infrequently, and
with no more than 3 to 5 workers of P.
parvula at a bait at one time. Recruitment
rates have been related to food patch size
and sucrose content (Taylor 1977). Perhaps
the size or content of the cheese baits used
in this study were not attractive enough to
elicit strong recruitment, although the small
baits were gathered by the first species to
find them. Lynch et al. (1980) reported that
P. melanderi (Wheeler) showed greater re-
cruitment to sugar baits. In the case of food
items similar in size and attractiveness to
large baits, it might be inefficient for small
ant species to recruit and mobilize several
workers only to lose virtually the entire food
items to unrelated ants.
Based on their bait studies, Lynch et al.
(1980) considered P. melanderi and L.
curvispinosus as timid ants and Prenolepis
imparis, an aggressive dominant speices. P.
imparis was common at the site of this study,
but since most of the trials were conducted
in mid-summer, this species, which is more
active in cooler weather, was not actively
involved. According to Lynch et al. (1980),
a single P. imparis worker can hold its own
at a bait against the large 4. rudis. No small-
ish ant seemed to fit this role in this study.
A detailed investigation of paired interspe-
cific interactions for control of small baits
might explain competitive relationships in
assemblages of ant species.
ACKNOWLEDGMENT
I thank Dr. R. Smith of the Systematic
Entomology Laboratory, Biosystematics and
Beneficial Insects Institute, U.S. Dept. of
Agriculture, U.S. National Museum of Nat-
500
ural History, Washington, D.C. for identi-
fying some of the ants involved in this study.
LITERATURE CITED
Bernstein, R. A. 1979. Evolution of niche breadth in
populations of ants. Amer. Natur. 114: 533-544.
Carroll, C. R. and D. H. Janzen. 1973. Ecology of
foraging by ants. Ann. Rev. Ecol. Sept. 4: 231-
DOT
Chew, R. M. and J. DeVita. 1980. Foraging char-
acteristics of a desert ant assemblage: Functional
morphology and species separation. J. Arid En-
viron. 3: 75-83.
Davidson, D. W. 1977a. Species diversity and com-
munity organization in desert seed-eating ants.
Ecology 58: 711-724.
1977b. Foraging ecology and community or-
ganization in desert seed-eating ants. Ecology 58:
725-737.
Hespenheide, H. A. 1973. Ecological inferences from
morphological data. Ann. Rev. Ecol. Syst. 4: 213-
229.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Levins, R., M. L. Pressick, and H. Heatwole. 1973.
Coexistence patterns in insular ants. Amer. Sci.
61: 463-472.
Lynch, J. F., E. C. Balinsky, and S. G. Vail. 1980.
Foraging patterns in three sympatric forest ant
species, Prenolepis imparis, Paratrechina melan-
deri and Aphaenogaster rudis (Hymenoptera: For-
micidae). Ecol. Entomol. 5: 353-371.
Oster, G. F. and E.O. Wilson. 1978. Caste and Ecol-
ogy in the Social Insects. Princeton University
Press, Princeton, N.J. 352 pp.
Rissing, S. W. and G. B. Pollock. 1984. Worker size
variability and foraging efficiency in Veromessor
pergandei (Hymenoptera: Formicidae). Behav.
Ecol. Sociobiol. 15: 121-126.
Taylor, F. 1977. Foraging behavior of ants—Exper-
iments with two species of myrmecine ants. Be-
hav. Ecol. Sociobiol. 2: 146-167.
Wilson, D. S. 1975. The adequacy of body size as a
niche difference. Amer Natur. 109: 769-784.
Wilson, E.O. 1978. Division of labor based on phys-
ical castes in fire ants (Hymenoptera: Formicidae:
Solenopsis). J. Kansas Entomol. Soc. 51: 615-636.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 501-507
HOST SPECIFICITY OF ADULT EUSTENOPUS HIRTUS (WALTL)
(COLEOPTERA: CURCULIONIDAE), A POTENTIAL
BIOLOGICAL CONTROL AGENT OF YELLOW STARTHISTLE,
CENTAUREA SOLSTITIALIS L. (ASTERACEAE, CARDUEAE)
S. L. CLEMENT, T. Mimmoccuti, R. SOBHIAN, AND P. H. DUNN
USDA, ARS Biological Control of Weeds Laboratory, Europe. % U.S. Embassy, APO
New York, New York 09794-0007; (SLC present address) USDA, ARS Plant Introduction
Station, 59 Johnson Hall, Washington State University, Pullman, Washington 99164-
6402; (TM present address) Via Baldo Degli Ubaldi 59, 00167 Rome, Italy.
Abstract.—Host specificity of the flower head weevil Eustenopus hirtus (Waltl) was
investigated in Italy and supported its potential and safety as a biological control agent
for yellow starthistle, Centaurea solstitialis L., in the United States. Adults damaged yellow
starthistle by feeding on young buds and larvae consumed developing seeds. In the lab-
oratory, adults fed and damaged capitula of several test plants but females oviposited
only on C. solstitialis and two congeneric species. Females experienced little or no oocyte
development when confined to plants other than Centaurea.
Key Words:
Yellow starthistle, Centaurea solstitialis
L. (Asteraceae, Cardueae), 1s a winter an-
nual or biennial plant of Eurasian origin that
has become a major threat to rangelands in
the western United States (Callihan et al.
1982, Maddox et al. 1985, Maddox and
Mayfield 1985, Roché et al. 1986). In the
late 1950°s, weed biocontrol workers began
surveying southern Europe to find potential
biological control agents for yellow star-
thistle and other weedy Centaurea (Zwolfer
1965, Zwolfer et al. 1971, Sobhian and
Zwolfer 1985, Clement and Mimmocchi
1988). To date, two natural enemies of C.
solstitialis in Europe have been found safe
for introduction into the U.S. but only one
of these, the flower head weevil Bangaster-
nus orientalis (Capiomont) (Coleoptera:
Curculionidae), has become established on
the plant in the U.S. (Maddox et al. 1986).
One of the first insects we considered as
a new biocontrol agent to supplement the
weevil, bud-feeding, seed-feeding
action of B. orientalis was Eustenopus hirtus
(Waltl) (Coleoptera: Curculionidae), a ca-
pitulum-infesting weevil that Sobhian and
Zwolfer (1985) indicated (cited as E. ab-
breviatus Faust) was probably restricted to
C. solstitialis in Greece. These workers also
reported that hibernating adults of this uni-
voltine weevil become active in late May or
June in northern Greece and lay eggs in well-
developed buds of C. solstitialis. Moreover,
they reported that a single larva destroys
almost all of the achenes in a small head.
Near Thermi, Greece, we have observed
beetles feed on small, young buds (Bu stages
1-2), copulate on mid-size, older buds (Bu
3-4), and lay eggs in the largest and most
mature closed buds (Bu 4) of C. solstitialis
(see Maddox 1981 for description of floral
bud [Bu] stages).
Csiki (1934) recognized a number of sub-
genera in the weevil genus Larinus of the
subfamily Cleoninae, tribe Lixini, including
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
and withered bud
the subgenus Eustenopus. However, Ter-
Minasyan (1978) and other workers (M. L.
Cox, pers. comm.) recognized Eustenopus
as a valid genus. In addition to E. villosus
(Boheman) (= FE. hirtus), Ter-Minasyan
(1978) listed two other species, F. /anugi-
nosus Faust and EF. abbreviatus.
Eustenopus hirtus 1s about 4-7 mm long
with white longitudinal stripes on its elytra
and is covered with long, erect hair-like se-
tae (Fig. 1). Itis recorded from Greece, Tur-
key, the Caucasus, Syria, and Iran (Ter-
Minasyan 1978, Sobhian and Zwolfer 1985).
In the literature, only C. solstitialis is re-
corded as a host plant of FE. hirtus (Sobhian
and Zwolfer 1985). The insect is not re-
corded from any crop plant in Europe, the
Middle East, or western Asia (Review of
Applied Entomology [Series A, 1913-1986],
Zoological Record [1950-1971], Grandi
1951, Hoffman 1954, Bonnemaison 1962,
Balachowsky 1963, Scherf 1964, Avidov and
Kotter 1966, Ter-Minasyan 1978, Fremuth
1982, Petney and Zwolfer 1985). Ter-Min-
asyan (1978) reported that FE. /anuginosus
was “found on” Cousinia (Compositae) in
Kazakhstan, USSR.
Adult Eustenopus hirtus and the yellow starthistle bud (Bu 2) it fed upon. Arrow indicates the damaged
After we found E. hirtus on C. solstitialis
in areas of Greece and Turkey with climates
similar to North American sites infested with
the weed, we initiated laboratory studies at
the USDA-ARS Biological Control of Weeds
Laboratory-Europe (BCWLE) in Rome, It-
aly, to learn more about the adult repro-
ductive behavior and host range of this
species. The results of these investigations
are presented in this paper.
MATERIALS AND METHODS
Test plants and insects.—Plants taxo-
nomically related to Centaurea solstitialis
(family Asteraceae; tribes Heliantheae, Car-
dueae [Cynareae], and Cichorieae) were used
in host-specificity tests, and were chosen
with 4 features in mind: (1) related weedy
species—Onopordum acanthium L.; Car-
thamus lanatus L.; Carthamus dendatus
(Forskal) Vahl; Cirsium arvense (L.) Scop.;
Centaurea nicaeensis All; Centaurea diffusa
Lam.; Centaurea cineraria L.; Cnicus bene-
dictus L.; Galactites tomentosa Moench; Ci-
chorium intybus L.; Scolymus hispanicus L.;
(2) related crop plants—Carthamus tincto-
rius L., ‘Hartman’ safflower; Cynara scoly-
VOLUME 90, NUMBER 4
mus L., “Green Globe’ artichoke; Helian-
thus annuus L., ‘Parendovik’ common
sunflower; Lactuca sativa L., ‘Bibb’ lettuce;
(3) U.S. native plants in the Cardueae— Cir-
sium undulatum (Nutt.) Spreng.; Cirsium
douglasti DC.; Centaurea americana Nutt.;
and (4) Palaearctic and Nearctic popula-
tions of the target weed—Centaurea solsti-
tialis L. grown from seed collected in Ther-
mi, Greece, Rome, Italy (control plants),
and Walla Walla and Yakima, Washington
(U.S.). Since breeding hosts of species closely
related to E. hirtus are unknown, we did not
consider this factor in the selection of test
plant species.
Entomologists (USDA, ARS) in Albany,
California provided seed of U.S. Cirsium
spp., Centaurea americana, Carthamus
tinctorius, the Washington forms of Cen-
taurea solstitialis, and rootstock of Cynara
scolymus. Other test plant species were
grown from young plants field collected in
Italy and Greece and from seed obtained in
the wild or from European botanical gar-
dens. Whenever possible, test plants were
allowed to flower and herbarium specimens
were deposited in the collection of the
BCWLE.
Reproductively active beetles were hand-
collected from C. solstitialis in northern
Greece, near the village of Doirani and on
the southern outskirts of Thessaloniki, in
June 1985 and 1986. In 1985 and 1986, 275
and 407 beetles respectively survived the
air shipments to Rome and were allowed to
feed on closed C. solstitialis buds (Bu 1-4)
for at least 48 hours before they were se-
lected for host-specificity tests. Groups of
teneral beetles, reared from C. solstitialis
capitula collected in central Greece (Xini-
ada) in August 1984 and northern Greece
(Thermi) in July and August 1985, were al-
lowed to overwinter in cages at the BCWLE
so pre-reproductive beetles would be avail-
able for feeding, mating, and oviposition
behavior studies in spring 1985 and adult
feeding and specificity studies in May 1986.
No external morphological differences were
503
found between the sexes, so males and fe-
males were selected from mating pairs in
holding containers.
Feeding and reproductive behavior.—
Adult feeding and reproductive behavior
were investigated by offering beetles a pro-
gression of C. solstitialis growth stages as
they would normally appear in the field in
northern Greece. Eight unsexed beetles (4
beetles and one plant per covered 500 cm?
cardboard carton) were first exposed to ro-
settes from March 5—April 28, then to bolt-
ing plants from April 29-May 7. Twice a
week, these plants (roots held in water-filled
vials plugged with cotton) were replaced with
fresh ones from a garden at the BCWLE.
On May 8, the eight beetles were placed in
a cage (clear plastic cylinder [diameter 20
cm: length 70 cm] with nylon organdy cov-
er) which enclosed a potted plant with Bu
1 buds, and on May 22 these heavily feeding
beetles were transferred to another plant with
Bu | buds. By June 4, this plant supported
all closed bud stages (Bu 1-4) and flowering
buds. Cartons and cages were observed at
least once but usually several times a day
to record beetle feeding, mating, and ovi-
position.
The reproductive rate (number of eggs laid
over time) of females was measured by plac-
ing a mating pair of beetles in each of five
cages (500 cm?’ carton) for 18 days and
counting the number of eggs laid per female
every three days when the C. solstitialis buds
were replaced with fresh ones. The ovipo-
sition substrates were two Bu 2, two Bu 3,
and two Bu 4 buds, with their stems held
in a water-filled vial. If a male died during
this study, it was replaced.
Host specificity tests. — Adult feeding and
ovipositional specificity, and mortality, were
measured under “no-choice”’ (tests 1-4; one
plant species per cage) and two choice (test
5; two plant species per cage) test condi-
tions. In tests | and 2, each nylon organdy
sleeve cage (diameter 14-20 cm; length 30-
42 cm) contained 2-9 beetles (1-4 females)
and branches of mature buds of one test
504
plant species. Each plant species was rep-
resented by 1-15 potted plants. Data were
recorded for each plant as soon as it started
to flower (after 3-11 days). Plants tested are
listed in Table 1. In test 3, each cardboard
cage (covered 500 cm? cartons) contained a
mating pair of beetles and Centaurea sol-
stitialis (Rome population) buds (one each
Bu 2, 3, and 4) or one closed bud (diameter
10-15 mm) of Carthamus tinctorius. There
were 20 cartons per plant species. Every
three days during this 15 day test, buds
(stems held in water-filled vials) were re-
placed with fresh ones from potted, green-
house-grown plants. If a male died during
this test, it was replaced with a new one. In
contrast to tests 1-3, pre-reproductive bee-
tles were used in test 4. For this test, two
unsexed beetles were confined to each nylon
sleeve cage for 15 days. Each cage enclosed
branches of closed buds of one test plant
species (Centaurea solstitialis [Rome pop-
ulation], Centaurea nicaeensis, Centaurea
cineraria, Galactites tomentosa, or Onopor-
dum acanthium). There were five potted
plants of Centaurea solstitialis and three of
each of the other plant species. In test 5,
beetles (two males and two females per
sleeve cage) were allowed to choose between
buds of Centaurea solstitialis (Therm1,
Greece population) and buds of another
plant species for nine days. Branches from
each potted plant supported all degrees of
bud development. There were five replica-
tions of each of the four Centaurea solsti-
tialis-test plant combinations listed in Ta-
ble 2.
