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VOL. 89 


NO. 1 

aSSN 0013-8797) 


of the 




J 8 84, 


AGNEW, C. W. — Status of Acigona Hubner (sensu Bleszynski) (Lepidopjjefa: TE*tralidag:^-j^ 

binae) with changes in nomenclature 100 

Crossing and chromosomal evidence for two additional sibling species within the taxon 
Anopheles dims Peyton and Harrison (Diptera: Culicidae) in Thailand 157 

BELLINGER, R. G. and R. L. PIENKOWSKI-Life history observations on the grasshopper 

Appalachia hebardi Rehn and Rehn (Orthoptera: Acrididae: Melanoplinae) 43 

CLARKE, J. F. G.— The correct identity of Acleris inana (Robinson) (Lepidoptera: Tortricidae) 175 

DIATLOFF, G. and W. A. PALMER— The host specificity of Neolasioptera lathami Gagne 

(Diptera: Cecidomyiidae) with notes on its biology and phenology 122 

HARRIS, S. C— A new species of Agarodes (Trichoptera: Sericostomatidae) from southeastern 

United States 74 

HOFFMAN, K. M. — Earwigs (Dermaptera) of South Carolina, with a key to the eastern North 

American species and a checklist of the North American fauna 1 

KONDRATIEFF, B. C. and R. F. KIRCHNER-Additions, taxonomic corrections, and faunal 

affinities of the stoneflies (Plecoptera) of Virginia, USA 24 

LOWRY, J. E. andH. D. BLOCKER— Two new species of Flexamia from the Nebraska Sand 

Hills (Homoptera: Cicadellidae: Deltocephalinae) 57 

MARI MUTT, J. A.— A new Seira from the United States, with a redescription of Seira bi- 
punctata (Packard) and new records for Seira distincta Man Mutt (Collembola: Entomobry- 
idae) 126 

NELSON, C. R. and R. W. BAUMANN— Scanning electron microscopy for the study of the 

winter stonefly genus Capnia (Plecoptera: Capniidae) 51 

PALMER, W. A.— The phytophagous insect fauna associated with Baccharis halimifolia L. and 

B. neglecta Britton in Texas, Louisiana, and northern Mexico 185 

{Continued on back cover) 




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Title of Publication: Proceedings of the Entomological Society of Washington. 
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89(1), 1987, pp. 1-14 




Kevin M. Hoffman 

Department of Entomology, Clemson University, Clemson, South Carolina 29634- 

Abstract. —Biological and distributional data are given for the nine species of Dermap- 
tera recorded from South Carolina. A key to the 1 7 eastern North American species is 
presented along with brief notes on the taxonomy and distribution of all 22 species 
occurring in North America north of Mexico. 

The most recent comprehensive checklist 
of North American Dermaptera was pub- 
lished over 35 years ago (Gumey, 1950), 
and the last key to the eastern North Amer- 
ican species was published by Blatchley 
(1 920). In the intervening years new species 
have been introduced, and the taxonomic 
status of others has changed. This paper 
documents the species occurring in South 
Carolina, presents a revised checklist for the 
species occurring in North America north 
of Mexico, and provides a key to eastern 
North American species. A similar study of 
the Dermaptera of California provided a key 
to most of the known western species 
(Langston and Powell, 1975). 

The order Dermaptera comprises some 
1200 species worldwide, occurring mostly 
in the tropics. Only 22 species are currently 
known from North America, and at least 
twelve of these were introduced from Eu- 
rope and the tropics. Seventeen species have 
been recorded from eastern North America, 
nine of which have been found in South 
Carolina. These nine are the most wide- 
spread species in the east; the other eight 
species are restricted to Florida and New 
Jersey. The remaining five North American 
species occur only in western United States. 

A brief summary of the North American 
records for non-South Carolina species fol- 

There are eight eastern species not found 
in South Carolina. Pyragropsis buscki (Cau- 
dell) is a Caribbean species that has become 
established in the Miami, Florida area (Gur- 
ney, 1959). Euborellia ambigua (Borelli) was 
originally described from Costa Rica and 
has since been recorded from a mangrove 
swamp near Miami, Florida (Hebard, 1922). 
Euborellia caraibea Hebard, a species that 
is widespread in the Caribbean Islands, has 
been collected in sandy areas on Virginia 
Key, Rorida (Brindle, 197 Id). The char- 
acters originally used by Hebard (1922) to 
differentiate E. caraibea from E. ambigua 
have shown considerable variation when a 
larger series of E. caraibea was examined, 
making it difficult to distinguish the two 
species (Brindle, 197 Id). This difficulty is 
reflected in the following key and awaits the 
collection of a larger series of E. ambigua 
before it can be resolved. Euborellia an- 
nulata (P.), a native of Indonesia, was re- 
corded by Townes (1946) from Miami, 
Florida (as Euborellia stali [Dohrn, 1864a], 
synonymized by Brindle, 1981). Specimens 
of this species are in the Florida State Col- 


lection of Arthropods from as far north as 
Daytona, Florida. 

Labia cunicauda (Motschulsky), origi- 
nally described from Ceylon, has been found 
on Long Key, Florida, occupying the bases 
of leaves in the tops of coconut palms (Rehn 
and Hebard, 1912). Labia rehni Hebard is 
known only from a single female found be- 
tween boards in a woodshed on Key West, 
Florida, on 7 July 1912 (Hebard, 1917; 
Brindle, 1971a). Doru davisi Rehn and He- 
bard is found only in southern Florida, es- 
pecially around Lake Okeechobee, and is 
usually associated with saw-grass (Gumey, 
1972). Marava arachidis (Yersin) was de- 
scribed from southern Europe and has sub- 
sequently been introduced into southern 
Florida (Caudell, 1913). This species has 
been recorded from two localities in New 
Jersey by Townes (1946) (as Marava wal- 
lacei [Dohrn, 1864b], synonymized by 
Boeseman, 1954). It has also been recorded 
from Texas, Arizona, and California (He- 
bard, 1943; Langston and Powell, 1975). 

Five species occur only in western United 
States. Chelisoches morio (F.) is a south Pa- 
cific species that has been introduced into 
Cahfomia (Langston and Powell, 1975). 
Euborellia cincticollis (Gerstaecker) is an 
African species that is now established in 
California and Arizona (Langston and Pow- 
ell, 1975). Euborellia femoralis (Dohrn), an 
Oriental species that is very similar to E. 
annulipes, has been recorded from Arizona 
and California (Steinmann, 1981). Vostox 
apicedentatus (Caudell) is a native species 
that has been recorded from California, Ar- 
izona, New Mexico, and Texas. It was listed 
in the genus Spongovostox by Langston and 
Powell (1975) but has been transferred to 
the genus Vostox by Brindle (1971b). Vos- 
tox excavatus Nutting and Gumey is known 
only from Arizona, New Mexico, and 
northwestern Mexico (Nutting and Gumey, 

Labia pilicornis (Motschulsky, 1863) was 
listed by Gumey ( 1 9 50) as being established 
in the United States, but no North Ameri- 

can collection records were cited. A foot- 
note associated with the listing stated that 
Morgan Hebard considered Labia rehni to 
be a junior synonym of L. pilicornis, so per- 
haps it was included in the list in antici- 
pation of this proposed synonymy. Because 
subsequent workers have maintained the 
two species as separate and no published 
Nearctic records have been found, it is 
doubtful that L. pilicornis occurs in North 
America and thus is omitted from the 
checklist below. In the Neotropical region, 
L. pilicornis is known only from Cuba (Brin- 
dle, 197 Id). 

Earwigs exhibit sexual dimorphism in 
both the number of abdominal segments and 
the shape of the forceps. Males have ten 
abdominal segments, whereas females have 
only eight apparent segments due to the fu- 
sion of the eighth and ninth segments with 
the tenth (Giles, 1 963). The forceps of males 
are generally curved and widely separated 
at the base, and many species have one or 
more prominent teeth on the inner margins 
(Figs. 1 1-25). Female forceps are more or 
less straight, usually closer together basally 
than those of the male and lack prominent 
teeth on their inner margins (Figs. 26-31). 
Both sexes possess a divided plate called the 
pygidium, which is located between the bas- 
es of the forceps (Fig. 18). The pygidium is 
thought to be derived from the epiproct and 
is useful taxonomically, especially in males. 