At the end of the tests, or every three days
in test 3, beetle mortality was recorded, and
buds were examined for feeding damage and
eggs. Feeding damage was classified in four
ways: (—), no feeding or very slight nibbling
on buds; (+), light to moderate feeding, some
buds with two or more feeding punctures;
(++), moderate to heavy feeding, less than
¥; of buds riddled with feeding punctures;
and (+++), heavy feeding, more than 3 of
buds riddled with punctures. All dissected
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
females in tests 2 (n = 50) and 3 (n = 20)
were examined to determine egg maturation
associated with each test plant.
All studies were conducted in the BCWLE
quarantine greenhouse under temperatures
of 16-33°C, 35-85% RH, and natural day-
light. Twist-ems® were used to close the
ends of nylon sleeve cages in tests 1, 2, 4,
and 5, and to bundle branches from separate
potted plants in test 5.
RESULTS
Feeding and reproductive behavior.—
When beetles were offered a progression of
C. solstitialis growth stages between March
5 and June 10, no feeding of any conse-
quence took place until beetles were ex-
posed to Bu | buds on May 8. The beetles
continued their heavy feeding on all closed
bud stages (Bu 1-4) until the end of the
study on June 10. From June 4-10, mating
occurred only on Bu 3 and 4 buds, and ovi-
position occurred only on Bu 4 buds. Beetles
did not feed on or oviposit in flowering buds.
The mean (+SE) number of eggs laid by
five females during six consecutive three-
day periods was 2.83 + 0.52, 3.67 + 0.61,
3.67 + 0:67, 3.17 2 0.52; and 3.671013)
(F = 0.59; df = 4, 25; means are not sig-
nificantly different, P > 0.05). The five fe-
males died on July 20 (n = 2) and 26 and
August 3 and 13 after laying 18, 23, 28, 34,
and 60 eggs, respectively.
Host specificity tests.—Although adult
feeding was moderate to heavy on most
plants in tests 1 and 2, beetles oviposited
only in Bu 4 buds of Centaurea solstitialis,
and mature closed buds of Centaurea ni-
caeensis and Centaurea diffusa (Table 1).
Only two larvae were found in Centaurea
diffusa buds and these died before molting
to second instar. Beetle mortality was sig-
nificant (60-100%) on all plant species ex-
cept Centaurea solstitialis, Centaurea ni-
caeensis, Centaurea americana, and Cirsium
douglasii (Table 1). In test 2, dissections of
50 females revealed oocyte development,
albeit rudimentary, in only two females, one
VOLUME 90, NUMBER 4
505
Table 1. Synopsis of host specificity screening of Eustenopus hirtus adults allowed contact with only one
plant species (tests | & 2), June-July, 1985-1986, Rome, Italy.
Total No. of
Closed
and No. Found No. Days Beetle
Flower- Amount in Buds Beetles Mortality
Test ing of Bud Confined (%) Dunng
Plant Species No. Plants Buds Beetles! Feeding? Eggs Larvae to Plants Test
Centaurea solstitialis 1&2 12 105 36(18) +++ 54 14 6-10 16.67
Greece
Centaurea solstitialis 1 3 18 6 (3) +++ 3 4 10-11 33:5
Washington State, US
Centaurea nicaeensis 1&2 8 713. 26 (13) ++ 12 12 4-10 11.54
Centaurea diffusa 1 > 203" 10/(5) aa as OQ 2 (dead) 7-8 60.0
Centaurea americana 2 5 23. + 20(10) +4++ 0 0 9-10 0.0
Carthamus tinctorius Wee 2 eS 72 56 (28) ++ 0 0 3-10 76.79
Carthamus lanatus 1 5 57 19 (8) + 0 0) 4-7 100.0
Carthamus dentatus 1 5 43 22 (12) = 0 0 4-6 100.0
Cynara scolymus 1 1 1 9 (4) + 0 0 5 100.0
Helianthus annuus 1 3 10 6 (3) — 0 0 47 100.0
Lactuca sativa I 3 «141 6 (3) 4F 0 0 3 100.0
Cnicus benedictus 1 5 21 10 (5) Te anar 0 0 8 80.0
Scolymus hispanicus 2 5) 56 20 (10) +r 0 0 5-6 100.0
Cirsium arvense 1 5 77 ~=—:10 (5) ++ 0 0 +8 60.0
Cirstum undulatum 1 2 15 12 (6) ++ 0 0 4 100.0
Cirsium douglasii 1 l 7h 6 (3) +44 0 0 5 16.7
' Numbers of females in parentheses.
>(—), no feeding or very slight nibbling; (+), light to moderate feeding; (+ +), moderate to heavy feeding;
(+++), heavy feeding (see text for more details).
each from a Carthamus tinctorius and a
Scolymus hispanicus plant.
In test 3, none of the closed buds of Car-
thamus tinctorius were accepted for ovi-
position, but they did suffer light to mod-
erate feeding damage. However, feeding was
insufficient for normal oogenesis because no
Table 2.
7, 1986, Rome, Italy.
eggs were found in the ovarioles of the 20
females examined. Moderate to heavy feed-
ing occurred on Centaurea solstitialis, and
the 18 females that lived to the end of the
15 day test laid an average of 12.83 (+0.74
SE) eggs. The 20 females confined to Cen-
taurea solstitialis had an average of 0.90
Feeding and oviposition of Eustenopus hirtus adults under two choice test conditions, June 29-July
Degree of Bud Feeding on!
Plant combinations
C. solstitialis
Number of Eggs and Larvae Found in Buds of
Other Plant C. solstitialis Other Plant
Centaurea solstitialis, Greece vs. +++
Carthamus tinctorius
Centaurea solstitialis, Greece vs. +++
Cirsium arvense
Centaurea solstitialis, Greece vs. trae
Cichorium intybus
Centaurea solstitialis, Greece vs. Seeiets
Centaurea nicaeensis
35 eggs; 5 larvae
+ 0
31 eggs; 4 larvae
= 0
21 eggs; 3 larvae
= 0
29 eggs; 5 larvae
35 10 eggs
'(—), no feeding or very slight nibbling; (+), light to moderate feeding; (+ +), moderate to heavy feeding;
(+++), heavy feeding (see text for more details).
506
(+0.14 SE; range 0-2) mature eggs in their
ovarioles. Beetle mortality during the test
was 42.5% in the Carthamus tinctorius car-
tons and 17.5% in the Centaurea solstitialis
cartons.
In test 4, feeding was heavy on Centaurea
solstitialis, moderate on C. nicaeensis and
negligible on Centaurea cineraria and Ga-
lactites tomentosa. Beetles did not feed on
closed buds of Onopordum acanthium and
83.3% died on this plant. Mortality was
16.6% or less on other plants. Eggs were not
found in the closed buds of any plant.
In test 5, when the beetles were given a
choice between Centaurea solstitialis and
either Centaurea nicaeensis, Cichorium in-
tybus, Carthamus tinctorius, or Cirsium ar-
vense, eggs were laid only in the mature
closed buds of the two Centaurea species
(Table 2). There was no evidence of feeding
on Cirsium arvense and Cichorium intybus,
but a few feeding punctures were observed
on closed buds of Carthamus tinctorius and
Centaurea nicaeensis. Feeding was heavy on
Centaurea solstitialis (Table 2).
DISCUSSION
Host specificity tests and field observa-
tions indicated that a strong relationship ex-
ists between E. hirtus and C. solstitialis. First,
C. solstitialis is the only known breeding
host of F. Airtus in nature. In the laboratory,
mating and oviposition only occurred on Bu
3-4 buds of C. solstitialis. Second, females
experienced little or no oocyte development
when confined to nonhost plants. Third, the
failure of females to oviposit on plants other
than C. solstitialis and two other Centaurea
species suggests that the egg-laying response
may be triggered by specific stimuli of ma-
ture closed buds of C. solstitialis and other
Centaurea. Fourth, although adult bud
feeding was moderate to heavy on several
nonhost plants in the absence of C. solsti-
tialis, it was absent or negligible when adults
were offered a choice between C. solstitialis
and a nonhost plant. Last, beetle longevity
was generally reduced when they were forced
to feed exclusively on nonhost plants.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Although we found that adults were able
to feed on some nonhost plants, and other
researchers found that a few larvae could
complete their development when placed as
neonates in safflower buds (Sobhian and
Zwolfer, 1985; Mimmocchi and Clement,
unpublished data), we still considered E.
hirtus a potential biocontrol agent. The adult
is the only mobile stage and our tests dem-
onstrated the specificity of oviposition by
females. Moreover, when beetles were ex-
posed equally to four plant species, includ-
ing safflower and artichoke, under field ex-
perimental conditions in Thermi, Greece,
the beetles fed and oviposited only on ma-
ture closed buds of C. solstitialis (Clement
and Sobhian, unpublished data). Like many
monophagous and oligophagous insects
(Force 1966), E. hirtus may have a broader
host-feeding range in artificial environ-
ments than in the field. The collective evi-
dence suggests that there 1s little or no dan-
ger of a crop like safflower being attacked
by E. hirtus as long as C. solstitialis is pres-
ent.
Eustenopus hirtus scored 40 by Goeden’s
system (1983) for rating the potential effec-
tiveness of biocontrol agents, indicating that
it could be a partially effective agent. This
monophagous beetle has been cleared for
introduction into the USDA quarantine fa-
cility in Albany, California, for additional
study.
ACKNOWLEDGMENTS
We thank L. Andres, G. Piper, C. Turner,
D. Whitehead and two anonymous review-
ers for helpful comments on the manuscript,
and M. Cristofaro and A. Laregina (BCWLE)
for technical assistance. We also thank E.
Colonnelli of the Dipartimento di Biologia
Animale e dell Uomo, Vaile dell Universita,
Rome, Italy, and M. Cox of the C.A.B. In-
ternational Institute of Entomology, Lon-
don, England, for determination of the bee-
tle. Mention of a proprietary product does
not constitute an endorsement or a rec-
ommendation for its use by the USDA.
VOLUME 90, NUMBER 4
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thistle infestations are on the increase. Calif. Agric.
39(11&12): 10-12.
Maddox, D. M., R. Sobhian, D. B. Joley, A. Mayfield,
and D. Supkoff. 1986. New biocontrol agent in-
troduced for yellow starthistle. Calif. Agric.
40(11&12): 4-5.
Petney, T. N. and H. Zwélfer. 1985. Phytophagous
insects associated with cynareae hosts (Asteraceae)
in Jordan. Israel J. Entomol. 19: 147-159.
Roche, B. F., G. L. Piper, and C. J. Talbott. 1986.
Knapweeds of Washington. Wash. State Coop. Ext.
Cir. EB1393, 41 pp.
Scherf, H. 1964. Die entwicklungsstadien der mit-
teleuropaischen Curculioniden (Morphologie,
bionomie, dkologie). Abh. Senckenberg. Naturf.
Gesellsch. 506, 335 pp.
Sobhian, R. and H. Zwélfer. 1985. Phytophagous
insect species associated with flower heads of yel-
low starthistle (Centaurea solstitialis L.) Z. ang.
Entomol. 99: 301-321.
Ter-Minasyan, M. E. 1978. Weevils of the subfamily
Cleoninae in the fauna of the USSR. Tribe Lixini.
(Transl. from Russian). Amerind Publ. Co., New
Dehli. 166 pp.
Zwolfer, H. 1965. Preliminary list of phytophagous
insects attacking wild Cynareae (Compositae) in
Europe, pp. 81-154. Commonwv. Inst. Biol. Con-
trol Tech. Bull., No. 6.
Zwolfer, H., K. E. Frick, and L. A. Andres. 1971. A
study of the host plant relationships of European
members of the genus Larinus (Col.: Curculioni-
dae), pp. 97-143. Commonw. Inst. Biol. Control
Tech. Bull., No. 14.
WANTED: Offprints or reprints of entomological articles and personal letters by the
Russian-American writer and entomologist, Vladimir Nabokov. For updating my stan-
dard bibliographic work on the author, I need to acquire or examine such papers. | will
send a list of his entomological journal appearances to anyone who asks. Contact Michael
Juliar, 355 Madison Ave., Highland Park, NJ 08904 USA, 201-846-4221.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 508-509
NOTE
A New Synonym and Revised Status in
Apterothrips (Thysanoptera: Thripidae)
Sericothrips apteris Daniel (1904, Ento-
mol. News 15: 295), synonymized under
Anaphothrips secticornis (Trybom) by Hood
(1927, Pan-Pac. Entomol. 3: 173), isa valid
species (REVISED STATUS) based on my
study of three paratypes of apteris, one para-
type of Apterothrips subreticulatus Bagnall
(1908, Trans. Nat. Hist. Soc. North. New-
castle-on-Tyne (N.S.) 3: 185) (= secticor-
nis), and an identified specimen of secticor-
nis from a type locality, Albany, Oregon. I
also conclude that apteris belongs in Apter-
othrips. The type depository of secticornis
is unknown. According to Mound and
Walker (1982, Fauna of New Zealand No.
1: 55). one of the localities mentioned in the
original description of secticornis by Try-
bom was Albany, Oregon. Sericothrips stan-
fordii Moulton (1911, U.S.D.A. Bur. Ento-
mol. Tech. Ser. No. 12, part III, p. 52) was
assigned to Anaphothrips by Moulton (1926,
Pan-Pac. Entomol. 3: 23) and later to Ap-
terothrips by zur Strassen (1973, Sencken-
bergiana Biol. 54: 142). A syntype of this
species examined in this study is identical
to apteris (NEW SYNONYMY). The types
of apteris and stanfordii were collected in
the same geographic area of California; ap-
teris at San Francisco in 1902? and stanfor-
dii at Stanford University, Palo Alto, in
1904. Only two species, apteris and secti-
cornis, are currently assigned to Apteroth-
rips.
Apterothrips apteris and secticornis are
apterous; their antennae are 8 or 9-seg-
mented with segment VI occasionally par-
tially divided; and abdominal tergites and
sternites have extensions of the posterior
margin (posteromarginal flange). The body
coloration of females varies from complete-
ly dark brown to the pterothorax or thorax
and first abdominal segment yellow with the
rest of the body brown. Antennae are com-
pletely brown or the bases of segment III
are yellowish brown or segment III and dis-
tal part of segment II are yellowish brown;
tarsi are yellow or occasionally brown, and
remainder of the legs varies from mostly
yellow to mostly brown. Body coloration of
the males is similar to those of the females;
however, secticornis males may also have
bodies that are mostly yellow with brown
head. The two species are readily differen-
tiated by the six major setae on the posterior
margin of abdominal sternites [V—VI: ap-
teris has the laterad-most setae (B3) at the
extreme side of the sternite, thus the inser-
tions of the six setae divide the postero-
marginal flange into five sections; converse-
ly, the B3 setae of secticornis are located
submarginally, thus the insertions of the six
setae divide the posteromarginal flange into
seven sections. Also, apteris usually has the
fifth dorsal seta from the middle on abdom-
inal tergites III-VII anterior to the posterior
margin and the setal insertion does not di-
vide the posteromarginal flange (occasion-
ally 6 dorsal setae may be present); whereas,
the fifth dorsal seta of secticornis 1s on the
posterior margin and the posteromarginal
flange is divided at the setal insertion.
Jacot-Guillarmod (1974, Ann. Cape Prov.
Mus. (Nat. Hist.) 7(3): 589) lists secticornis
from Russia, Europe, Canada, United States
(California, Oregon, Hawaii), Argentina,
Chile, South Georgia I., Juan Fernandez Is.,
Easter I., and New Zealand. Mound and
Houston (1987, Occ. Pap. Syst. Entomol. 4:
5) report it from Australia, Crozet I., Falk-
land Is. and Peru. Some of these records are
based apparently on misidentifications of
apteris. | have examined, in the collection
VOLUME 90, NUMBER 4
of Thysanoptera in the United States Na-
tional Museum of Natural History, secti-
cornis specimens from Europe, Canada (AI-
berta, British Columbia, Labrador) and
United States (Alaska, Colorado, Idaho,
Nevada, Oregon, Washington). The follow-
ing apteris records from Argentina, Chile,
Ecuador, Guadelupe I., Mexico, Panama,
Peru, Australia and New Zealand are based
on reexaminations of previously identified
secticornis material.
I thank W. H. Ewart, University of Cal-
ifornia, Riverside for reviewing the manu-
509
script and for additional distribution rec-
ords, and to the following reviewers for their
comments and suggestions: T. Kono, Cali-
fornia Dept. of Food and Agriculture, Sac-
ramento; R. J. Gagne and F. C. Thompson,
Systematic Entomology Laboratory, ARS,
USDA, Washington D. C. and M. B. Stoet-
zel, same laboratory, Beltsville, Maryland.
Sueo Nakahara, Systematic Entomology
Laboratory, PSI, Agricultural Research Ser-
vice, USDA, Beltsville, Maryland 20705.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 510-512
NOTE
Feeding by Medetera Species (Diptera: Dolichopodidae) on
Aphids and Eriophyid Mites on Apple,
Malus domestica (Rosaceae)
Adults and most known larvae of doli-
chopodids are predaceous on soft-bodied
arthropods (Robinson, H. and J. R. Vock-
eroth. 1981. Jn J. F. McAlpine et al., eds.,
Manual of Nearctic Diptera, Vol. 1. Agric.
Canada Monogr. No. 27. Biosystematics
Research Institute, Ottawa. 674 pp.). Sev-
eral Medetera larvae are subcortical pred-
ators living under the bark of dead and dying
trees. Immatures of Medetera aldrichii
Wheeler are major predators of scolytid lar-
vae (Schmid, J. M. 1971. Can. Entomol.
103: 848-853.), (Hopping, G. R. 1947. Can.
Entomol. 79: 150-153), (Nagel, W. P. and
T. D. Fitzgerald. 1975. Entomophaga 20:
121-127). The feeding habits of adult Med-
etera remain obscure. Bickel (1985.
U.S.D.A. Tech. Bull. 1692. 109 pp.) re-
ported Medetera petulca Wheeler (Fig. 1)
preying on arthropods such as spiders, mites,
small centipedes, Collembola, Diptera
(Sciaridae, Psychodidae, and Cecidomyi-
idae), Homoptera (including Aphididae) and
small lepidopterous larvae. Here we de-
scribe feeding behavior of 3 species of Med-
etera from central Washington: Medetera
petulca Wheeler, M. n. sp. nr. alpina and
M. n. sp. nr. xerophila or utahensis. Med-
etera petulca was the most common species
encountered on apple and has previously
been collected on Salix and cultivated apple
trees (Bickel 1985). Present observations
cannot be linked to any one of the 3 species.
Medetera n. sp. nr. alpina and Medetera n.
sp. nr. xerophila or utahensis will be de-
scribed in a later publication.
During our study adult Medetera spp. were
observed on unsprayed 3-year-old apple
trees, Malus domestica (Borkhausen) var.
“Red Delicious.” The trees (ca. 1-1.2 m
high) were in 2 experimental “mini-or-
chards” planted within a deciduous riparian
habitat near the Wenatchee River (Wash-
ington: Chelan Co., elev. 220 m). The 2
““mini-orchards,”’ Sunnyslope (SS) and Riv-
er County Park (RCP), were being studied
by the senior author (RJR) to determine the
contribution of colonists from native hab-
itat to young apple trees. Vegetation sur-
rounding the 2 ““mini-orchards”’ was an im-
portant source pool for Medetera adults
colonizing apple and included: Rosa woodsii
Lindley (Rosaceae), Populus trichocarpa
Torrey & Grey (Salicaceae), Cornus stolo-
nifera Michaux (Cornaceae), Salix exigua
Nutall (Salicaceae), Crataegus douglasii
Lindley (Rosaceae), and 4melanchier alni-
folia Nutall (Rosaceae). Medetera adults
were collected in sweep samples of plants
surrounding the apple trees at both the RCP
and SS sites. The SS orchard, 10 km north-
west of Wenatchee, was partially shaded and
the RCP orchard, 12 km northwest of We-
natchee, was in direct sunlight. At 2-week
intervals from late May until late July, 1987,
visual observations were made on the trees
at each site. The behavior of adult Medetera
on the main stem, lateral branches, and fo-
liage of the apple trees was recorded by
ground level observations. Approximately
10 hours were spent recording Medetera
feeding behavior. All observations were
made between 8 a.m. and noon.
Flies remained in a stationary position for
ca. 90% of the total observation time with
prothoracic legs fully extended and meso-
and metathoracic legs pressed against the
body. Individual flies oriented themselves
at 30-40° angles to the branch or leaf upon
which they rested. During feeding the pro-
VOLUME 90, NUMBER 4
thoracic legs were retracted and the meso-
and metathoracic legs were extended, forc-
ing the head downward. Striking at leaf and
bark surfaces was repeated as a “pecking”
motion until either the prey had been se-
cured between the labellae or had escaped.
Medetera spp. adults were first noted
feeding on apple rust mites, Aculus schlecht-
endali (Nalepa) (Acari: Eriophyidae) on 2
June 1987 at the SS site. Flies were observed
making a “pecking” motion on bark and
leaf surfaces. Medetera spp. could have been
feeding on any micro-arthropods present
(scales, whiteflies, other mite species). How-
ever, examination of leaf and bark samples
in the laboratory with a binocular micro-
scope revealed eriophyids were the only ar-
thropods present in high numbers. Other
mite species, including 7etranychus spp. and
Panonychus ulmi (Koch) (Tetranychidae),
were absent from the trees at RCP and SS
in 1987. From May through August, 1987,
counts of eriophyids were made at 2-week
intervals. On 4 June at SS the rust mite
population peaked at ¥ = 232.0 + 80.39
mites/leaf [Mean + SE (n = 75)]. Feeding
by Medetera on rust mites at SS was also
observed on 16 and 20 June. Twenty-eight
“pecking actions” were recorded during an
8-minute observation period of one fly on
20 June. The observed fly concentrated
“pecking” at the veins on the upper leaf
surface. Examination of the leaf with a hand
lens (16 x) showed the highest density of A.
schlechtendali near the veins on the upper
surface of the leaf. One fly feeding on the
upper surface of a leaf was collected with
an aspirator and immediately transferred to
alcohol. In the laboratory the mouth parts
of this fly were examined, and a rust mite
was found between its labellae.
On 27 June, Medetera spp. were again
observed feeding on eriophyid mites at SS.
They were also seen consuming apple
aphids, Aphis pomi De Geer. Within a 15
minute period, 9 Medetera adults were ob-
served attacking apterous aphids which were
moving up and down a single apple tree.
511
Fig. 1.
Line drawing of male Medetera petulca
(Magnification = 30).
After capture, duration of feeding was ca.
60 seconds/aphid. Medetera were observed
preying on aphids on 3 additional dates: 11,
13, and 15 July at both the RCP and SS
sites. Estimates of mean aphid abundances
for 20 trees were 103.55 + 31.64 aphids/
tree and 1384 + 679.4 aphids/tree, at SS
on | and 15 July, respectively, and 332.5
+ 120.10 aphids/tree on 13 July at RCP.
The importance of predation by adult
dolichopodids on 2 apple pests, rust mites
and aphids, remains to be determined. Based
on these preliminary observations, we con-
clude that Medetera probably does not have
a significant impact as a biological control
agent on apple. To our knowledge this is the
first record of feeding by dolochopodid
adults on 4. schlechtendali and A. pomi. We
also observed Medetera feeding on imma-
ture thrips at both RCP and SS. Further
studies are planned to tie feeding observa-
tions with the different Wedetera species.
We thank W. J. Turner, C. H. Shanks, S.
C. Hoyt, and W. W. Cone for critical review
of the manuscript. W. J. Turner is also ac-
knowledged for taxonomic determination
of Medetera species, and J. C. Rodriguez is
acknowledged for the line drawing. We thank
T. Kono for identification of the eriophyid
mites. This research was supported by the
512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Washington State Tree Fruit Research ern Ave., Wenatchee, Washington 98801;
Commission and conducted under project Scott J. Hulbert, Department of Entomol-
1090. ogy, Washington State University, Pullman,
: Washington 99164.
Robin J. Rathman and Jay F. Brunner,
Tree Fruit Research Center, 1100 N. West-
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 513-523
OBITUARY
= ba x =e >
Carl Muesebeck in the 1920s
Carl Muesebeck in 1976
Carl Frederick William Muesebeck
1894-1987
Honorary President, Entomological Society of Washington,
1971-1987
The long career of Carl Muesebeck, one
of the 20th century’s most noted and pro-
ductive entomologists, came to an end at
age 93 on 13 November 1987. He was born
24 September 1894 in Medina, New York,
to William and Marie Koch Muesebeck.
Both parents emigrated to America in the
1880s. William (christened Carl Friedrich
Wilhelm Miisebeck'), came from Stettin,
Pomerania, in eastern Germany (now part
' Carl’s first cousin, Carl W. Muesebeck, traces the
Muesebeck ancestry as far back as the 10th century,
the family name being variously spelled Musbach, also
Muespach, and later Meusebach. A patent (or charter)
of nobility was conferred in 1690 on Johann Gregor
von Meusebach, Imperial Aulic Counselor, and Lord
Lieutenant and Treasurer of Upper and Lower Saxony.
of Poland), and Marie from Angermunde,
northeast of Berlin. They met and were
married in America. The father served an
apprenticeship as a tailor in Stettin, and
worked as a tailor in Medina, later moving
to Brockport to establish his own tailoring
business. The parents were Lutheran and
Carl was brought up in that faith. He left
the church after college, not caring for for-
mal ritual, but had a deep and abiding belief
in God throughout his life.
Carl attended elementary school in Medi-
na and then Brockport High School. He also
helped in his father’s tailoring business, and
greatly enjoyed working on a farm for six
months after high school graduation.
He entered Cornell University in 1912,
514
and was impressed at once by the Victorian
inscription on the gates of what was at that
time the principal entrance to the univer-
sity: “So enter that daily thou mayest be-
come more learned and more thoughtful; so
depart that daily thou mayest become more
useful to thy country and to mankind.” This
admonition guided Carl’s subsequent life,
and he quoted it admiringly to friends. He
thought that he might prepare himself to
become a professor of English, and he was
also keenly interested in mathematics. Dur-
ing his later undergraduate years he excelled
in tutoring foreign born students at Cornell
in English.
He had no particular interest in natural
history when he began his studies at Cornell,
but several courses under the inspired
teaching of John Henry Comstock and his
wife, Anna Botsford Comstock, were a rev-
elation to young Carl. Comstock was the
founder of the first department of ento-
mology in an American university, and his
wife was a gifted teacher also, but of general
natural history. Carl once told a reporter
that Comstock “made the study of insects
sound so fascinating that I switched to ento-
mology.”
Carl received a Bachelor of Science degree
in 1916. Inspired by his love of entomology,
he applied for and received his first profes-
sional appointment in December, 1916,
from the U.S. Department of Agriculture
(USDA) in the laboratory of the Bureau of
Entomology at Melrose Highlands, Mas-
sachusetts. He served there for nearly two
years in the grade of Scientific Assistant,
working on a biological study of the para-
sites of the introduced gypsy and brown-tail
moths.