Immature earwigs undergo five to six in- 
stars before reaching adulthood. Antennal 
segments are added at each molt, and wing 
pads begin to develop in the second or third 
instar. Immatures are lighter in color than 
adults and possess conspicuous ecdysial lines 
on the head and thoracic terga. Immatures 
have male-like, ten-segmented abdomens, 
but have female-like, straight forceps. 

Material for this study was examined from 
the following institutions: North Carolina 
State University, University of Georgia, 
Florida State Collection of Arthropods, Na- 
tional Museum of Natural History, and 
Clemson University. The body length of 


specimens was measured from the head to 
the tip of the forceps. A brief diagnosis and 
information on the synonymy, bionomics, 
world distribution, and South Carolina dis- 
tribution are given for those species record- 
ed from South Carolina. The synonymy giv- 
en for each South Carolina species is 
complete unless otherwise noted, in which 
case synonyms commonly used in previous 
literature on Nearctic species are provided, 
and a reference is cited which contains a 
more complete synonymy. South Carolina 
county records for each species include every 
county from which the species has been col- 
lected, but not necessarily every record from 
those counties. The known North American 
distribution of each species is included in 
the following checklist, with states desig- 
nated by the two-letter codens assigned by 
the United States Postal Service. The rec- 
ords used in compiling the checklist are con- 
sidered to represent established popula- 
tions, although it is possible that some may 
represent transient infestations that have 
since disappeared. The classification system 
used in the checklist is that of Popham 
(1965a, b). 

Checklist of the North American 


Superfamily Pygidicranoidea 
Family Pygidicranidae 
Subfamily Pygidicraninae 
Pyragropsis buscki (Caudell, 1907); 
Superfamily Labioidea 
Family Carcinophoridae 
Subfamily Carcinophorinae 
Anisolabis maritima (Bonelli, 1832); 

widespread on sea coasts. 
Euborellia ambigim (Borelli, 1906); 

Euborellia annulata (F., 1793); FL. 
Euborellia annulipes (Lucas, 1847); 

Euborellia caraibea Hebard, 1922; 

Euborellia cincticollis (Gerstaecker 

1883); AZ, CA. 
Euborellia femoralis (Dohm, 1863) 
AZ, CA. 
Family Labiidae 
Subfamily Labiinae 

Labia curvicauda (Motschulsky 

1863); FL. 
Labia minor {h., 1758); widespread 
Labia rehni Hebard, 1917; FL. 
Subfamily Spongiphorinae 
Marava arachidis (Yersin, 1 860); AZ 

CA, TX, NJ, FL. 
Marava pulchella (Audinet-Serville 
1839); southeastern U.S., TX. 
Vostox apicedentatus (Caudell, 1 905) 

AZ, CA, NM, TX. 
Vostox brunneipennis (Audinet-Ser- 
ville, 1839); eastern U.S., TX 
Vostox excavatus Nutting and Gur- 
ney, 1961; AZ, NM. 
Superfamily Forficuloidea 
Family Labiduridae 
Subfamily Labidurinae 

Labidura riparia (Pallas, 1773) 
southeastern U.S., AZ, CA, TX 
Family Chelisochidae 
Subfamily Chelisochinae 
Chelisoches morio (F., 1775); CA. 
Family Forficulidae 
Subfamily Forficulinae 

Doru aculeatum (Scudder, 1876) 

eastern U.S., Ontario. 
Doru davisi Rehn and Hebard, 1914: 

Doru taeniatum (Dohrn, 1862) 
southeastern U.S., AZ, CA, TX. 
Forficula auricularia L., 1758; wide- 

Key to Adults of the Eastern North 
American Species of Dermaptera 

L Tegminae absent or present only as 
rounded flaps that do not meet at the inner 
basal margins (Figs. 1 and 2); right forceps 






Figs. 1-10. Thoracic terga, metatarsi, and antennal bases of Dermaptera. 1, Euborellia annulata, thoracic 
terga, dorsal view. 2, E. ambigim. thoracic terga (redrawn from Hebard, 1922), dorsal view. 3, Marava pulchella, 
pronotum and wings of fully winged form, dorsal view. 4, M. arachidis, pronotum (redrawn from Brindle, 197 1), 
dorsal view. 5, Doru aculeatum. right metatarsus, lateral view. 6, D. aculeatum, right metatarsus, dorsal view. 
7, M. pulchella, right metatarsus, lateral view. 8, M. pulchella, right metatarsus, dorsal view. 9, Forficula 
auricularia, base of right antenna, dorsal view. 10, D. aculeatum, base of right antenna, dorsal view. a4 = fourth 
antennal segment, cl = tarsal claw, hw = hindwing, ms = mesonotum, mt = metanotum, pr = pronotum, te = 
tegmina, ts 1 = first tarsal segment, ts2 = second tarsal segment, ts3 = third tarsal segment. 

of male curved inward more strongly than 
left (Figs. 1 1-12) (Carcinophoridae) .... 2 
Tegminae normally developed and meet- 
ing along entire midline (Fig. 3); forceps 

of male symmetrical (Figs. 14-25) 6 4(3). 

2( 1 ). Tegminae present as rounded lateral flaps 

on mesonotum (Figs. 1 and 2); Florida . 3 

- Tegminae absent; widespread 5 

3(2). Tegminal flaps widely separated on meso- 
notum (Fig. 1); legs with brown rings 
around femora and tibiae; left forceps of 

male curved only at apex (Fig. 12) 

Euborellia annulata (F.) 5(2). 

Tegminal flaps covering most of meso- 
notum (Fig. 2); legs unicolorous or femora 
darkened; left forceps of male curved al- 
most as much as right (Fig. 13) 4 

Legs uniformly pale; tegminal flaps not 
always meeting at midline (Fig. 2); found 
only in mainland mangrove swamps . . . 

Euborellia ambigua (Borelli) 

Legs pale or with femora darkened; teg- 
minal flaps always meeting or overlapping 
at midline; found in various habitats on 

islands Euborellia caraibea Hebard 

Antennae entirely dark brown, 20-24 seg- 


:— a10 



Figs 1 1-18. Male forceps of Dermaptera, dorsal views. 11, Anisolabis maritima. 12, Euborellia annulipes. 
13 E awZ)/^/a (redrawn from Hebard, 1922). U, Labia minor. 1 5, L. CMmcflM^a (redrawn from Bnndle, 1971a). 
le! Marava arachidis (redrawn from Brindle, 1971b). 17, M. pulchella. 18, Vostox bmnneipenms. alO 
abdominal segment, fc = forceps, pg = pygidium. 



Figs. 19-25. Male forceps of Dermaptera, dorsal views. 19, Dom taeniatum. 20, D. aculeatum. 21, D. davisi 
(redrawn from Gumey, 1972). 22, Pyragropsis buscki. arcuate form (redrawn from Gumey, 1959). 23, P. buscki, 
elongate form (redrawn from Gumey, 1959). 24, Labidura riparia. 25, Forficula auhculaha. 

mented; femora and tibiae without encir- 
cling dark bands; body length 20-25 mm 

Anisolabis mahtima (Bonelli) 

Antennae brown with 1-3 white subapical 
segments, 14-18 segmented; femora and 


tibiae pale yellow with encircling dark 

bands; body length 12-18 mm 

Euborellia annulipes (Lucas) 

Second tarsal segments cylindrical and not 
wider than third, at most only slightly ex- 


Figs. 26-31. Female forceps of Dermaptera, dorsal views. 26, Euborellia annulipes. 27, Labia minor. 28, L. 
rehni (redrawn from Hebard, 1917). 29, Marava pulchella. 30, Vostox brunneipennis. 31, Doru aculeatum. 

tended beneath third (Figs. 7 and 8) . . . 7 
Second tarsal segments dilated and much 
wider than third, extending conspicuously 
beneath third (Figs. 5 and 6) (Forficulidae) 

7(6). Large pad-like arolium between tarsal 
claws; hindwings visible; body length 14- 
1 9 mm; male forceps of two types, both 
forms curved strongly inward (Figs. 22 

and 23) Pyragropsis buscki (Caudell) 

No arolium between tarsal claws; hindw- 
ings visible or not; body length variable; 
male forceps not strongly incurved (Figs. 
14-18) 8 

8(7). Antennae 25-30 segmented; pronotum 
light brown with two dark longitudinal 
stripes; body length 20-30 mm; male py- 
gidium reduced and not visible in dorsal 
view (Fig. 24) .... Lahidura riparia (Pallas) 
Antennae 10-16 segmented; pronotum 
unicolorous; body length less than 20 mm; 

male pygidium prominent (Figs. 14-18) 
(Labiidae) 9 

9(8). Tegminae pubescent; body length less than 

8 mm 10 

Tegminae glabrous; body length 8-18 
mm 12 

10(9). Tegminae as long as pronotum; visible 
portion of hindwings only half length of 
pronotum; inner margin of female forceps 

notched basally (Fig. 28) 

Labia rehni Hebard 

Tegminae much longer than pronotum; 
visible portion of hindwings as long as 
pronotum; inner margin of female forceps 
not notched basally (Fig. 27) 11 

11(10). Head and thorax black; abdomen reddish; 
male pygidium quadrate; inner margin of 

male forceps curved (Fig. 15) 

Labia curvicauda (Motschulsky) 

Body yellowish-brown to brown; male py- 
gidium elongated medially; inner margin 


of male forceps straight (Fig. 14) 

Labia minor (L.) 