He resigned from USDA in August, 1918,
to return to Cornell where he registered for
a doctorate. His graduate committee con-
sisted of Professors R. Matheson (insect
parasitology), J. C. Bradley (systematics),
and Moore (middle initials unknown, in
bacteriology). Moore was later replaced by
O. A. Johannsen (morphology) when Carl
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
changed his minor in bacteriology to par-
thenogenesis in insects. He also served as
an instructor in entomology at Cornell with
the responsibility for conducting courses in
biological control of insects and in taxon-
omy of parasitic Hymenoptera. The need to
provide more adequate support for his fam-
ily led him to terminate his graduate studies,
just a semester short of completing the man-
datory residency requirements for the doc-
torate. His doctoral thesis, the large revision
of North American Apanteles (#3), carried
the byline, “C. F. W. Muesebeck, Instructor
in Entomology, Cornell University.” He
mentioned in the introductory paragraph
that the contribution was the result of stud-
ies made during a temporary USDA ap-
pointment in the summer of 1919 to work
on the great mass of valuable material in
the collection at the U.S. National Museum,
and that contributory data had been ob-
tained earlier during his work at the Melrose
laboratory and during the winter and spring
of 1918-1919 at Cornell.
He returned to the USDA laboratory at
Melrose Highlands in April, 1921. For the
next decade he conducted research on bio-
systematics of parasitic Hymenoptera and
directed the work of other employees in this
field. His taxonomic studies during this pe-
riod resulted in the completion and publi-
cation of additional large revisionary stud-
ies on the Braconidae (#4, 6, 7, and 10).
Early in 1926 he was assigned temporarily
for 22 years to Budapest, Hungary, where
he had the responsibility for directing USDA
studies in Europe on the biology of Euro-
pean parasites of the gypsy and brown-tail
moths. He accepted this assignment with
the provision that he would be able to spend
periods during the winter months traveling
to various museums to study type speci-
mens of earlier workers in parasitic Hy-
menoptera. These visits included the col-
lections in Kiel, Dublin, London, Berlin and
2 Numbers in parentheses refer to articles listed in
the appended bibliography.
VOLUME 90, NUMBER 4
others. He travelled extensively in central
Europe searching for effective parasites of
these two forest pests, and was responsible
for collecting, rearing and shipping the bi-
ologically effective species to America.
During the stay in Europe he received the
first of a number of honors to be conferred
during his lifetime, that of honorary mem-
bership in the Hungarian Entomological So-
ciety. His entomological career was me-
morialized at the 583rd meeting of that
society on 19 February 1988.
In the fall of 1931 he was reassigned to
Washington to serve as Assistant Leader of
the Division of Insect Identification, USDA,
with additional responsibilities of carrying
on taxonomic research on several groups of
parasitic Hymenoptera, primarily the Bra-
conidae, Bethyloidea and Proctotrupoidea,
and providing identification service in these
groups. He also supervised the updating and
development of the insect host-parasite card
catalog that had been initiated by L. O.
Howard. Most of the national collection of
insects on which Carl worked was in the
U.S. National Museum, now known as the
National Museum of Natural History of the
Smithsonian Institution (SI). In 1935 he be-
came Leader of the Division of Insect Iden-
tification with his administrative office in
the South Building, USDA, 5 minutes across
the Mall from the Museum. He continued
his research and identification work on
parasitic Hymenoptera, and, as older staff
members retired or died, he added identi-
fication and curatorial responsibility for lice,
fleas and ticks to an already superhuman
load. In connection with his work on these
latter three groups of ectoparasites, he con-
tinued to add data to the card files of host-
parasite lists, and made these files available
to other investigators.
Carl was always a prodigious, dedicated
and conscientious professional, working 12
to 14 hours or more on weekdays, and fre-
quently additional hours on the weekend.
The normal working period for USDA em-
ployees early in 1941 was 8:45 a.m. to 5:15
215
p.m. By the time all of us had arrived at the
Museum, he would already have been work-
ing there from 6 a.m., and would have left
for his administrative office by 8:30 a.m.
Usually he returned at noon to have lunch
with us in the cafeteria of the Internal Rev-
enue Service Building across the street from
the Museum, bringing with him specimens
that required urgent identification or letters
that had been typed for us in the South
Building. He then went back to administra-
tive work for the afternoon, returning to the
Museum at 5 p.m. to spend another 3-4
hours over the microscope.
World War II imposed additional bur-
dens on him, for some of the younger spe-
cialists in the Division, J. F. Gates Clarke,
A. B. Gurney, D. G. Hall, K. V. Krombein,
P. W. Oman and B. E. Rees, entered mili-
tary service, most of them to serve as Army
entomologists. A caring, warmhearted per-
son, Carl wrote occasional lengthy personal
letters to each of us to send news of what
was happening at the Museum and what our
colleagues in military service were doing.
He looked forward with keen anticipation
to receiving letters with news of our ento-
mological activities in distant parts of the
world. He even found time to write several
times to the wife of one of our group to
inquire about her wellbeing and that of a
prematurely born child, and to ask for any
recent news from her husband.
During the war he was able to bring to
the Museum for shorter or longer periods
such specialists as M. T. James, V. S. L.
Pate and H. K. Townes, to curate parts of
the collection and to provide identification
service. Wartime transportation difficulties
meant that he was one of the few people to
continue working at the Museum on Sun-
days, usually between 8 a.m. and 3 p.m.
Earlier on Sunday and again late in the after-
noon, he visited the USDA research facility
at Beltsville to care for the silkworms being
reared there because no one else was avail-
able on that day to care for them.
He retired from Federal service in 1954,
516
having suffered a serious heart attack two
years earlier. Upon his retirement he was
appointed a Collaborator in the Insect Iden-
tification and Parasite Introduction Section,
USDA, and a Research Associate in the
Smithsonian Institution, the latter appoint-
ment enabling him to have an office and a
parking space at the Museum. Relieved of
the burden of administrative responsibili-
ties, he continued industriously working
over his microscope and cataloguing at the
Museum seven days a week from early in
the morning until noontime. After returning
home, he continued work on other ento-
mological projects such as editorial work
and additional cataloguing. From 1962 un-
til he left the Washington area in 1980 he
was editor for Entomological Review, the
translation of the important Russian peri-
odical, Entomologicheskoye Obozreniye.
With his usual meticulous attention, he
mastered the difficult Russian language, thus
ensuring that translations by the language
experts made good entomological sense.
Carl’s marriage in 1917, to Ida C. Praedel
of Brockport, New York, ended with her
death in 1975. An only son, Carl, Jr., died
suddenly from a brain tumor at age 16 in
1935. The loss of their gifted son was a
grievous blow to Carl and Ida. For many
years he took fresh flowers weekly from his
garden to the boy’s grave. Carl moved to
the state of Washington in 1980, and mar-
ried Luella M. Walkley, a former cataloguer
and specialist on Ichneumonidae in the Di-
vision of Insect Identification (USDA). She
died a year later, and he returned to New
York state in 1982 to live in Schenectady
with a cousin, Elfrieda Geissler. He was
physically active through 1986. Although
handicapped by occasional cardiac or res-
piratory problems, he enjoyed daily walks
along the Mohawk River and regularly drove
his car nearly 250 miles on the New York
Thruway several times a year between Sche-
nectady and a vacation home in western
New York. He remained in full mental vigor
until his death, enjoying occasional long dis-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tance conversations about Bethylidae,
friends in Washington, and the Washington
Redskins football team.
Carl was keenly interested in sports
throughout his life. During his high school
years he was a catcher on the baseball team.
At Cornell he participated annually in the
5-mile cross-country run in Ivy League
competition, never finishing less than 5th
in the race, and once coming in 2nd. He was
an active bowler during the late 1920s and
early 1930s, a sport in which he was joined
by several entomological colleagues. He be-
came particularly proficient at duckpins and
maintained a high average. He was so en-
thusiastic about this sport, that sometimes,
playing by himself, he used two adjacent
alleys, bowling a frame on one while the pins
were being set up in the other alley. An in-
jury to his foot while on the bowling alley
necessitated his withdrawal from active
sports. After the advent of television he
watched various sporting events, particu-
larly football, with great enthusiasm.
Another avocational interest was garden-
ing to which he was very dedicated. He
maintained a large garden at his home in
suburban Maryland, containing about 100
varieties of hybrid tearoses and other flow-
ers.
Carl was active in a number of profes-
sional societies including the Entomological
Society of America (Fellow in 1934, Second
Vice President in 1938, President in 1946,
Honorary Member in 1959), Entomological
Society of Washington (President in 1940,
Honorary Member in 1957, Honorary Pres-
ident from 1971 until his death), Biological
Society of Washington, Society of System-
atic Zoologists, Washington Academy of
Sciences, and Sigma Xi. He was also a mem-
ber of the Cosmos Club (1936-1954). The
Department of Agriculture recognized his
exceptional service by awarding him its Dis-
tinguished Service Award in 1951. The el-
oquent citation for this award paid tribute
to him, “For his contributions to science
and public welfare as an internationally rec-
VOLUME 90, NUMBER 4
ognized insect taxonomist; for his ability to
inspire and guide his associates in ento-
mology throughout the world in the acqui-
sition and dissemination of information on
destructive and beneficial insects; and for
his devotion to duty.”” The Eastern Branch
of the Entomological Society of America
honored his professional and societal ac-
complishments in 1978 by presenting him
its L. O. Howard Distinguished Achieve-
ment Award.
He was the author of more than a hundred
scientific contributions, many of them long-
er revisionary studies of the highest quality
such as his contributions on the genera
Apanteles (#3), Meteorus (#6), Macrocen-
trus (#21), Orgilus (#123), Macroteleia
(#135), Psilus and Coptera (#137), and his
revisions of higher categories such as Mi-
crogasterinae (#4), Braconinae (#10), and
Euphorinae (#32). His papers in systematics
were models of clarity and precision, re-
flecting his early love of the English lan-
guage.
His single most important contribution
to hymenopterology was the leadership that
resulted in the monumental synoptic cata-
log (#60), ““Hymenoptera of America North
of Mexico.” This catalog, published by
USDA as Agriculture Monograph No. 2 in
1951, was the first on the North American
fauna since 1887. It was a collaborative ef-
fort by a couple of dozen American and
Canadian specialists in various groups of
Hymenoptera. Carl served as one of three
co-editors and prepared 1n co-authorship the
sections on Braconidae and parts of Proc-
totrupoidea. His stature in the profession
ensured the reasonably prompt completion
of manuscripts by the 20 collaborators, and
his influence within the Department of Ag-
riculture facilitated the allotment of funds
from that agency for publication of this in-
dispensable tool of 1400+ pages. Twenty
years later, when the volume of publications
on North American Hymenoptera necessi-
tated preparation of an updated, comput-
erized edition by the Smithsonian and Ag-
S17;
riculture hymenopterists at the Museum, he
participated by preparing the sections on
Pelecinoidea, Proctotrupoidea and Cera-
phronoidea (#1 16-118).
The impact and stimulation generated by
the 1951 catalog were assessed astutely by
E. O. Wilson in a review of the revised 1979
edition (Science, 208: 721-722, 1980). He
stated in part: “In 1951 C. F. W. Muesebeck
and a group of associates provided a com-
plete taxonomic record of the species of
North America north of Mexico, with sum-
maries of natural history data. Important
syntheses always work toward their own ob-
solescence, and so it was that the Muesebeck
catalog stimulated a rush of new systematic
and biological studies.”
The Entomological Society of Washing-
ton dedicated the September 1969 issue of
its Proceedings to Carl in recognition of his
75th birthday. Dedicatory remarks by three
contributors are included here to give an
appreciation of the impact of his work in
three diverse groups of arthropods.
The late, eminent tick expert, Harry
Hoogstraal, honored his work on ticks by
describing a new argasid tick, Ornithodoros
(Alectorobius) muesebecki, from a remote
island off the coast of Saudi Arabia. In a
deft allusion to Carl’s great work on the
common names of insects, he christened this
bird tick, Muesebeck’s Arabian Booby Ar-
gasid. Carl, possessed of a wonderful sense
of humor, must have given a hearty guffaw
when he first read this name. Hoogstraal’s
dedicatory remarks were: “This new species
is dedicated to Mr. C. F. W. Muesebeck on
his 75th birthday in token recognition of
the selfless and devoted service that he has
provided to several generations of ento-
mologists throughout the world. The Jubi-
lee for the ever helpful Curator Muesebeck
thus extends to one of the most remote spots
of arthropod-inhabited land on the globe.”
The renowned flea specialist, Robert
Traub, prefaced his description of the new
flea genus, Muesebeckella, with the follow-
ing perceptive dedication: “Scientists fa-
518
miliar with the multitude of significant con-
tributions made by C. F. W. Muesebeck to
the study of Hymenoptera may be surprised
to see an article on Siphonaptera in a Jubilee
Volume dedicated in his honor. It would
seem that an entomologist who had accom-
plished so much in such a difficult field and
who also nevertheless somehow managed
to serve as an authority on the principles of
taxonomy and on the standardization of
common names of insects, could surely not
find the time to master the systematics of
fleas and lice. The fact is, however, that Mr.
Muesebeck is expert enough on both of these
groups of ectoparasites to have repeatedly
provided definitive identifications of ob-
scure species, despite the dearth of taxo-
nomic keys, proper descriptions and illus-
trations for which at least the Siphonapteran
literature is notorious. Even if this were not
the case, it would be fitting to include a
paper on fleas in this volume because of all
that Mr. Muesebeck has done to stimulate
students of ectoparasites and to ensure that
the U.S. Department of Agriculture and the
Smithsonian Institution would always be
able to identify specimens and answer quer-
ies submitted from all parts of the world.
When one considers the serious problems
posed over the years by the shortage of funds
to support such activities, the true value of
Mr. Muesebeck’s ‘extracurricular’ activities
becomes apparent.”
A fellow hymenopterist, Karl Krombein,
dedicated to him the new cuckoo wasp ge-
nus, Muesebeckidium, and offered these
thoughts: “During his many years at the U.S.
National Museum, formerly as the head of
systematic entomology investigations for the
U.S. Department of Agriculture, and pres-
ently as an honored Research Associate of
the Smithsonian Institution, I have come to
know and cherish Carl Muesebeck very
highly. His quietly skillful and dynamic
leadership of the Insect Identification Di-
vision, USDA, brought it to a peak in
professional and support strength not
equaled before or since. Had it not been for
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
his effective and tactful direction, an ep-
ochal contribution such as the catalog of
North American Hymenoptera might not
be a reality today. Finally, his warm per-
sonal interest in the professional develop-
ment and well-being of his staff endeared
him to all of those privileged to work under
his leadership.”
In another article in the issue of the Pro-
ceedings celebrating Carl’s 75th birthday, R.
H. Nelson, Executive Director Emeritus,
Entomological Society of America (ESA),
wrote at length of Carl’s numerous services
to that society. Among other activities in
ESA Carl was a member of the small, joint
committee that proposed the merger of the
original ESA with the American Associa-
tion of Economic Entomologists to form the
present, larger national society representing
all of entomology.
All of his professional life Carl had a kind-
ly concern for his associates, and for the
friends and neighbors outside of his profes-
sion. Always generous in his praise of sci-
entific contributions by others, as late as
1986 he expressed his great delight in re-
ceiving the monumental study of the Rus-
sian Braconidae by Tobias and his esteem
for the quality of the work.
One reviewer of the revised Hymenop-
tera catalog commented felicitously (Quart.
Rey. Biol., 55: 444-445, 1980) that “.. . all
among its users will appreciate and find most
fitting its dedication, with admiration and
affection, to Mr. C. F. W. Muesebeck, him-
self a paragon among entomologists.”” Carl
was a gentleman and a gentle man, loved
by his friends and close colleagues, and held
in affectionate regard and respect by his as-
sociates all over the world.
Acknowledgments.—We are grateful to
Carl’s cousin, Miss Elfrieda Geissler, Sche-
nectady, for sharing with us her recollec-
tions of things he had told her about his
family and early years. We also thank Victor
E. Krantz, Chief of the Natural History
Branch of the Photographic Division, SI, for
his skillful copying of the photographs from
VOLUME 90, NUMBER 4
Miss Geissler that are used in our sketch.
Several colleagues, J. F. Gates Clarke, Ken-
neth W. Cooper, John G. Franclemont, Beth
and Arnold Norden, Paul W. Oman, Louise
M. Russell, Curtis W. Sabrosky, and Robert
Traub were kind enough to comment on a
preliminary draft and/or to provide helpful
information. Mrs. Isabel T. S. Gardner, Of-
fice of the Dean, The Graduate School, pro-
vided the available information from Cor-
nell archives on Carl’s registration for
graduate work.
Karl V. Krombein, Department of Ento-
mology, Smithsonian Institution, NHB 105,
Washington, D.C. 20560.
Paul M. Marsh, Systematic Entomology
Laboratory, USDA, % Smithsonian Insti-
tution, NHB 168, Washington, D.C. 20560.
Bibliography of
Carl F. W. Muesebeck
1. Muesebeck, C. F. W. 1918. Two important in-
troduced parasites of the brown-tail moth. J. Agric.
Res. 14: 191-206.
2. Muesebeck, C. F. W. 1919. Three new species
of Braconidae. Canad. Entomol. 51: 113-115.
3. Muesebeck, C. F. W. 1920. A revision of the
North American species of ichneumon-flies be-
longing to the genus Apanteles. Proc. U.S. Natl.
Mus. 58: 483-576.
4. Muesebeck, C. F. W. 1922. A revision of the
North American ichneumon-flies belonging to the
subfamilies Neoneurinae and Microgasterinae.
Proc. U.S. Natl. Mus., Vol. 61, Art. 15, 76 pp.
5. Muesebeck, C. F. W. 1922. Zygobothria nidi-
cola, an important parasite of the brown-tail moth.
Bull. U.S. Dept. Agric. No. 1088, 9 pp.
6. Muesebeck, C. F. W. 1923. A revision of the
North American species of ichneumon-flies be-
longing to the genus Meteorus Haliday. Proc. U.S.
Natl. Mus., Vol. 63, Art. 2, 44 pp.
7. Muesebeck, C. F. W. 1925. A revision of the
parasitic wasps of the genus Microbracon occur-
ring in America north of Mexico. Proc. U.S. Natl.
Mus. Vol., 67, Art. 8, 85 pp.
8. Muesebeck, C. F. W. 1926. Descriptions of new
reared parasitic Hymenoptera and some notes on
synonymy. Proc. U.S. Natl. Mus., Vol. 69, Art.
7, 18 pp.
9. Muesebeck, C. F. W. and C. McCann. 1926. A
new ichneumon fly of the genus Apanteles from
24. Muesebeck, C. F. W.
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519
the death’s head moth Acerontia styx, with a note
on its life history. J. Bombay Nat. Hist. Soc. 31:
726-728.
1927. A revision of the
parasitic wasps of the subfamily Braconinae oc-
curring in America north of Mexico. Proc. U. S.
Natl. Mus., Vol. 69, Art. 16, 73 pp.
. Muesebeck, C. F.W. 1927. New species of chal-
cid flies parasitic on the gypsy-moth parasite,
Apanteles melanoscelus (Ratzeburg). J. Agric. Res.
34: 331-333.
. Muesebeck, C. F. W. and S. M. Dohanian.
1927. A study in hyperparasitism, with partic-
ular reference to the parasites of Apanteles mel-
anoscelus (Ratzeburg). Bull. U.S. Dept. Agric. No.
1487, 35 pp.
1928. A new European
species of Apanteles parasitic on the gypsy moth.
Proc. Entomol. Soc. Wash. 30: 8.
. Muesebeck, C. F.W. 1928. Family Braconidae,
pp. 895-920. In M. D. Leonard, ed., List of the
Insects of New York. Cornell Univ. Agric. Exp.
Sta. Mem. No. 101, 1121 pp.
1929. Two new species
of Apanteles (Hymenoptera: Braconidae). Proc.
Entomol. Soc. Wash. 31: 118-120.
1931. Description of a
new genus and eight new species of ichneumon-
flies with taxonomic notes. Proc. U.S. Natl. Mus.,
Vol. 79, Art. 16, 16 pp.
1931. Monodontomerus
aereus Walker, both a primary and a secondary
parasite of the brown-tail moth and the gypsy
moth. J. Agric. Res. 43: 445-459.
1932. The genus Meso-
coelus Schulz (Hymenoptera, Braconidae). Proc.
Biol. Soc. Wash. 45: 227-230.
1932. Two new species
of Phanomeris Foerster (Hymenoptera, Bracon-
idae) parasitic on leaf-mining sawflies. Proc.
Entomol. Soc. Wash. 34: 81-83.
1932. Four new North
American species of Bassus Fabricius (Hyme-
noptera: Braconidae), with notes on the genotype.
J. Wash. Acad. Sci. 22: 329-333.
1932. Revision of the
Nearctic ichneumon-flies belonging to the genus
Macrocentrus. Proc. U.S. Natl. Mus., Vol. 80,
Art. 23, 55 pp.
1932. Two new species
of phytophagous Eurytomidae (Hymenoptera:
Chalcidoidea). Proc. Entomol. Soc. Wash. 34:
109-112.
. Muesebeck, C. F. W. 1933. Five new hymenop-
terous parasites of the Oriental fruit moth. Proc.
Entomol. Soc. Wash. 35: 48-54.
1933. Seven new species
520
25. Muesebeck, C. F. W. and D. L. Parker.
28. Muesebeck, C. F. W.
34.
35.
36.
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. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of reared Braconidae (Hymenoptera). Proc. Ento-
mol. Soc. Wash. 35: 193-200.
1933.
Hyposoter disparis Viereck, an introduced ich-
neumonid parasite of the gypsy moth. J. Agric.
Res. 46: 335-347.
1934. Seven new species
of Indian Bethylidae (Hymenoptera). Rec. Indian
Mus. 36: 223-232.
. Muesebeck, C. F. W. 1935. Synonymical notes
on Ecphylus Foerster, with description of one new
species (Hym., Braconidae). Proc. Entomol. Wash.
37: 21-24.
1935. Three new reared
parasitic Hymenoptera, with some notes on syn-
onymy. J. Wash. Acad. Sci. 25: 270-283.
1935. On the genus On-
cophanes Foerster, with descriptions of two new
related genera (Hymenoptera: Braconidae). Ann.
Entomol. Soc. Amer. 28: 241-250.
1935. A new species of
parasite of Typhlocyba pomaria McAtee (Hy-
menoptera: Bethylidae). Proc. Entomol. Soc.
Wash. 37: 167-168.
1935. On two little known
genera of Braconidae (Hymenoptera). Proc. Ento-
mol. Soc. Wash. 37: 173-177.
. Muesebeck, C. F. W. 1936. The genera of para-
sitic wasps of the Braconid subfamily Euphori-
nae, with a review of the Nearctic species. Misc.
Publ. U. S. Dept. Agric. No. 241, 36 pp.
1937. Anew West Indian
species of Mirax Haliday, parasitic on the coffee
leaf-miner (Hymenoptera: Braconidae). Proc.
Entomol. Soc. Wash. 39: 139-141.
Muesebeck, C.F. W. 1937. Three new Brazilian
species of Heterospilus (Hym., Braconidae), par-
asites of Gasterocercodes gossypii Pierce. Rev.
Entomol. 7: 8-11.
Muesebeck, C. F. W. 1938. Two reared North
American species of the genus Stantonia Ash-
mead (Hymenoptera: Braconidae). Proc. Ento-
mol. Soc. Wash. 40: 89-91.
Muesebeck, C. F. W. 1938. New species and
synonymy in the genus Macrocentrus (Hymenop-
tera: Braconidae). Proc. Entomol. Soc. Wash. 40:
170-172.
1938. Three new reared
species of Apanteles from California (Hymenop-
tera: Braconidae). Proc. Entomol. Soc. Wash. 40:
201-204.
. Muesebeck, C. F. W. 1938. The genus Dendro-
soter Wesmael in the Unites States (Hymenop-
tera: Braconidae). Proc. Entomol. Soc. Wash. 40:
281-287.
1939. Three new hymen-
opterous parasites of the Lemna fly. Proc. Ento-
mol. Soc. Wash. 41: 58-62.
40.
41.
44.
46.
47.
48.
49.
1;
33:
54.
55.
56.
a7.
2. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
Muesebeck, C. F. W. 1939. A new species of
Heterospilus parasitic on an injurious anobuid
(Hymenoptera: Braconidae). Proc. Entomol. Soc.
Wash. 41: 62-63.
Muesebeck, C. F. W. 1939. Five new species
of Meteorus (Hymenoptera: Braconidae). Proc.
Entomol. Soc. Wash. 41: 83-87.
1939. A new reared Me-
teorus from Tasmania (Hymenoptera: Braconi-
dae). Proc. Entomol. Soc. Wash. 41: 172-173.
1939. A new mealybug
parasite (Hymenoptera: Scelionidae). Canad.
Entomol. 71:158-160.
Muesebeck, C. F. W. 1939. The North Amer-
ican species of the genus Laelius (Hymenoptera:
Bethylidae). Proc. Biol. Soc. Wash. 52: 171-176.
1940. Two new reared
species of Bassus (Hymenoptera: Braconidae).
Proc. Entomol. Soc. Wash. 42: 91-93.
Muesebeck, C. F.W. 1940. Two new hymenop-
terous parasites of sugar-cane borers in India. Proc.
Entomol. Soc. Wash. 42: 120-122.
Muesebeck, C. F. W. 1941. Two new reared
species of Doryctes (Hymenoptera: Braconidae).
Proc. Entomol. Soc. Wash. 43: 149-152.
Muesebeck, C. F. W. 1941. A new ant parasite
(Hymenoptera, Braconidae). Bull. Brooklyn
Entomol. Soc. 36: 200-201.
Muesebeck, C. F. W. 1942. Common names.
J. Econ. Entomol. 34: 862-863.
1942. Fundamental taxo-
nomic problems in quarantine and nursery in-
spection. J. Econ. Entomol. 35: 753-758.
Muesebeck, C. F. W. 1942. Two new species
of Allotropa (Platygasteridae, Serphoidea) para-
sitic on the Comstock mealybug. Bull. Brooklyn
Entomol. Soc. 37: 170-173.
1945. On the interpre-
tation of Article 23 of the International Code in
cases where a species is originally described as
being both in a genus and in a subgenus and later
the subgenus is elevated to generic rank and the
species is transferred to the genus so erected. Bull.
Zool. Nomen., August 1945, p. 92.
Muesebeck, C. F. W. 1946. A new Apanteles
from Hawaii (Hym., Braconidae). Proc. Hawai-
ian Entomol. Soc. 12: 615-616.
Muesebeck, C. F. W. 1947. Two new species
of Apanteles from California (Hymenoptera: Bra-
conidae). Pan-Pac. Entomol. 23: 21-24.
Muesebeck, C. F. W. 1949. A new flightless
Phaenopria (Hymenoptera: Diapriidae). Canad.
Entomol. 81: 234-235.
Muesebeck, C. F. W. 1949. Two new species
of Opius from rose hips. Canad. Entomol. 81:
254-256.
Muesebeck, C. F. W. 1950. Two new genera
VOLUME 90, NUMBER 4
58.
60.
61.
64.
65.
66.
67.
68.
69.
70.
7K.
. Muesebeck, C. F. W.
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and three new species of Braconidae. Proc. Ento-
mol. Soc. Wash. 52: 77-81.
Muesebeck, C. F. W. 1950. Common names of
insects approved by the American Association of
Economic Entomologists. J. Econ. Entomol. 43:
117-138.
. Muesebeck, C. F. W., A. B. Gahan, and E. R.
Sasscer. 1951. Sievert Allen Rohwer, 1888-1951
{obituary]. Proc. Entomol. Soc. Wash. 53: 227-
230.
Muesebeck, C. F. W., K. V. Krombein, and H.
K. Townes [eds.]. 1951. Hymenoptera of Amer-
ica North of Mexico—Synoptic Catalog. U.S.
Dept. Agric., Agric. Monog. 2, 1420 pp.
Muesebeck, C. F. W. and L. M. Walkley. 1951.
Family Braconidae, pp. 90-184. In C. F. W.
Muesebeck et al., eds., Hymenoptera of America
North of Mexico—Synoptic Catalog. U.S. Dept.