12(9). Male pygidium large, produced medially 
(Fig. 18); hindwings always present; an- 
tennae entirely brown; body length 12-16 

mm Vostox brunneipennis 

Male pygidium not produced medially 
(Figs. 16-17); hindwings often absent or 
concealed; basal 2-3 antennal segments 
yellow, remainder brown; body length 8- 
12mm 13 

13(12). Pronotum broader than long (Fig. 4); male 
pygidium pentagonal; male forceps with- 
out inner basal tooth (Fig. 16) 

Marava arachidis (Yersin) 

Pronotum as broad as long (Fig. 3); male 
pygidium quadrate; male forceps with in- 
ner basal tooth (Fig. 17).. Marava pulchella 

14(6). Fourth antennal segment less than twice 
as long as broad (Fig. 9); tegminae dark 
brown; male pygidium truncate; male for- 
ceps broadened, almost touching basally 
(Fig. 25) Forficula auricularia L. 

- Fourth antennal segment more than twice 
as long as broad (Fig. 1 0); tegminae yellow 
with brown inner margins; male pygi- 
dium spine-like; male forceps widely sep- 
arated at base (Figs. 19-21) 15 

15(14). Hindwings visible beyond tegminae ... 

Doru taeniatum (Dohm) 

Hindwings not visible beyond tegminae 1 6 

16(15). Male forceps with tooth near apex (Fig. 
20); male pygidium distinctly shorter than 

length of last abdominal segment 

Doru aculeatum (Scudder) 

Male forceps without tooth near apex (Fig. 
21); male pygidium as long as length of 

last abdominal segment Doru davisi 

Rehn and Hebard 

Dermaptera of South Carolina 

Anisolabis maritima (Bonelli, 1832) 

Fig. 1 1 

Forficula maritima Bonelli, 1832, in Gene, 
Ann. Sci. Regn. Lomb.-Venet. 2: 224. 

Anisolabis maritima (Bonelli) Fieber, 1853, 
Lotos 3: 257. 

Steinmann (1984) gives a more complete 

Anisolabis maritima can be distinguished 
from the other wingless species occurring in 

South Carolina by the 20-24 segmented an- 
tennae, lack of dark encircling bands on the 
legs, and 20-25 mm body length. This 
species has been cited as A. maritima (Gene, 
1832) in earlier papers. Anisolabis maritima 
is usually found underneath litter and drift- 
wood along seashores (Langston, 1 974). This 
earwig forages at night and is predaceous; 
its prey includes crickets, sandfleas, and 
smaller earwigs (Bennett, 1 904). 

This species is essentially cosmopolitan. 
Introduced into North America, A. mari- 
tima now occurs locally along the Pacific 
coast from British Columbia south to Cal- 
ifornia, and along the Atlantic and Gulf 
coasts from Maine to Florida and westward 
to Texas (Langston and Powell, 1975). 

South Carolina Records.— Anisolabis 
maritima probably occurs locally along the 
entire South Carolina coast. Beaufort Co.: 
Beaufort, 14 June 1930, coll. O. L. Cart- 
wright, 1 9; Bluffton, 2 April 1933, coll. D. 
Dunavan, 1 3, 2 9, 2 immatures. Horry Co.: 
30 August 1981, coll. S. Mudge, 1 6. Pickens 
Co.: Clemson, 23 June 1980, under board 
in bam, coll. C. Lay, 19. 

Euborellia annulipes (Lucas, 1847) 

The Ringlegged Earwig 
Figs. 12, 26 

Forficesila annulipes Lucas, 1847, Ann. Soc. 
Entomol. Fr. 5: 84. 

Anisolabis annulicornis Blanchard, 1 85 1 , /« 
Gay, Hist. Fisica Pol. Chile, Zool. 6: 10. 

Euborellia annulipes (Lucas) Burr, 1915, J. 
R. Microsc. Soc. 35: 545. 

Sakai (1970a) gives a more complete syn- 

The ringlegged earwig can be distin- 
guished from the only other wingless species 
in South Carolina by its 14-18 segmented 
antennae, dark encircling bands on the fem- 
ora and tibiae, and 12-18 mm body length. 
The dark brown antennae generally have the 
third, fourth, and sometimes fifth subapical 
segments pale yellow to white, although a 
few specimens show only one or no pale 


Euborellia annulipes is a general scav- 
enger that is commonly found on the ground 
underneath rocks, logs, and other debris. 
This earwig can cause minor damage to 
plants and stored foods when it enters 
greenhouses, nurseries, and warehouses 
(Bharadwaj, 1966). However, it is also pre- 
daceous and will attack other insect pests. 
The ringlegged earwig is an introduced 
species that occurs worldwide. It has been 
established in North America for at least 
1 40 years and now occurs virtually through- 
out the United States and into British Co- 
lumbia (Langston and Powell, 1975; Scud- 
der and Vickery, 1985). 

South Carolina Records.— The ringlegged 
earwig is the most commonly encountered 
species in the state. Aiken Co.: Windsor, 25 
November 1933, coll. O. L. Cartwright, 1 
9. Anderson Co.: Pendleton, 31 October 
1975, in cotton trash, coll. P. Zinsmelster, 
1 5, 1 2, 2 immatures. Barnwell Co.: Elko, 
30 July 1981, debris under pig feeding 
trough, coll. J. R. Brushwein, 1 3, 2 9, 1 
immature. Beaufort Co.: Beaufort, 30 Oc- 
tober 1925, under trash, coll. F. Sherman, 
1 immature. Charleston Co.: Charleston, 5 
May 1951, coll. D. Dunavan, 1 9. Cherokee 
Co.: Gaffney, 15 August 1939, coll. L. M. 
Sparks, 1 9. Clarendon Co.: Summerton, 31 
March 1929, coll. O. L. Cartwright, 1 im- 
mature. Colleton Co.: Bear Island, 30 Sep- 
tember 1979, grass, coll. B. Hendrix, 1 9. 
Darlington Co.: Darlington, 19 June 1929, 
coll. J. M. Napier, 1 9. Dorchester Co.: Sum- 
merville, 15 August 1931, F. Sherman, 1 6. 
Edgefield Co.: Trenton, 1 9 November 1959, 
trunk of peach tree, coll. T. E. Skelton, 1 
immature. Florence Co.: Florence, 17 July 
1981, leaf litter, coll. R. D. Simpson, 2 9. 
Lee Co.: Meredith, 22 June 1926, coll. O. 
L. Cartwright, 1 3, 1 9. Oconee Co.: Fairplay, 
18 October 1984, under feathers in chicken 
house, coll. W. Barton, 2 immatures. 
Orangeburg Co.: Orangeburg, 22 July 1927, 
coll. F. Sherman, 1 9. Pickens Co.: Clemson, 
1 December 1983, beneath rock in garden, 
coll. J. Joly, 1 9, 2 immatures. Richland Co.: 
Columbia, 21 February 1926, O. Cart- 

wright, 1 9. Spartanburg Co.: Greer, 31 Oc- 
tober 1976, coll. M. McClimon, 1 9. 

Labia minor (L., 1758) 

Figs. 14, 27 

Forficula mmor Linnaeus, 1758, Syst. Nat. 
(10) 1:423, no. 2. 

Labia minor (L.) Stephens, 1835, 111. Brit. 
Entomol., Mand. 6: 8. 

Labia minuta Scudder, 1 862, Boston J. Nat. 
Hist. 7: 415. 