Agric., Agric. Monog. 2, 1420 pp.
2. Muesebeck, C. F. W. and L. M. Walkley. 1951.
Superfamily Bethyloidea, pp. 726-734. In
C. F. W. Muesebeck et al., eds., Hymenoptera of
America North of Mexico— Synoptic Catalog. U.S.
Dept. Agric., Agric. Monog. 2, 1420 pp.
1952. What kind of insect
is it? Yearbook of Agriculture for 1952, pp. 43-
55:
Muesebeck, C. F. W. 1952. Progress in insect
classification. Yearbook of Agriculture for 1952,
pp. 56-60.
Muesebeck, C. F. W. 1953. Three new reared
Braconidae (Hymenoptera). Proc. Entomol. Soc.
Wash. 55: 149-151.
Muesebeck, C. F. W. 1953. Scientific names of
the body and head lice. J. Econ. Entomol. 46:
524.
Muesebeck, C. F. W. 1953. National entomo-
logical societies merge. Science 117: 546-547.
Muesebeck, C. F. W. 1954. Three new mealy-
bug parasites of the genus 4//otropa (Hymenop-
tera: Platygasteridae). Bull. Brooklyn Entomol.
Soc. 49: 18-21.
Muesebeck, C. F. W. 1954. Some braconid par-
asites of the pink bollworm Pectinophora gossy-
piella (Saunders). Boll. Lab. Entomol. Agric. Fi-
lippo Silvestri 33: 57-68.
Muesebeck, C. F. W. 1955. New reared Bra-
conidae from Trinidad (Hymenoptera). Proc.
Entomol. Soc. Wash. 57: 161-164.
Muesebeck, C. F. W. 1955. A remarkable new
species of Perilitus from Mexico (Hymenoptera:
Braconidae). Proc. Biol. Soc. Wash. 68: 143-144.
2. Shands, W. A., G. W. Simpson, F. S. Roberts,
and C. F. W. Muesebeck. 1955. Parasites of
potato-infesting aphids and of some other aphids
in Maine. Proc. Entomol. Soc. Wash. 57: 131-
136.
1956. New synonymy in
74.
73:
76.
Wis
78.
19:
80.
81.
84.
85.
86.
87.
88.
. Muesebeck, C. F. W.
. Shenefelt, R. D. and C. F. W. Muesebeck.
521
the Braconidae. Proc. Entomol. Soc. Wash. 58:
34.
Muesebeck, C. F. W. 1956. A braconid parasite
of a psocid (Hymenoptera). Proc. Entomol. Soc.
Wash. 58: 148-149.
Muesebeck, C. F. W. 1956. Two new braconid
parasites of the avocado looper (Hymenoptera:
Braconidae). Pan-Pac. Entomol. 32: 25-28.
Muesebeck, C. F. W. 1956. Two new parasites
of the yellow clover aphid and the spotted alfalfa
aphid (Hymenoptera: Braconidae). Bull. Brook-
lyn Entomol. Soc. 51: 25-28.
Muesebeck, C. F. W. 1956. On Opius ferrugi-
neus Gahan and two closely similar new species
(Hymenoptera: Braconidae). Entomol. News 67:
99-102.
Muesebeck, C. F. W. 1956. New generic syn-
onymy in the Scelionidae (Hymenoptera). Proc.
Entomol. Soc. Wash. 58: 24.
Muesebeck, C. F. W. and L. M. Walkley. 1956.
Type species of the genera and subgenera of para-
sitic wasps comprising the superfamily Procto-
trupoidea (order Hymenoptera). Proc. U.S. Natl.
Mus. 105: 319-419.
Muesebeck, C. F. W. 1957. New World Apan-
teles parasitic on Diatraea (Hymenoptera: Bra-
conidae). Entomol. News 68: 19-25.
Muesebeck, C. F. W. 1957. Four new species
of Eubadizon from Western United States (Hy-
menoptera: Braconidae). Bull. Brooklyn Ento-
mol. Soc. 52: 51-56.
1957. Robert Asa Cush-
man, 1880-1957 [obituary]. Proc. Entomol. Soc.
Wash. 59: 247-248.
1957.
Ashmead’s Meteoridea (Hymenoptera: Braconi-
dae). Proc. Entomol. Soc. Wash. 59: 129-134.
Muesebeck, C. F. W. 1958. Family Braconidae,
pp. 18-36. Jn F. V. Krombein et. al., eds., Hy-
menoptera of America North of Mexico—Syn-
optic Catalog. U.S. Dept. Agric., Agric. Monog.
2, Suppl. 1, 305 pp.
Muesebeck, C. F. W. 1958. Superfamily Proc-
totrupoidea, pp. 88-94. In K. V. Krombein et al.,
eds., Hymenoptera of America North of Mexi-
co—Synoptic Catalog. U.S. Dept. Agric., Agric.
Monog. 2, Suppl. 1, 305 pp.
Muesebeck, C. F. W. 1958. Three new aphid
parasites from the Pacific coast (Hymenoptera:
Braconidae, Aphidiinae). Entomol. News 69: 141-
145.
Muesebeck, C. F. W. 1958. New Neotropical
wasps of the family Braconidae (Hymenoptera)
in the U.S. National Museum. Proc. U.S. Natl.
Mus. 107: 4054461.
Muesebeck, C. F. W. and B. R. Subba Rao.
1958. A new braconid parasite of Hymenia re-
522
89.
90.
Ot
94.
95:
96.
97:
98.
99.
100.
101.
102.
103.
104.
105.
106.
2. Cory, E. N. and C. F. W. Muesebeck.
. Muesebeck, C. F. W.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
curvalis (Fabricius). Indian J. Entomol. 20: 27-
28.
Muesebeck, C. F. W. 1959. New reared species
of Lygocerus Foerster (Hymenoptera: Ceraphron-
idae). Entomol. News 70: 91-96.
Muesebeck, C. F. W. 1960. New reared Neo-
tropical species of Rogas Nees (Hymenoptera:
Braconidae). Entomol. News 71: 257-261.
Muesebeck, C. F. W. 1960. A fossil braconid
wasp of the genus Ecphylus (Hymenoptera). J.
Paleont. 34: 495-496.
1960.
Arthur Burton Gahan, 1880-1960 [obituary].
Proc. Entomol. Soc. Wash. 62: 198-204.
1961. A new Opius and
two new species of Microctonus (Hymenoptera:
Braconidae). Bull Brooklyn Entomol. Soc. 56: 57—
61.
Muesebeck, C. F. W. 1961. A new Japanese
Trichopria parasitic on the house fly (Hymenop-
tera: Diapriidae). Mushi 35: 1-2.
Muesebeck, C. F. W. 1961. Annual Review of
Entomology, volume 6, 1961 [book review]. Bull.
Entomol. Soc. Amer. 7: 193.
Muesebeck, C. F. W. 1962. A new Hawaiian
Opius from a leaf-mining pest of beans (Hyme-
noptera: Braconidae). Proc. Hawaiian Entomol.
Soc. 18: 289-290.
Muesebeck, C. F. W. 1962. Annual Review of
Entomology, volume 7, 1962 [book review]. Bull.
Entomol. Soc. Amer. 8: 93.
Muesebeck, C. F. W. 1963. Six new reared
species of Bracon (Hymenoptera: Braconidae).
Entomol. News 74: 157-165.
Muesebeck, C. F. W. 1963. Host relationships
of the Euphorini (Hymenoptera: Braconidae).
Proc. Entomol. Soc. Wash. 65: 300.
Muesebeck, C. F. W. 1963. A new ceraphronid
from cretaceous amber (Hymenoptera: Procto-
trupoidea). J. Paleont. 37: 129-130.
Muesebeck, C. F.W. 1963. A platygasterid par-
asite of certain wasp larvae (Hymenoptera: Proc-
totrupoidea, Platygasteridae). Beitr. Entomol. 13:
391-394.
Muesebeck, C. F. W. 1963. Annual Review of
Entomology, volume 8, 1963 [book review]. Bull.
Entomol. Soc. Amer. 9: 120.
Muesebeck, C. F.W. 1964. Identity of the genus
Miocolus Foerster (Hymenoptera: Braconidae).
Proc. Entomol. Soc. Wash. 66: 88.
Muesebeck, C. F.W. 1964. Insects of Campbell
Island. Hymenoptera: Braconidae. Pac. Ins.
Monogr. 7: 494-495.
Muesebeck, C. F. W. 1964. Annual Review of
Entomology, volume 9, 1964 [book review]. Bull.
Entomol. Soc. Amer. 10: 115.
Muesebeck, C. F. W. 1965. Two new braconid
107.
108.
109.
110.
111.
114.
116.
1) Ug
118.
IMs
2. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
5. Muesebeck, C. F. W.
. Masner, L. and C. F. W. Muesebeck.
parasites of the spruce budworm (Hymenoptera).
Entomol. News 76: 71-74.
Muesebeck, C. F.W. 1965. A new diapriid (Hy-
menoptera: Diapriidae) from termite nests from
South Africa. J. Entomol. Soc. S. Afr. 27: 188-
190.
Muesebeck, C. F. W. 1965. Annual Review of
Entomology, volume 10, 1965 [book review]. Bull.
Entomol. Soc. Amer. 11: 95-96.
Shands, W. A., G. W. Simpson, C. F. W. Muese-
beck, and H. W. Wane. 1965. Parasites of po-
tato-infesting aphids in northeastern Maine.
Maine Agric. Exp. Sta. Bull. T19, 77 pp.
Muesebeck, C. F. W. 1966. Annual Review of
Entomology, volume 11, 1966 [book review]. Bull.
Entomol. Soc. Amer. 12: 233.
Muesebeck, C. F. W. 1967. Agathis metzneria
Muesebeck, new species. /n C. Juhala, ed., Notes
on parasitic Hymenoptera associated with a ge-
lechiid moth, Metzneria lappella, in the common
burdock, and description of a new species of
Agathis (Braconidae). Ann. Entomol. Soc. Amer.
60: 95-97.
1967. Three new south-
eastern parasitic Hymenoptera. Fla. Entomol. 50:
57-61.
1967. A new braconid
parasite of the potato tuberworm (Hymenoptera).
Proc. Entomol. Soc. Wash. 69: 177-178.
Muesebeck, C. F. W. 1967. Three new reared
Braconidae (Hymenoptera). Entomol. News 78:
135-139.
1967. Annual Review of
Entomology, volume 12, 1967 [book review]. Bull.
Entomol. Soc. Amer. 13: 134.
Muesebeck, C. F.W. 1967. Family Braconidae,
pp. 27-60. Jn K. V. Krombein and B. D. Burks,
eds., Hymenoptera of America North of Mexico
—Synoptic Catalog. U.S. Dept. Agric., Agric.
Monogr. 2, Suppl. 2, 584 pp.
Muesebeck, C. F. W. 1967. Superfamily Pele-
cinoidea, p. 285. Jn K. V. Krombein and B. D.
Burks, eds., Hymenoptera of America North of
Mexico—Synoptic Catalog. U.S. Dept. Agric.,
Agric. Monogr. 2, Suppl. 2, 584 pp.
Muesebeck, C. F. W. and L. Masner. 1967. Su-
perfamily Proctotrupoidea, pp. 285-305. Jn
K. V. Krombein and B. D. Burks, eds., Hyme-
noptera of America North of Mexico—Synoptic
Catalog. U.S. Dept. Agric., Agric. Monogr. 2,
Suppl. 2, 584 pp.
Muesebeck, C. F. W. 1968. Aphid parasites of
Czechoslovakia, by P. Stary [book review]. Bull.
Entomol. Soc. Amer. 14: 80.
1968. The
types of Proctotrupoidea (Hymenoptera) in the
VOLUME 90, NUMBER 4
126.
129.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
United States National Museum. U.S. Natl. Mus.
Bull. No. 270, 143 pp.
1969. Annual Review of
Entomology, volume 14, 1969 [book review]. Bull.
Entomol. Soc. Amer. 15: 382-383.
1970. A new mealybug
parasite from Japan. Proc. Entomol. Soc. Wash.
72: 318-319.
. Muesebeck, C. F.W. 1970. The Nearctic species
of Orgilus Haliday (Hymenoptera: Braconidae).
Smith. Contr. Zool. No. 30, 104 pp.
1970. Annual Review of
Entomology, volume 15, 1970 [book review]. Bull.
Entomol. Soc. Amer. 16: 130-131.
1971. Annual Review of
Entomology, volume 16, 1971 [book review]. Bull.
Entomol. Soc. Amer. 17: 291-292.
Burks, B. D., C. F. W. Muesebeck, and R. D.
Gordon. 1971. Edward Albert Chapin, 1894—
1969 [obituary]. Proc. Entomol. Soc. Wash. 73:
99-104.
1972. On the identity of
Agonophorus Dahlbom (Proctotrupoidea, Dia-
priidae). Proc. Entomol. Soc. Wash. 74: 131.
1972. Nearctic species of
Scelionidae (Hymenoptera: Proctotrupoidea) that
parasitize the eggs of grasshoppers. Smith. Contr.
Zool. No. 122, 33 pp.
Muesebeck, C. F.W. 1972. A new reared species
of Trichopria (Proctotrupoidea, Diapriidae).
Entomol. News 83: 141-143.
130.
136.
137,
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
. Muesebeck, C. F. W.
523
Muesebeck, C. F. W. 1972. Annual Review of
Entomology, volume 17, 1972 [book review]. Bull.
Entomol. Soc. Amer. 18: 156-157.
. Cushman, H. G., C. F. W. Muesebeck, and G.
B. Vogt. 1972. Warren Samuel Fisher, 1878-
1971 [obituary]. Proc. Entomol. Soc. Wash. 74:
344-352.
1973. Annual Review of
Entomology, volume 18, 1973 [book review]. Bull.
Entomol. Soc. Amer. 19: 125-126.
1974. A new Nearctic
species of Telenomus (Hymenoptera: Proctotru-
poidea, Sceiionidae). Folia Entomol. Hung. 27:
135-137.
1974. Annual Review of
Entomology, volume 19, 1974 [book review]. Bull.
Entomol. Soc. Amer. 20: 271-272.