Sakai (1970b) gives a more complete syn- 

Labia minor is distinguished from all oth- 
er earwigs in South Carolina by both the 
abundance of golden hair covering most of 
its body and its small body size (less than 
8 mm long). It resembles a small rove beetle 
(Staphylinidae) and as such is often put into 
unsorted collections of these beetles. Labia 
minor is both a scavenger and a predator 
and can be found under various kinds of 
debris. It flies readily and is frequently at- 
tracted to lights. Labia minor is found in 
temperate and subtropical zones worldwide 
and is another species that has been intro- 
duced into the Nearctic region. Like E. an- 
nulipes, it is now distributed throughout 
North America (Langston and Powell, 1975; 
Scudder and Vickery, 1985). 

South Carolina Records.— La^/« minor is 
presently known only from the extreme 
northwestern part of the state. It may well 
occur in other parts of the state, being com- 
monly overlooked because of its small size. 
Anderson Co.: Pendleton, 28 July 1972, in 
cottonseed, coll. R. P. Griffin, 1 9. Pickens 
Co.: Clemson, 23 June 1956, in light trap, 
coll. D. Dunavan, 1 3; Clemson, 19 Septem- 
ber 1985, in cattle feed at Clemson Univ. 
dairy bam, coll. K. M. Hoffman, 1 imma- 
ture; Clemson, 5 October 1984, flying in 
field, coll. J. Barron, 1 6; Clemson, 20 April 
1939, at light, coll. F. T. Arnold, 1 6. 

Marava pulchella (Audinet-Serville, 1839) 

Figs. 3, 7,8, 17,29 

Forficula pulchella Audinet-Serville, 1839, 
Hist. Nat. Ins. Orthop. p. 42. 



Labia guttata Scudder, 1 876b, Proc. Boston 
Soc. Nat. Hist. 18: 265. 

Labia burgessi Scudder, 1876b, Proc. Bos- 
ton Soc. Nat. Hist. 18: 266. 

Labia melancholica Scudder, 1876b, Proc. 
Boston Soc. Nat. Hist. 18: 267. 

Prolabia pulchella (Audinet-Serville) He- 
bard, 1917, Entomol. News 28: 319. 

Laprobia pulchella (Audinet-Serville) 
Hincks, 1960, Proc. R. Entomol. Soc. 
Lond. (B)29: 156. 

Marava pulchella (Audinet-Serville) Brin- 
dle, 1971a, J. Nat. HisL 5: 557. 

Marava pulchella is the only earwig in 
South Carolina that exhibits both fully 
winged and brachypterous adults. When the 
tegminae and hindwings are fully devel- 
oped, the hindwings are visible, somewhat 
lighter in color basally, and the compound 
eyes are large. Brachypterous individuals 
have shorter tegminae, hindwings that are 
either absent or concealed, and smaller 
compound eyes. Factors governing the rel- 
ative proportions of these morphs in a pop- 
ulation are not known for M. pulchella, but 
temperature has been shown to be influ- 
ential for a different species (Knabke and 
Grigarick, 1971). Each branch of the male 
forceps generally has two teeth located api- 
cally and basally on the inner margin, al- 
though in some specimens the apical tooth 
may be missing. 

This earwig can be found under the bark 
of dead trees and in debris. Fully winged 
adults can be attracted to lights. Marava 
pulchella is native to southeastern United 
States, being found from North Carolina 
south to Florida and westward to Texas. It 
has also been recorded from Cuba (Alayo 
and Hernandez, 1980). 

South Carolina Records.— Marava pul- 
chella probably occurs statewide. Anderson 
Co.: Pendleton, 28 July 1972, in cotton seed, 
coll. R. P. Griffin, 1 <5. Rorence Co.: Flor- 
ence, 29 March 1930, coll. O. L. Cartwright, 
1 (5, 1 immature. Hampton Co.: Yemassee, 
5 January 1928, coll. J. A. Berly, 1 6; Ye- 

massee, 28 December 1941, coll. O. L. Cart- 
wright, 1 3. Lee Co.: Meredith, 19 April 
1928, coll. O. L. Cartwright, 1 $. Pickens 
Co.: Clemson, 7 March 1935, coll. J. G. 
Watts, 1 immature; Clemson, 20 October 
1972, hickory log, coll. R. P. Griffin, 1 <5; 
Isaqueena Forest, 2 August 1984, UV light 
trap, coll. K. M. Hoffman, 1 2; Clemson, 15 
August 1986, porchlight, coll. K. M. Hoff"- 
man, 1 $. Richland Co.: Columbia, 20 Feb- 
ruary 1926, coll. O. Cartwright, 1 imma- 

Vostox bmnneipennis 
(Audinet-Serville, 1839) 

Figs. 18, 30 

Psalidophora brunneipennis Audinet-Ser- 
ville, 1839, Hist. Nat. Ins. Orthop. p. 30. 

Vostox brunneipennis (Audinet-Serville) 
Burr, 1911, Genera Insect. 122: 51. 

This earwig is a reddish-brown to dark 
brown species with yellow hindwings that 
are bordered on their apical and inner mar- 
gins with brown. The male forceps usually 
have a single tooth located on the inner mar- 
gin near the midpoint, although some spec- 
imens possess a second tooth distally (Fig. 
1 8). This species is most commonly found 
under the bark of dead trees, but adults can 
also be attracted to lights. 

Vostox brunneipennis, a native species, is 
found from Panama northward to the 
southern United States. The holotype is from 
Pennsylvania, and in the United States this 
species is known from Illinois, Indiana, and 
Virginia south to Florida and westward to 
Texas and Oklahoma (Brindle, 1971b; Ar- 
nold and Drew, 1979). 

South Carolina Records. — Vostox brun- 
neipennis probably occurs statewide. An- 
derson Co.: Anderson, 1 1 March 1982, coll. 
G. Jones, 1 3. Greenwood Co.: Greenwood, 
5 February 1977, coll. P. Ellenburg, 1 9. Lee 
Co.: Meredith, 12 February 1927, coll. O. 
L. Cartwright, 1 3, 3 9. Pickens Co.: Clem- 
son, 9 January 1986, under bark of dead 
standing oak tree, coll. J. R. Brushwein, 1 



(5; Clemson, 1 August 1956, in trap light, 
coll. D. Dunavan, 1 9; Six Mile, 1 7 October 
1973, under log, coll. R. Peigler, 1 imma- 
ture. Saluda Co.: Ridge Spring, 25 July 1984, 
under bark of dead oak tree on ground, coll. 
D. E. Scotten, 2 immatures. 

Labidum riparia (Pallas, 1773) 

The Striped Earwig 

Fig. 24 

Forficula riparia Pallas, 1773, Reise Russ. 

Reiches 2: 727. 
Forficula bidens Olivier, 1791, Encycl. 

Method. 6: 466. 
Forficula erythrocephala Fabricius, 1793, 

Entomol. Syst. 2: 4. 
Labidura riparia (Pallas) Dohm, 1863, Stett. 

Entomol. Zeit. 24: 313. 
For a more complete synonymy, see Stein- 


The striped earwig is most easily recog- 
nized by both its large size and the two dark 
bands running the length of the pronotum. 
Also, there is a broad, dark, medio-dorsal 
band extending the length of the abdomen. 
This species is primarily predaceous, ac- 
tively seeking its prey at night (Schlinger et 
al., 1959). It can be collected on the ground 
at night or under debris during the day. 

Labidura riparia is an introduced species 
that occurs worldwide in both tropical and 
temperate zones. In the United States this 
earwig is established in the southern third 
of the country, from North Carolina south 
to Florida and westward to Texas, Arizona, 
and California (Langston and Powell, 1975). 

South Carolina Records.— The striped 
earwig is found throughout the state. Bam- 
well Co.: Blackville, 24 October 1975, Ed- 
isto Expt. Sta., pitfall traps from soybeans, 
coll. J. F. Price, 5 <5, 5 9. Beaufort Co.: Sea- 
brook, 18 October 1980, coll. M. K. Disney, 
1 immature. Charleston Co.: Charleston, 7 
February 1934, coll. J. A. Berly, 1 6. Edge- 
field Co.: Trenton, 19 November 1959, 
trunk of peach tree, coll. T. E. Skelton, 1 $. 
Greenville Co.: Greenville, 30 September 

1933, coll. H. T. Poe, Jr., 1 3. Horry Co.: 
Myrtle Beach, 27 December 1955, in dwell- 
ing of Joe C. Ivey, coll. D. Dunavan, 1 $, 1 
9. Pickens Co.: Clemson, 10 October 1961, 
under wood slat, coll. R. E. O'Brien, 1 im- 
mature; Clemson, 26 September 1984, on 
cement beneath lights at night, coll. K. M. 
Hoffman, 1 9. Sumter Co.: Sumter, Febru- 
ary 1955, coll. unknown, 1 immature. 

Dom aculeatum Scudder, 1876 

Figs. 5, 6, 10, 20,31 

Forficula aculeata (Scudder, 1876a, Proc. 
Boston Soc. Nat. Hist. 18: 262. 

Doru aculeatum (Scudder) Rehn and He- 
bard, 1914, J. N.Y. Entomol. Soc. 22: 93. 

Doru aculeatum is 12-18 mm long, and 
can be distinguished from the closely related 
species D. taeniatum by the lack of visible 
hindwings. This earwig is generally found 
on grasses and sedges growing near water 
(Hebard, 1934; Cantrell, 1968). Doru acu- 
leatum is native to eastern North America, 
ranging from southern Michigan and On- 
tario to Florida and westward to Illinois, 
Nebraska, and Louisiana. 

South Carolina Records.— Dorw aculea- 
tum has only been found in the extreme 
northwestern part of the state and at Myrtle 
Beach. Anderson Co.: Portman Shoals, 24 
November 1927, coll. F. Sherman, 2 3, 1 9. 
Horry Co.: Myrtle Beach, 1 March 1965, 
sedge, coll. V. M. Kirk, 2 <5. Oconee Co.: 
Seneca, 10 September 1969, on jewel weed, 
coll. J. W. Van Duyn, 1 5. Pickens Co.: 
Clemson, 1 4 October 1 96 1 , in com ear, coll. 
J. A. DuRant, 3 3, 1 9; Pickens, 28 March 
1978, coll. J. Keith, 1 3; Rocky Bottom, 22 
May 1934, coll. O. L. Cartwright, 1 3; Six 
Mile, 26 October 1963, under dead willow 
bark, coll. D. G. Bottrell, 1 3, 1 9. 

Dom taeniatum (Dohrn, 1862) 

Fig. 19 

Forficula taeniata Dohm, 1862, Stett. Ento- 
mol. Zeit. 23: 230. 



Forficula californica Dohm, 1865, Stett. 
Entomol. Zeit. 25: 85. 

Forficula exilis Scudder, 1876a, Proc. Bos- 
ton Soc. Nat. Hist. 18: 262. 

Doru exile (Scudder) Burr, 1911, Genera In- 
sect. 122: 79. 

Doru lineare (Eschscholtz) Burr, 1911, Gen- 
era Insect. 122: 79 (partim). 

Doru taeniatum (Dohm) Brindle, 1971c, 
Papeis Avulsos Zool. 23: 191. 

Doru taeniatum is distinguished from the 
closely related D. aculeatutn by the presence 
of visible hindwings. The species Doru lin- 
eare (Eschschlotz, 1822) is listed by Blatch- 
ley (1920), Gumey (1950), and Langston 
and Powell (1975) as occurring in the United 
States. However, a revision of the genus 
Doru has shown the range of this species to 
be limited to Brazil, Argentina, and Para- 
guay (Brindle, 1971c). Records of this species 
in North America are now referrable instead 
to D. taeniatum (Gumey, 1972). 

Doru taeniatum ranges from Bolivia and 
Colombia northward to the southeastem 
United States (Brindle, 1971c). It has been 
found as far north as Maryland and as far 
west as Texas, Arizona, and Califomia. It 
is unclear whether it is native to North 
America or was introduced from Central 
and South America (Gumey, 1972). 

South Carolina Records.— Dorw taenia- 
tum probably occurs locally statewide, but 
seems to be more prevalent in the southern 
and eastem counties. Bamwell Co.: Black- 
ville, 18 October 1978, on soybeans, coll. 
G. Sanders, 1 9. Berkeley Co.: St. Stephen, 
17 September 1983, coll. T. Johnson, 1 5. 
Charleston Co.: Charleston, 17 March 1980, 
coll. M. Wallace, 1 $. Dorchester Co.: Four 
Holes Swamp, 13 August 1976, Mellards 
Lake, coll. J. Morse and J. Chapin, 1 9. 
Hampton Co.: 1 October 1983, coll. D. Ed- 
wards, 1 $. Horry Co.: Conway, 20 March 
1976, woods, coll. T. Thompson, 1 9. Pick- 
ens Co.: Clemson, 16 March 1976, on 
ground, coll. C. Wilson, 1 3; Clemson, 10 
October 1976, coll. T. Currin, 1 aduh; 

Clemson, 18 September 1978, coll. C. Whit- 
mire, 1 9. 

Forficula auricularia L., 1758 

The European Earwig 
Figs. 9, 25 

Forficula auricularia Linnaeus, 1758, Syst. 
Nat. (10) 1:423, no. 1. 

Sakai (1973) gives a more complete syn- 

The European earwig is 15-20 mm in 
length and can be recognized by the basally 
broadened forceps of the male (Fig. 25). It 
is the most economically destmctive ear- 
wig, occasionally causing substantial dam- 
age to cereals, fmits, seedling vegetables, and 
flowers. Unlike most earwigs, the European 
earwig does not survive well in warm cli- 
mates, preferring instead cool, moist re- 
gions (Crumb et al., 1941). The European 
earwig occurs in temperate regions around 
the world and has become widely estab- 
lished in North America. It has been re- 
corded locally across southern Canada 
southward to North Carolina and westward 
to Arizona and Califomia (Langston and 
Powell, 1975; Scudder and Vickery, 1985). 

South Carolina Records. — The only re- 
cord of this species in South Carolina is of 
a female taken in Charleston. Due to this 
species' preference for cool climates and the 
fact that Charleston is a major seaport, it is 
possible that this specimen is an adventive 
and not a member of an established popu- 
lation. Further collecting is necessary to de- 
termine whether the European earwig is es- 
tablished in South Carolina. Charleston Co.: 
Charleston, 30 October 1983, coll. B. F. Ce- 
lek, 1 9. 


I am grateful to the following for allowing 
me to examine material: Lewis Deitz and 
Carol Parron (North Carolina State Uni- 
versity), Cecil Smith (University of Geor- 
gia), Lionel Stange (Florida State Collection 
of Arthropods) and David Nickle (Agricul- 



tural Research Service, USDA, National 
Museum of Natural History). Gratitude is 
also extended to Peter H. Adler and John 
C. Morse (Clemson University) for review- 
ing the manuscript. This is Technical Con- 
tribution No. 2593 of the South Carolina 
Agricultural Experiment Station, Clemson 

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89(1), 1987, pp. 15-23 




Michael D. Schwartz and Gary M. Stonedahl 

Department of Entomology, American Museum of Natural History, Central Park West 
at 79th Street, New York, New York 10024. 

Abstract.— The new genus Oaxacacoris is diagnosed and described, including three new 
species: cygnus, pueblensis, schaffnerL All species occur in southcentral Mexico. The male 
genitalia of all species are illustrated, and a dorsal habitus provided for the male oi cygnus. 
Scanning electron micrographs of the scale-like setae and pretarsus of schajfneri also are 

The idea for this paper originated during 
our revision of the genus Pseudopsallus 
(Stonedahl and Schwartz, 1 986). Among the 
material on loan to us from Joseph C. 
Schaffner, Texas A&M University, College 
Station were specimens with a resemblance 
to Pseudopsallus, but which we now place 
in the new genus Oaxacacoris. The rela- 
tionship of Oaxacacoris to other orthotyline 
genera is not well understood. This is due 
to the limited knowledge of Mexican Miri- 
dae in general, as well as the poorly defined 
limits of many New World genera of Or- 

In our revision of the western North 
American genus Pseudopsallus (Stonedahl 
and Schwartz, 1986), we selected Oaxaca- 
coris as the sister group of the former genus 
because these two genera shared derived 
characters of the male genitalia that have 
yet to be discovered in other genera of the 
tribe. Further, we suggested that Oaxaca- 
coris and Pseudopsallus belonged to a com- 
plex of genera united by similarities in type 
and fine structure of setae and by the general 
form of the male genitalia. Oaxacacoris can 
be distinguished from other genera of this 
group by the characters given in the generic 

diagnosis. The only known host association 
for the genus is Mimosa rhododactyla B. L. 
Robinson (Fabaceae). 

All holotypes and some paratypes are de- 
posited in the collection of the American 
Museum of Natural History, New York 
(AMNH). The remaining paratypes and ad- 
ditional specimens are deposited in the Tex- 
as A&M University collection (TAM). All 
measurements are in milHmeters. 