. Muesebeck, C. F. W. 1977. The parasitic wasps
of the genus Macroteleia Westwood of the New
World (Hymenoptera, Proctotrupoidea, Scelion-
idae). U.S. Dept. Agric. Tech. Bull. No. 1565, 57
pp.
Muesebeck, C. F. W. 1978. A new egg parasite
(Hymenoptera: Scelionidae) of the elm span-
worm, Ennomos subsignarius (Lepidoptera: Geo-
metridae). Proc. Entomol. Soc. Wash. 80: 91-93.
Muesebeck, C. F. W. 1980. The Nearctic para-
sitic wasps of the genera Psi/us Panzer and Cop-
tera Say (Hymenoptera, Proctotrupoidea, Dia-
priidae). U.S. Dept. Agric. Tech. Bull. No. 1617,
71 pp.
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 524-528
Book REVIEW
Acarology. Mites and Human Welfare.
Tyler A. Woolley. Wiley-Interscience
(John Wiley & Sons), New York. 1988,
484 pp. $57.75.
The mites, more than any other group,
have successfully explored the advantages
of combining the arthropod body plan with
small size. Evidence lies in their tremen-
dous diversity, whether the latter term is
used in the context of taxonomy or of ad-
aptations affecting morphology, nutrition,
or life-cycle. This book is intended as a gen-
eral textbook of acarology, and thus an in-
troduction to this diversity.
The book contains 23 chapters, arranged
in four parts, plus a brief Appendix con-
taining references relating to study tech-
niques and control. The first part presents
glimpses of biological diversity (Chapter 1),
offers some generalizations on morphology
(Chapter 2), and attempts a general discus-
sion of the Arachnida and the relationships
of mites within the class. The second part,
comprising more than 60% of the book, deals
with external morphology (Chapters 4—6),
internal anatomy (Chapters 7-12), and
pheromones (Chapter 13). Coverage of oth-
er aspects of biology is interspersed in ways
which are not immediately obvious from
chapter headings. As examples, feeding bi-
ology is discussed in Chapter 7 (Digestive
System), behavior is treated in Chapter 11
(Nervous System), and embryology and life
cycles are covered in Chapter 12 (Repro-
ductive System). The third part (about 20%
of the book) deals with classification. As an
introduction, Chapter 14 contains a brief
historical review and an ineffectually illus-
trated key to orders; following this is one
chapter on each of the seven recognized or-
ders in the subclass Acari (Chapters 1 5-21).
Very brief, superficial chapters on the his-
tory of acarology (Chapter 22) and ecology
(Chapter 23) compose the fourth part of the
book. There is no glossary. References are
listed at the end of each chapter, which fa-
cilitates access with only a minor cost in
redundancy for those which are repeated
throughout the book. Including such repe-
tition, there are slightly more than 2000 ci-
tations; about 17% are of non-English lan-
guage sources, but often these are indirect
(“fide’’) references.
At first opening, the book seems to be an
attractive, polished, informative product,
the kind we would expect from a major pub-
lisher of scientific texts. Yet, after even a
quick perusal of chapter headings, one starts
to question its subtitle, “Mites and Human
Welfare.”’ Most readers will expect to find
substantive discussions of mites important
to medical and veterinary science, mites as
agricultural pests, mites as biocontrol agents
acting on arthropod pests of crops and for-
ests, and mites of interest in stored product
protection. They will be disappointed; there
is no significant treatment of these topics.
Reading lists provided at the end of the
three-page chapter on ecology (Chapter 23)
and after an Appendix section entitled
“Control,” will lead the ambitious reader to
a small portion of the important literature,
but the subtitle is nonetheless greatly mis-
leading. Indeed it is mysterious that it could
even be considered. As examples: there is
only cursory mention of the important hon-
eybee parasites Acarapis woodi and Varroa
Jacobsoni, which are causing widespread
concern in North America; there is no
mention of the much-publicized Lyme dis-
ease or its tick vector; and the human par-
asite Sarcoptes scabei is briefly mentioned
in several places, but the important disease
it engenders (scabies) is ignored.
The book is illustrated with 220 text fig-
ures, many of which are compound plates.
They include both line drawings and scan-
ning electron micrographs (SEMs). Most of
the figures are redrawn from original sources,
VOLUME 90, NUMBER 4
but a few are clearly copies of originals; no
distinction is made in the captions, and at
least one (Figs. 6-10A) is wrongly attrib-
uted. A frequent problem with SEMs (and
some line drawings) is the absence of la-
beling, such that the caption describes only
a small part of the figure, which the reader
may not be able to locate (e.g. Figs. 2-3, 4—
5, 11-11). Occasionally, figure labels do
not match caption descriptions, or are not
explained at all, and sometimes the absence
of notes on orientation will cause confusion.
Text references to figures are sometimes i1n-
correct; for example, in a discussion of mus-
culature in gamasid mites (p. 103) the cited
figure is of Listrophorus, one of the Astig-
mata.
Like any text of wide scope, a book at-
tempting to cover the field of acarology is
dificult to write, and one cannot hope to
include all relevant data and literature. But
success is governed by more than the num-
ber of facts or citations one can squeeze into
the text. Just as important are effectiveness
of synthesis and various mechanical con-
cerns, such as organization, clarity of writ-
ing, and the ease with which the book can
be used for reference. In these areas, which
are within the purview of the editors as well
as the author, the book is deficient. Much
of it seems to have been written as a mosaic,
with sections simply juxtaposed after being
written at different times, using different
sources. The result is often a conspicuous
lack of synthesis and an unusually large
number of mechanical problems. The qual-
ity of organization within major sections of
text is inconsistent, and at times the flow of
ideas is nearly impossible to discern. Fre-
quently a subject is introduced, dropped for
another, then returned to just as abruptly.
Ideas or summaries may be repeated several
times within a chapter, with the same or
different wording; reinforcement is often a
valid writing technique, but clearly that is
not the intent in most cases. The discussion
in Chapter 3, regarding phylogenetic rela-
tionships with other arachnid groups and
525
the question of monophyly of the Acari, is
especially disjointed and difficult to follow,
as are many parts of Chapter 12, on repro-
duction. Statements out of context also may
be misleading. For example, the discussion
of the chemical “‘gyplure”’ (p. 328) implies
that it is a mite pheromone; no statement,
reference, or even implication informs the
reader that it is a gypsy moth sex phero-
mone. The quality of sentence structure is
inconsistent, with one or more examples of
poor syntax found on most pages. The result
may be simply awkward, or make no gram-
matical sense, but sometimes poor structure
makes sentences factually misleading.
Inconsistency in usage of names and spe-
cialized terminology is especially damaging
in an introductory text, and it is common
in this book. In Part 3 (Classification), the
cohort names Parasitiformes and Acari-
formes are used for the two principal lin-
eages, but the respective synonyms Anac-
tinotrichida and Actinotrichida are used in
most of the preceding text. The ordinal name
Actinedida is used in preference to its syn-
onyms, but the latter are commonly seen in
adjectival form (i.e. trombidiform mites,
prostigmatic mites). Examples of other
names and terms used interchangeably in-
clude: Acaridae/Tyroglyphidae, Histiosto-
matidae/Anoetidae, Dermatophagoides fa-
rinae/Tyroglyphus farinae, Sancassania/
Caloglyphus, Eriophyoidea/Tetrapodili,
Bimichaeloidea/Pachygnathoidea, Cla-
paréde’s organ/urstigma, segment/joint/ar-
ticle, eupathidion/acanthion, areae poro-
sae/porose areas, and subcapitulum/
infracapitulum. Sometimes terms are used
as synonyms when they clearly are not (e.g.
hysterosoma/notogaster, Ovoviviparity/Vvi-
viparity, integument/cuticle, trichobothria/
pseudostigmata). The author is also incon-
sistent in giving the taxonomic afhliations
of species used as examples; commonly these
names are not mentioned in sections on
classification, or even in the index, so a
reader must turn to other references. Some-
times terms are criticized, then incon-
526
gruously used with regularity. For example,
a paragraph (p. 132) is devoted to explaining
the inappropriateness of the term “podo-
cephalic canal’’, then the term is used con-
sistently in subsequent chapters. Similarly,
the term “‘opisthosoma” is said to be not
applicable to mites (p. 17), but it is used
without comment in later sections.
An abundance of contradictory state-
ments will frustrate the reader. For exam-
ple, on p. 18 the author implies an absence
of trichobothria from the legs of mites, then
discusses their presence on several later
pages. On p. 84 we learn that members of
the Astigmata do not possess a rutellum; on
p. 411 we learn that they do. The esopha-
gous is said to be of midgut origin, with no
cuticular lining (p. 136), then a cuticular
lining (implying ectodermal origin) is dis-
cussed (p. 141). On p. 199 the excretory
product guanine is said to be nontoxic, fol-
lowed by a discussion of how some mites
prevent it from reaching toxic level in the
hemolymph. Sometimes there are multiple
contradictions. The normal palp of mem-
bers of the Astigmata is variously described
or implied to be four-segmented (p. 84), two-
segmented (p. 96), or one-segmented (p.
411). On p. 391 we learn that actinedid mites
have no sejugal body division, on p. 393 we
learn that they do, and later (p. 394 and Fig.
19-2) we learn the truth by implication and
illustration; it may be present or absent.
Contradictions also occur with regard to
classification. On p. 230 Heterochthonius is
considered a member of the Cosmochtho-
noidea (sic); in the classification (p. 432) it
is in its own superfamily, the Heterochtho-
noidea (sic). On p. 367 Uropodellidae and
Ichthyostomatogasteridae are listed sepa-
rately; on p. 368 they are considered syn-
onyms. On p. 430 the Parhypochthonoidea
is included in the suborder Enarthronota;
on p. 433 of the same classification it com-
prises a separate suborder, the Parhypoch-
thonata.
There are relatively few typographical
errors, but they are concentrated in the sec-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tions on classification, where mispelled
names will not be obvious to many readers.
In some cases the errors introduce “new”
names or terminology, for example “‘di-
gophagous”’ for oligophagous (p. 145) or
“Bujobia” for Bryobia (p. 231).
An important part of a general text deal-
ing with an unfamiliar subject is a glos-
sary—or in its absence, a good index. The
author addresses the lack of a glossary, de-
ferring to the two volumes of a glossary of
mite terminology prepared by van der
Hammen (mentioned on p. viii, but without
specific citation until p. 320). But the latter
publications are expensive, and have a much
more restricted distribution than will this
book. Many specialized terms are not even
indexed, and some of them are first intro-
duced well before their definition, or else
lack definitions altogether. The index is also
difficult to use in places; as many as four
levels of indentation subdivide a main en-
try. In some cases, cited terms have abso-
lutely no relevance to the main entry; for
example, under the entry “rutellum” incon-
gruously are listed pages for “‘lyrifissure,”
‘“‘palpal-apoteles,”’ and ‘‘picket-fence-se-
tae.”’ Instances of incorrect page references
were also noted. As a test of the index, page
references to the mammalian follicle mites
were examined. Under the family name
“Demodicidae”’ two pages were listed, yet
the name is used in discussions of family-
level information on at least 12 pages, from
acursory examination of the text. Similarly,
under the species name “‘folliculorum” (one
of the two human follicle mites) three pages
are listed. In fact, there is no actual reference
to the species on one of these, and at least
five pages with such information are not
indexed, one of which (p. 303) contains most
of the biological information regarding this
species. Five other species of Demodex are
discussed at various places in the text, some
on multiple pages, but are not listed in the
index under the generic or specific names.
Curiously, the second species parasitizing
humans, D. brevis, is never mentioned.
VOLUME 90, NUMBER 4
There are no index entries under such log-
ical words as “‘follicle mite” or “Shuman fol-
licle mite’; in fact, the relationship of D.
folliculorum with humans is never men-
tioned in the text.
The book contains numerous factual
errors, of which only a small sample can be
mentioned here. Usually errors seem to be
introduced by the author, such as the as-
sociation of Tsutsugamushi disease with
ticks instead of chiggers (p. 386), or the par-
asitism of locusts by the honeybee mite (p.
41). Sometimes obviously incorrect state-
ments are taken intact from literature
sources—for example, the greatly inflated
numbers of solenidia on the legs of oribatid
mites (p. 430, taken from Johnston, 1982).
Misinterpretations of literature sources were
also noted. For example, a misreading of
the French apparently caused the author to
attribute to F. Grandjean (uncited, but cer-
tainly his 1964 paper; Acarologia 6: 170-
198) the erroneous idea that some oribatid
mites use their pteromorphs as gliding air-
foils. This was actually introduced as an un-
supported speculation by A. P. Jacot (an
uncited 1930 paper; Amer. Naturalist 64:
285-288) but is treated by the author as
observed fact (pp. 107, 238). Witalinski’s
cited (1979) paper was apparently the source
for a suggestion (p. 364) that peritremes of
gamasid mites function in hygroreception,
despite his clear statement that they lack
any association with sensory cells.
Errors also commonly appear in the form
of incorrect generalizations. Page 107 alone
has three such instances. The author gen-
eralizes that ““Phoretic mites use the legs to
clasp hairs or to hold to their transport
hosts”; in fact, chelicerae, sucker plates, anal
stalks, or other structures are more com-
monly used than legs, and some of these
mechanisms are discussed elsewhere in the
book. He states that “The first legs of Onb-
atids (sic) are tactile and carried above the
substrate”; in fact, this obtains for a small
minority of species. He also claims that
“among mites leaping is limited to a single
527
family of Oribatida” (the Zetorchestidae);
in fact, jumping is encountered in several
well-known families of Actinedida, as dis-
cussed in several papers which are cited
elsewhere in the text, as well as one or two
other families of oribatid mites. Some in-
correct generalizations are clearly more im-
portant, such as the statement that“. . . rel-
atively few mites consume particulate food”
(p. 146); in fact, particulate feeding is the
rule for two of the three major groups of
Acariformes (Oribatida, Astigmata) and for
early derivative members of the third (Ac-
tinedida), and is found as well in the Opi-
lioacarida.
Errors of omission can be expected in a
work of this magnitude, but some are sur-
prising. In a discussion of gamasid mor-
phology (p. 365), the author claims that no
“function is ascribed to the tritosternum
...’, but the excellent study of Wernz and
Krantz (Can. J. Zool. 54: 202-213; 1976),
clearly demonstrated its role in fluid distri-
bution during feeding.