Oaxacacoris Schwartz and Stonedahl, 
New Genus 

Diagnosis. — Recognized by the green or 
yellow green, slightly shining general ap- 
pearance; dorsal vestiture with silvery and 
golden brown to black, narrow, scale-like 
setae with converging ridges (Fig. 2a); and 
male genitalia with the following character- 
istics: anterodorsal margin of aperture of 
genital capsule with three tergal processes, 
one right lateral and two left lateral of mid- 
line (Figs. 3, 10, 17); lateralmost pair of 
processes very large, slightly flattened, ex- 
tending well beyond margins of aperture; 
left medial tergal process small, simple, ori- 
ented ventrad into aperture; dorsal lobe of 
left paramere greatly expanded distally, with 



4-9 Strong spines (Figs. 6, 13, 19); right par- 
amere subquadrate in lateral view, with 
prominent, sometimes bifurcate, basal lobe 
(Figs. 8, 15, 22); and right spicula of vesica 
bisected to base, dorsal portion much small- 
er than ventral portion (Figs. 9, 16, 23). 

Description.— Male. Macropterous. Small 
to moderate subovate body form; length 
3.50-4.90 (apex of tylus to apices of hemel- 
ytra); width (across humeral angles of 
pronotum) 1.15-1.40; surface texture 
smooth; coloration dark to light green, vari- 
ably suffused or mottled with yellow, with 
frons, anterior portion of pronotum, em- 
bolium, scutellum, and cuneus sometimes 
extensively yellow; dorsal vestiture with re- 
cumbent, narrow, golden scale-like setae and 
long, suberect, golden simple setae, some- 
times also with recumbent, narrow, black 
scale-like setae and short to long, suberect, 
black simple setae. Head: Subtriangular; 
length of portion of head anterior of eye 
subequal to length of head posterior of an- 
terior margin of eye in dorsal or lateral view; 
posterior margin without carina; frons and 
vertex weakly convex, gently sloping anteri- 
ad, meeting tylus with weak suture; tem- 
poral area obsolete; eyes large, subovate, 
projecting slightly beyond anterolateral an- 
gles of pronotum in dorsal view, occupying 
six-sevenths of head height in lateral view, 
posterior margin sublinear, anterior margin 
slightly emarginate; antennal fossae small, 
contiguous with anterior margin of eye, ven- 
tral margin of fossae slightly to obviously 
dorsad of ventral margin of eye in lateral 
view; tylus smoothly curving to apex; juga 
triangular, flattened to slightly convex; lora 
rectangular, slightly swollen; gena present, 
width equal to diameter of antennal seg- 
ment I; gula small; length of buccular flange 
and cavity equal, not reaching posterior 
margin of head; labium reaching apices of 
metacoxae or sometimes to sixth stemite; 
antennal segment I with length equal to or 
slightly greater than width of vertex, linear, 
tapered basally, with a few erect bristle-like 
setae on distal and interior surface; seg- 

ments II-IV cylindrical, linear, diameter 
smaller than segment one, with densely dis- 
tributed, reclining, short, black simple setae. 
Pronotum: Trapezoidal, broader than long, 
slightly sloping transversely and longitudi- 
nally; lateral margins straight, smoothly 
curved at junction with propleura; posterior 
margin straight; anterior and posterior an- 
gles broadly rounded; anterior margin 
slightly arcuate medially; calli weakly con- 
vex, reaching anterior angles and lateral 
margin of pronotum, confluent anterome- 
dially, bordered posteriorly by faint to shal- 
low depression; mesoscutum narrowly ex- 
posed; scutellum weakly convex or flattened. 
Hemelytra: Subparallel-sided, widest me- 
dially; claval vein weakly elevated; radial 
vein elevated basally, obsolete distally; cu- 
neal incisure small; cuneal fracture angled 
slightly anteriad; cuneus longer than broad; 
membrane lightly to strongly suffused with 
uniform fuscous, light species with short spot 
or band of dark fuscous or black near in- 
tervein of inner cell; inner cell longer than 
cuneus, slightly narrowed distally; outer cell 
triangular. Legs: Coloration uniformly yel- 
low, or mixed yellow and yellow green with 
variable number of green spots; femora 
slightly flattened, tapered distally and ba- 
sally, with short, reclining, black or fuscous 
simple setae, sometimes with setae restrict- 
ed to anterior portion; meso- and meta- 
tibiae with several rows of minute, dark 
spinulae; tibiae with suberect, light, simple 
setae; tibial spines dark or light fuscous; api- 
ces of tibiae dark fuscous; apices of third 
tarsal segment black; tarsal segment one half 
as long as segment two and three; claws 
strongly curved, pulvilli connate to interior 
surface (at angle) of claw (Fig. 2b). Genitalia: 
Genital capsule: Large, subrectangular in 
ventral view, width greater than length, 
anterodorsal margin with three large var- 
iously shaped, posteriorly directed sclero- 
tized tergal processes; processes lateral of 
midline of capsule, and not supporting rec- 
tal opening; left processes consisting of sin- 
gle large, basally spinose process, originat- 



Fig. 1 . Oaxacacoris cygnus. dorsal habitus of male. 

ing on posterior edge of anterodorsal margin 
(left tergal process, LP) and single small pro- 
cess originating on ventral surface of antero- 
dorsal margin, oriented ventrad into aper- 
ture (left ventral tergal process, LVP); right 

process solitary, large, usually orientated to 
left side of aperture (right tergal process, 
RP); postero ventral margin of capsule with 
deep paramere and phallotheca sockets, and 
without posteriorly directed processes; ap- 



Fig. 2. Oaxacacoris schaffneri. 2a, Scale-like setae on dorsum. 2b, Pretarsal structures. 

erture large, subcircular. Left paramere: 
Dorsal lobe (DLL) very large, flattened, 
strongly spinose, and usually expanded dis- 
tally; lateral lobe (LAL) narrow, rounded, 
and curved or sometimes notched distally. 
Right paramere: Subquadrate in lateral view; 
interior surface with prominent medial lobe 
(MIL), sometimes obsolete; dorsal lobe 
(DLR) usually spinose marginally and pro- 
jecting dorsad of medial lobe; basal lobe 
(BLR) solitary or strongly bifurcate, strong- 
ly spinose. Phallotheca: laterally com- 
pressed, open on dorsal and distal surfaces, 
opening narrow and linear dorsally, merg- 
ing with larger irregularly shaped distal por- 
tion; without secondary ventrobasal open- 
ing. Vesica: Ductus seminis simple, 
cylindrical, flexible with ribs; secondary 
gonopore horseshoe-shaped, base short and 
weakly sclerotized; two long posteriorly di- 
rected vesical spiculae narrowly attached 
right and left of midline of ductus basad of 
secondary gonopore, and practically sur- 
rounding ductus; right spicula bifurcate; 
dorsal portion (DS) short, simple or weakly 
serrate; ventral portion (RVS) with thick 
basal half, tapering to long, flattened or 
rounded, strongly and marginally serrate 
distal half; left spicula (LVS) with long, 
sometimes sinuate, rounded shaft, distal 
third recurved, flattened and strongly ser- 
rate marginally, base of recurved portion 
with fine serrations. 

Female. Macropterous. Similar to male 
in color, vestiture, and structure, except 
usually somewhat larger with slightly 
broader head and vertex. Genitalia: Follow- 
ing the terminology of Slater (1950). Scler- 
otized rings: Large, elongate, widely sepa- 
rated; lateral margin of ring moderately 
folded dorsally, not extending mesad. Pos- 
terior wall: K structures of moderate size, 
heart-shaped with bifid apex, well separated 
medially; L structure and portion of J struc- 
ture visible in anterior view. 

Etymology.— Named for the state of Oa- 
xaca in Mexico, the type locality of the type 
species; masculine. 

Type spQcies. — Oaxacacoris cygnus, new 

Distribution. — Guerrero, Michoacan, 
Oaxaca, and Puebla, Mexico (Fig. 24). 

Discussion. — Identification of the species 
of Oaxacacoris is most easily accomplished 
by careful examination of the male genitalic 
structures. To faciliate identification we 
provide illustrations and descriptions of all 
the structures for each species. 

Oaxacacoris cygnus, New Species 

Figs. 1, 3-9, 24 

Diagnosis. — Recognized by the large size; 
labium reaching apices of metacoxae or 
slightly beyond; dorsal vestiture always with 
densely distributed, dark brown or black 
scale-like setae; and by the structure of the 



male genitalia, especially the shape of the 
tergal processes (Figs. 3, 4), the swan-shaped 
left paramere with 6-9 strong spines on dis- 
tal portion of dorsal lobe (Fig. 6), and basal 
lobe of right paramere bifurcate, with very 
large, laterally directed basalmost spine 
(Figs. 7, 8). 

Description. — Male (Fig. 1). Length 4.15- 
5.20; mottled dark green general coloration; 
dorsal vestiture with recumbent, narrow, 
black scale-like setae and narrow, golden 
simple setae. Head: Width across eyes 0.85- 
0.86, width of vertex 0.37-0.39; dark green, 
tinged with extensive yellow green or yellow 
areas overall; antennae yellow or yellow 
brown, segment IV and distal end of seg- 
ment III fuscous; length of segment I 0.35- 
0.45, segment II 1 .5 1-1.65; labium reaching 
apices of metacoxae or slightly beyond. 
Pronotum: Posterior width 1 .40-1 .59; whit- 
ish green anterior of calli, calli and disk dark 
green, disk narrowly adjoining posterior 
margin of calli yellow, posterior angles faint 
green; mesoscutum yellow laterally and me- 
dially; scutellum with yellow apex. Hemel- 
ytra: Dark mottled green; cuneus, emboli- 
um, clavus and corium bordering embolium, 
and apex of clavus sometimes pale yellow 
or yellow green; membrane lightly suffused 
with fuscous and with band of dark fuscous 
bordering apex of inner cell; veins pale. Legs: 
Yellow, metafemora and sub-basal portion 
of metatibiae tinged with small green spots; 
femora with short, reclining black simple 
setae. Genitalia: RP very long, flattened, 
parallel-sided, distal fourth tapered and with 
several strong spines, or with only single 
large secondary serrate spine, apex pointed 
and directed to left; LP long, with bifurcate 
basal spine, or with simple basal spine, and 
recurved pointed apex; LVP cylindrical, 
sinuate with bifurcate pointed apex. Left 
paramere: DLL palmate with six to nine 
spines; LAL flattened, sinuate with pointed 
apex. Right paramere: MIL simple; DLR 
somewhat pointed distally with several small 
spines; BLR bifurcate, with long simple 
slightly recurved basalmost spine, distal- 

most spine solitary, or weakly to deeply bi- 
furcate. Phallotheca: dorsodistal edge re- 
curved. Vesica: DS faintly serrate, shorter 
than RVS; RVS flattened and tapered, with 
strongly serrate margin, apex pointed; LVS 
with long, tapering recurved distal third, 
apex pointed. 

Female. Length 4.35-5.05; width across 
eyes 0.78-0.85, width of vertex 0.40-0.43; 
length of antennal segment I 0.43-0.44, seg- 
ment II 1.50-1.64; posterior width of 
pronotum 1.50-1.64. 

Etymology. — From the latin cygnus 
(swan), referring to the swan-like shape of 
the left paramere in lateral view. 

Distribution. — Figure 24. 

Holotype $.- MEXICO: Oaxaca. 4 mi 
NE of Miltepec, July 21 1984, Carroll, 
Schaflher, Friedlander. 

Paratypes.— M£'A'/CO.- Oaxaca. 6 3, 5 9 
same data as holotype. 

Additional specimens. — M£'A'/C6>.- 
Guerrero. 13 3, 1 5 2 6 mi E of Xochipala, 
July 13, 1985, Jones, Schaff'ner. Puebla: $, 
9 4 mi SW of Acatepec, July 21, 1984, Car- 
roll, Schaflher, Friedlander; 3, 3 9 4.4 mi 
SW of Acatepec, July 9, 1977, Schaflher. 

Oaxacacoris schaffneri, New Species 
Figs. 2a, 2b, 10-16, 24 

Diagnosis. — Similar to cygnus, but dif- 
ferentiated by its smaller size; labium reach- 
ing well beyond apices of metacoxae, some- 
times to sixth abdominal sternite; dorsal 
vestiture without, or with only a few dark 
brown, scale-like setae; and by the structure 
of the male genitalia, especially the shape 
of the tergal processes (Figs. 10, 11) and 
dorsal lobe of left paramere with only four 
strong spines distally (Fig. 1 3). 

Description. — Male. Length 3.58-4.05; 
mottled green general coloration; dorsal 
vestiture with recumbent, narrow golden or 
brown, scale-like setae and narrow, golden 
simple setae. Head: Width across eyes 0.72- 
0.78, width of vertex 0.33-0.34; yellow or 
yellow green; antennae yellow or yellow 
brown, segments III and IV slightly fuscous; 



Figs. 3-16. Male genitalic structures. 3-9, Oaxacacoris cygnus. 3, 4, Tergal processes of genital capsule. 3, 
Dorsal view (LP, left process; LVP, left ventral process; RP, right process). 4, Posterior view. 5, 6, Left paramere. 



length of segment I 0.34-0.35, segment II 
1.19-1.26; labium surpassing apices of 
metacoxae, sometimes reaching sixth ster- 
nite. Pronotum: Posterior width 1.16-1.20; 
yellow or yellow green anteriad of posterior 
depression of calli, disk mottled green and 
yellow or sometimes entirely yellow green 
with margin of disk faint green; mesoscu- 
tum orange yellow; scutellum yellow with 
faint orange cast; area adjoining mesoscu- 
tum mostly orange. Hemelytra: Mottled 
green and yellow, or sometimes mostly 
mottled yellow; cuneus and embolium yel- 
low; membrane very lightly suffused with 
fuscous, more so peripherally, sometimes 
with small testaceous band near apex of in- 
ner cell; veins testaceous. Legs: Yellow; tarsi 
and apices of tibiae brownish yellow; fem- 
ora with short, reclining, yellow simple se- 
tae. Genitalia: RP long, flattened, parallel- 
sided, distal portion tapered and weakly 
serrate marginally, with apex pointed and 
directed to left; LP long, thick, with several 
large serrations basally, tapered and narrow 
distally, apex pointed; LVP cylindrical and 
simple. Left paramere: DLL palmate with 
four spines; LAL somewhat flattened, sin- 
uate with pointed apex. Right paramere: 
MIL simple; DLR with several small spines 
distally; BLR deeply bifurcate, with long 
simple basalmost spine, distalmost spine 
short with bifid apex. Vesica: DS serrate, 
shorter than RVS; RVS wide and flattened, 
with strongly serrate margin, apex truncate 
and serrate; LVS long, broad recurved distal 
third, medially tapering to narrow apex. 

Female. Length 3.81-4.25; width across 
eyes 0.76-0.80, width of vertex 0.38-0.40; 
length of antennal segment I 0.35-0.40, seg- 

ment II 1.37-1.55; posterior width of 
pronotum 1.27-1.31. 

Etymology. — Named for Joseph C. 
Schaffher, who graciously provided all spec- 
imens for this study. 

Distribution. — Figure 24. 

Holotype S.- MEXICO: Michoacan. 30 
mi S of Nueva Italia, August 8, 1978, Plitt, 

Paratypes.— M£'X/CO.- Michoacan. 5 $, 
1 2 9 same data as holotype. 

Additional specimens. — MEXICO. Mi- 
choacan. 20 (5, 30 9 28.5 mi S of Nueva 
Italia, July 9, 1 985, Jones, Schaffher, ex Mi- 
mosa rhododactyla B. L. Robinson (Faba- 

Oaxacacoris pueblensis, New Species 
Figs. 17-24 

Diagnosis. — Readily distinguished from 
cygniis and schaffneri by the following char- 
acteristics of the male genitalia: left tergal 
process bulbous with numerous, tiny ser- 
rations (Figs. 17,1 8); lateral lobe of left par- 
amere notched apically (Fig. 20); basal lobe 
of right paramere solitary, with large sec- 
ondary spines (Figs. 2 1 , 22); and dorsal spic- 
ula of vesica small and nonserrate (Fig. 23). 
The scale-like setae of the dorsal vestiture 
are silvery to golden brown, never dark 
brown or black as in cygnus and sometimes 
schaffneri. Further distinguished from 
schaffneri by the shorter labium. 