Perhaps the most unfortunate shortcom-
ing of the book is its failure to invoke any
sense of wonder or challenge. Other than
suggesting how many undescribed species
there are, the author rarely makes a state-
ment which resembles an unanswered ques-
tion, a testable hypothesis, or even original
speculation. We know so little about the
most basic aspects of the biology of even
the most common species that questions
should flow freely in the text. Indeed, the
function of such a book should be as much
stimulation as elucidation. As one of earth’s
dominant arthropod groups, mites offer un-
limited opportunities for experimental study
in ecology, physiology, genetics, evolution,
and nearly every other subdiscipline of bi-
ology, opportunities which are nearly un-
tapped. A seminal, comprehensive synthe-
sis of our current knowledge of this group,
which could attract the attention of nonspe-
cialists and influence the direction of stu-
dents, has been conspicuously overdue, and
remains so.
528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
In summary, the book is not recom-_ ingful way. Factual errors are common, so
mended as a textbook of acarology, nor as__ original sources should always be consulted.
a source for comparative information on
mites, although the reference lists may be Roy A. Norton, S.U.N.Y. College of En-
useful. It seems carelessly written and ed- vironmental Science and Forestry, Syracuse,
ited, and generally fails to synthesize the New York 13210.
diversity of cited information in a mean-
PROC. ENTOMOL. SOC. WASH.
90(4), 1988, pp. 529-532
SocieTY MEETINGS
937th Regular Meeting—January 7, 1988
The 937th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Gene Wood in the
Naturalist Center, National Museum of
Natural History, at 8 p.m. on 7 January
1988. Because of a severe snowstorm, only
16 members and one guest were present.
Minutes of the previous meeting were read
and approved. No old business was trans-
acted, nor were there any applicants for
membership.
Mignon Davis noted that the staff of the
Naturalist Center would very much appre-
ciate donations of local natural history spec-
imens—including insects. She then dis-
played various insects that had been
embedded in plastic blocks; these are es-
pecially suitable for the sort of “hands on”
instruction offered at the Center.
President Wood announced that there
would be a meeting of the Executive Com-
mittee in the Museum of Natural History
at 10 a.m. on Monday, 8 February.
F. C. Thompson exhibited the Diptera
volume of the 1986 Zoological Record and
observed that this invaluable taxonomic
reference 1s once again being printed within
a year of the date that 1s indexed. Moreover,
the publishers have agreed to resume offer-
ing the Zoological Record on file cards, only
now the cards will be laser-printed.
The speaker for the evening was the Re-
cording Secretary, R. G. Robbins, whose
talk was entitled “Ticks: An Introduction
to the Ixodoidea and to the National Tick
Collection.” At the conclusion of his pre-
sentation, copies of the text Bloodsucking
Ticks (Ixodoidea)— Vectors of Diseases of
Man and Animals, by Yu. S. Balashov, were
distributed to the membership.
The sole guest was introduced and the
meeting was adjourned at 9:30 p.m., after
which refreshments were served.
Richard G. Robbins, Recording Secretary
938th Regular Meeting—February 4, 1988
The 938th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Gene Wood in the
Naturalist Center, National Museum of
Natural History, at 8 p.m. on 4 February
1988. Thirty members and 10 guests were
present. Minutes of the January meeting
were read and approved.
Membership Chairman G. B. White read
the names of the following applicants for
membership: Alfred P. Arthur, Agriculture
Canada, Saskatoon, Saskatchewan; Art Bor-
kent, Biosystematics Research Centre, Ot-
tawa, Ontario, Canada; Hans K. Loechelt,
Tieton, Washington; Stephen R. Moulton
II, Lusby, Maryland; and Robin J. Rath-
man, Tree Fruit Research Center, Wenatch-
ee, Washington.
R. G. Robbins exhibited a picture of an
amblyommine tick crafted from peacock
feathers for the late Harry Hoogstraal. This
and other acarological objets d’art were
Christmas gifts from Dr. Hoogstraal’s Egyp-
tian colleague Sherif Tewfik.
W.E. Bickley announced the death on 26
February of George S. Langford, former
Maryland State Entomologist. Dr. Langford
was the oldest active member of this Soci-
ety, having paid dues since 1924. He is
widely remembered for his leadership in a
program aimed at controlling the Japanese
beetle (Popillia japonica) through dissem1-
nation of milky white disease.
J. M. Kingsolver announced the recent
death of Lewis J. Stannard, an active ESW
member since 1948. Dr. Stannard worked
530
for the Illinois Natural History Survey where
he specialized in thrips and mites.
The speaker for the evening was Thomas
E. Wallenmaier, Program Planning Staff,
U.S. Department of Agriculture, APHIS-
PPQ. His much anticipated talk, “A New
Theoretical Foundation for Systematics,”
advocated the primacy of genetic characters
in evolutionary analysis.
President Wood reminded Society offi-
cers that the meeting of the Executive Com-
mittee previously scheduled for 8 February
had been postponed to the 11th at 2 p.m.
Guests were introduced and the meeting
was adjourned at 9:30 p.m. Refreshments
followed.
Richard G. Robbins, Recording Secretary
939th Regular Meeting— March 3, 1988
The 939th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Gene Wood in the
Naturalist Center, National Museum of
Natural History, at 8 p.m. on 3 March 1988.
Twenty-five members and six guests were
present. Minutes of the February meeting
were read and approved.
Membership Chairman G. B. White read
the names of the following applicants for
membership: Robert M. Eisenberg and
Lawrence E. Hurd, both at the School of
Life Sciences, University of Delaware,
Newark; Edward A. Lisowski, Illinois Nat-
ural History Survey, Champaign; Terry D.
Miller, Moscow, Idaho; Roy A. Norton,
SUNY College of Environmental Science
and Forestry, Syracuse; John D. Sedlacek,
Department of Entomology, University of
Kentucky, Lexington; and Gary J. Steck,
Laurel, Maryland.
R. G. Robbins read the minutes of the
Executive Committee meeting held at the
National Museum of Natural History on 8
February between 2 and 3:45 p.m. Among
the items discussed were obituaries for
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
C.F.W. Muesebeck and Frederick W. Poos;
possible increases in membership dues and
subscription rates for 1989; future reduc-
tions in the net press run of the Proceedings:
committee review of manuscripts submit-
ted for publication as Memoirs; procedures
to be followed when members request
Emeritus status; and the location of this
year’s banquet.
President Wood explained that during the
Executive Committee’s meeting a motion
had been submitted by M. B. Stoetzel rec-
ommending that Honorary Member Curtis
W. Sabrosky be nominated Honorary Pres-
ident. Also during this meeting, T. E. Wal-
lenmaier had moved that Louise M. Russell
be nominated to fill the position of Hon-
orary Member that would be vacated if Dr.
Sabrosky were elected Honorary President.
Both motions were unanimously passed by
the Executive Committee. President Wood
now placed these nominations before the
membership, which approved them by ac-
clamation.
E. S. Saugstad displayed photoenlarge-
ments of the epaulets worn by officers in the
army of the Ivory Coast. These insignia are
entomologically remarkable in that they are
ornamented with the likenesses of several
kinds of African insects.
J. H. Fales reported that his paper, “The
Butterflies of Rock Creek Park, Washing-
ton, D.C.,” has appeared in the revived
Maryland Naturalist (31: 5-24), published
by the Natural History Society of Maryland.
Fales’ 45 years of collecting in Rock Creek
Park have yielded 58 of the 97 species of
butterflies and skippers known from the
District of Columbia. Photocopies of this
valuable reference were distributed to the
membership.
W.E. Bickley announced that Alan Stone,
a Life Member of this Society, has been hon-
ored with the John N. Belkin Award of the
American Mosquito Control Association in
recognition of Dr. Stone’s many outstand-
ing contributions to mosquito systematics.
R. G. Robbins exhibited some of the
VOLUME 90, NUMBER 4
hundreds of metal plates used to illustrate
the acarological papers of the late Harry
Hoogstraal. Noteworthy examples include
a zinc plate depicting the distribution of
Haemaphysalis longicornis in the western
Pacific and a copper halftone plate showing
St. Mary’s Church in Krakow, Poland, the
type locality of Argas polonicus.
The speaker for the evening was Chris-
topher K. Starr, Postdoctoral Fellow,
Smithsonian Institution, whose talk was en-
titled ““Getting Around the Archipelago: The
Distribution of Bugs and a Bug-Watcher in
Southeast Asia.” During his presentation,
Dr. Starr distributed handouts showing the
zoogeographic tracks of various genera of
pachyrrhynchine weevils and stenogastrine
wasps in the Malay Archipelago and adja-
cent areas. He also exhibited several insect
specimens that he had collected in South-
east Asia and provided the membership with
a list of references to the natural history of
this region.
Visitors were introduced and the meeting
was adjourned at 9:30 p.m. Refreshments
followed.
Richard G. Robbins, Recording Secretary
940th Regular Meeting—April 7, 1988
The 940th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Gene Wood in the
Naturalist Center, National Museum of
Natural History, at 8:08 p.m. on 7 April
1988. Twenty-eight members and six guests
were present. Minutes of the March meeting
were read and approved.
Membership Chairman G. B. White read
the names of the following applicants for
membership: Christopher H. Dietrich, De-
partment of Entomology, North Carolina
State University, Raleigh; and John D.
Glaser, Baltimore, Maryland.
President-Elect F. C. Thompson an-
nounced that the Society’s annual banquet
531
will be held on Monday, 6 June, at the Uni-
versity of Maryland’s Center of Adult Ed-
ucation. The guest speaker will be James M.
Carpenter, Department of Entomology,
Museum of Comparative Zoology, Harvard
University. The title of his presentation will
be “Testing Scenarios: Wasp Social Behav-
10K:
President Wood called for a round of ap-
plause for new Honorary Member Louise
M. Russell and new Honorary President
Curtis W. Sabrosky. Dr. Sabrosky was pres-
ent and warmly thanked the membership.
C. K. Starr reported on three species of
Philippine jumping spiders (Salticidae), each
of which apparently is a species-specific
mimic of a different otiorrhynchine weevil.
Dr. Starr illustrated this relationship with
slides that he had taken of one of the spiders
and its model.
R. G. Robbins exhibited a set of three
Malaysian stamps commemorating the In-
stitute for Medical Research, Kuala Lum-
pur. One of these stamps features a well-
executed drawing of a chigger belonging to
the subgenus Leptotrombidium, which in-
cludes all known vectors of scrub typhus
(Rickettsia tsutsugamushi).
T. J. Spilman observed that a recent U.S.
commemorative stamp honoring the State
of Massachusetts includes the minute figure
of a grasshopper atop the weather vane on
Boston’s Faneuil Hall. He also distributed
an assortment of surplus entomological
publications.
Mignon Davis again asked that members
and their friends contribute local natural
history specimens to the Naturalist Center.
Insects are an especially important com-
ponent of several courses offered at the Cen-
ter.
The speaker for the evening was Gary F.
Hevel, Collections Manager, Department of
Entomology, Smithsonian Institution. His
talk, entitled ““Durians and Trilobite Bee-
tles: Collecting Experiences in Sabah,” re-
counted his entomological, botanical and
cultural adventures with teammate Warren
532 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
E. Steiner, Jr., during their ascent of Kin- Visitors were introduced and the meeting
abalu, the highest mountain on the island was adjourned at 9:45 p.m. Refreshments
of Borneo (indeed, in all Southeast Asia). followed.
Misilevcls pueseutation was accompanied Richard G. Robbins, Recording Secretary
by some unusual sound effects provided by
the President-Elect.
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Cynipid Galls of the Eastern United States, by Lewis H. Weld icc
Gynipid:Gallsiofithe: Southwest: byWewis H. Weld 25s Se ee
Bothipapers Onicynipidsealls =e rw sae tk ee ee ee ee
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman eee eeceeeee eee
Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R.
Traver.
A Short History of the Entomological Society of Washington, by Ashley B. Gurney...
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C.
Ste ys kcal newt cee coe Fees Ae ee Re OE ea ee ee ee
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
Steyskal
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939.
No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
P/E Faye ICY eS Ne ha AN eon a ee ag eM Oe eas ne a ed a ee De eee ES
No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
PISS 0) 00 Es CERT Ee ne es ae a oe Se I ERE Se SNP TE A Ds
No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952.00
No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp.
SG face tN PS i eae ame ea i Aa Nd ae eee ee ee
No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
LLNS EE AS UD) Yo) onl PLoS ahaa aD ie o me ece ieaset eRe ENN tetera ee a
No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
Oi Geers ee ee Ew Ee Ie Be A one 2 Oe eee Se De tS ae AOE Bi eS
No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cera-
topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979. eeseeeeeeeeee eee
No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200
DD UGS less Mise Nother ase Mee ee NINE Be Ee ee a Ts ee ee Oe
No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982...
No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp.
OR Seka eee BNR CN ay Sa ek ee re re ee aS Ne ee
No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
SETS ol OR Otero Neca Sd I UN ee Oe REN ce SY Se ae ey
No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hy-
menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987 _.... #5
$ 5.00
3.00
6.00
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Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume
to non-members and $13.00 per volume to members of the Society.
Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10
per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with
the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
WILKERSON, R. C.—Notes and redescriptions of some Anopheles series Arribalzagia holotypes
(Diptera: Culicidae) in the British Museum (Natural History)
NOTES
NAKAHARA, S.—A new synonym and revised status in Apterothrips (Thysanoptera: Thripidae) 508
RATHMAN, R. J., J. F. BRUNNER and S. J. HULBERT—Feeding by Medetera species
(Diptera: Dolichopodidae) on aphids and eriophyid mites on apple, Malus domestica
(Rosaceae)
OBITUARY
KROMBEIN, K. V. and P. M. MARSH—Carl Frederick William Muesebeck, 1894-1987 ... 513
BOOK REVIEW
NORTON, R. A.—Acarology. Mites and Human Welfare
SOCIETY MEETINGS
INDEX OF VOLUME 90 (1988) CONTENTS WILL APPEAR IN VOLUME 91(1)
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