Description. — Male. Length 3.43; mot- 
tled green and yellow general coloration; 
dorsal vestiture with recumbent, long, nar- 
row, golden scale-like setae and long, sub- 
erect, golden to nearly black simple setae; 
metacuneus, clavus and corium bordering 

5, Posterior view (DLL, dorsal lobe; LAL, lateral lobe). 6, Lateral view (DLL, LAL; as in Fig. 5). 7, 8, Right 
paramere. 7, Dorsal view (BLR, basal lobe; DLR, dorsal lobe; MIL, medial interior lobe). 8, Inside lateral view 
(BLR, DLR, MIL; as in Fig. 7). 9, Spiculae of vesica (DS, dorsal; LVS, left ventral; RVS, right ventral). 10-16, 
Oaxacacoris schaffneri. 10, 11, Tergal processes of genital capsule. 10, Dorsal view. 1 1, Posterior view. 12, 13, 
Left paramere. 12, Posterior view. 13, Lateral view. 14, 15, Right paramere. 14, Dorsal view. 15, Inside lateral 
view. 16, Spiculae of vesica. 



Figs. 17-23. Male genitalic structures of Oa.vacacom pw?W^«5/5. 17, 18, Tergal processes of genital capsule. 
17, Dorsal view. 18, Posterior view. 19, 20, Left paramere. 19, Lateral view. 20, Posterior view. 21, 22, Right 
paramere. 21, Dorsal view. 22, Inside lateral view. 23, Spiculae of vesica. 

claval suture sometimes with darker brown 
scale-like setae. Head: Width across eyes 
0.73, width of vertex 0.34; yellow; antennae 
testaceous, segment IV and apex of segment 
III fuscous; length of segment I 0.38, seg- 
ment II 1.13; labium reaching apices of 
metacoxae. Pronotum: Posterior width 1.16; 
yellow anteriad of posterior depression of 
calli, calli somewhat darker, disk mottled 
green and yellow, peripheral border faint 
green; mesoscutum and scutellum orange 
yellow. Hemelytra: Mottled green and yel- 
low, interior of cuneus yellow; membrane 
darkly suffused with fuscous; veins fuscous, 
rufous adjacent to cuneus. Legs: Yellow; 
tarsi and apices of tibiae lightly fuscous, with 
long, suberect, yellow simple setae. Geni- 
talia: RP long, thick basally, narrowed me- 

dially, apex truncate with single large spine 
directed to left; LP bulbous, surface densely 
serrate, with single apical and basal spines; 
LVP cylindrical, short with pointed apex. 
Left paramere: DLL long, broad basally, ta- 
pering to pointed apex, proximal surface 
with five spines; LAL fairly broad, with 
notched apex. Right paramere: MIL obso- 
lete; DLR small, with several small spines 
on interiobasal surface; BLR large and sol- 
itary, with large spines, apex extending dis- 
tad of distal end of DLR, and with large 
basal spines. Vesica: DS very small, simple; 
RVS narrow with strong marginal serra- 
tions, much smaller than LVS; LVS with 
curved portion strongly hooked and mar- 
ginally serrate. 

Female. Length 4.01-4.32; width across 



Fig. 24. Distribution of Oaxacacoris cygnus •, 
Oaxacacoris schaffneri A, Oaxacacoris pueblensis ■. 

eyes 0.76-0.77, width of vertex 0.39-0.40; 
length of antennal segment I 0.44-0.46, seg- 
ment II 1.40-1.65; posterior width of 
pronotum 1.31-1.38. 

Etymology. — Named for its occurrence in 
the state of Puebla, Mexico. 

Distribution.— Figure 24. 

Holotype $.- MEXICO: Puebla. 16 mi 
NW of Acatlan, July 14, 1974, Clark, Mur- 
ray, Ashe, Schaffner. 

Paratypes.-M£'A'/CO.- Puebla. 3, 15 $ 
same data as holotype. 

Additional specimens.— M£'X/C(9.' Pu- 
ebla: 4 9 5 mi SE of Izucar de Matamoros, 
July 20, 1974, Carroll, Schaffner, Friedlan- 
der; 9 7.3 mi SW Izucar de Matamoros, July 

22, 1981, Bogar, Schaffner, Friedlander. In- 
dividuals of these populations are slightly 
greater in length (4. 1 4-4.68) and darker than 
the holotype and paratypes but are indis- 
tinguishable from the later specimens with 
regard to external morphology and vesti- 
ture. We are tentatively recognizing them 
as conspecific with the type, pending the 
acquisition of male specimens from other 
areas for detailed comparison. All three lo- 
calities for this species are within a 35 mile 


We thank Kathleen Schmidt, Hillsdale, 
New York for the fine illustration of the 
adult male of Oaxacacoris cygnus, Lauren 
Duffy, Interdepartmental Laboratory, 
American Museum of Natural History, who 
assisted with the preparation of the scanning 
electron micrographs, and an anonymous 
reviewer for improving the manuscript. 

The authors contributed equally to all fac- 
ets of this paper. 

Literature Cited 

Slater, J. A. 1950. An investigation of the female 
genitalia as taxonomic characters in the Miridae. 
Iowa St. J. Sci. 25: 1-81. 

Stonedahl, G. M. and M. D. Schwartz. 1986. Revi- 
sion of the plant bug genus Pseudopsallus Van Du- 
zee (Heteroptera: Miridae). Am. Mus. Novit. No. 
2842, 58 pp. 


89(1), 1987, pp. 24-30 




Boris C. Kondratieff and Ralph F. Kirchner' 

(BCK) Colorado State University, Department of Entomology, Fort Collins, Colorado 
80523; (RFK) U.S. Army Corps of Engineers, Huntington District, Water Quality Section, 
502 8th Street, Huntington, West Virginia 25701. 

Abstract.— One hundred and forty-nine species of stoneflies are recorded from Virginia. 
Of these, 80 species have distributions associated with the Appalachian Mountains, 57 
are widely distributed east of the Rocky Mountains, 6 are associated with the Coastal 
Plain Physiographic Province, and 6 have ranges which include both the Appalachians 
and the Coastal Plain. There are apparently 7 regional endemic species. Twenty-one species 
are recorded from Virginia for the first time. 

Kondratieff and Voshell (1979) reported 
1 1 6 species of Plecoptera known from Vir- 
ginia in their checklist of the stoneflies of 
Virginia. Since 1979, 21 state records have 
been added and seven new species described 
from Virginia. There also have been several 
recent important taxonomic changes: Pel- 
toperlidae (Stark and Stewart, 1981), Ptero- 
narcyidae (Stark and Szczytko, 1982), Per- 
lodidae (Stewart and Stark, 1984; Szczytko 
and Stewart, 1981), Chloroperiidae (Sur- 
dick, 1985), and Perlidae (Zwick, 1984). 
Three genera, Isoperla by Szczytko, Sweltsa 
by Surdick, and Perlesta by Stark, are being 
revised and will add additional taxa to Vir- 

This species list is presented to provide a 
framework for future studies of the Plecop- 
tera of Virginia, including an identification 
manual. The 149 species reported herein 
from Virginia are the most recorded from 
any state or province from the Nearctic re- 

' The views of the Author do not purport to reflect 
the position of the Department of the Army or the 
Department of Defense. 

gion. The following numbers of species have 
been reported from surrounding states: 
Kentucky- 77 (Tarter et al., 1984), North 
Carolina— about 130 (Unzicker and Mc- 
Caskill, 1982), and West Virginia- 106 
(Tarter and Kirchner, 1980). 

The richness of the stonefly fauna from 
Virginia is indicative of the range of topog- 
raphy which allows for heterogeneously di- 
verse habitats. Five major physiographic 
provinces of eastern North America (Coast- 
al Plain, Piedmont Plateau, Blue Ridge, 
Ridge and Valley, and the Appalachian Pla- 
teau) are represented in Virginia (Hoffman, 
1969). Fig. 1 delimits the physiographic 
provinces of Virginia and the major river 
systems. Virginia has eight major river ba- 
sins, several of old geographical systems such 
as the New River (Hoffman, 1969). The to- 
pographic relief of Virginia ranges from sea 
level to 1743 m (Mt. Rogers, 5720 ft.). 

The distribution of Plecoptera of Virginia 
has three basic patterns. Eighty species are 
closely linked with the central Appalachians 
and foothills of the upper Piedmont Plateau 
region. This area has one of the most diverse 



\x_'r 1 '1 


y^ New ^ V/ y^ ^N| ^"^ -^ 5^r^ 

^,,..c^vX'^''^^'^nif ^ "T\ 

Fig. 1 . Physiographic provinces of Virginia (according to Hoffman, 1 969). CP = Coastal Plain; PP = Piedmont 
Plateau; BR = Blue Ridge; RV = Ridge and Valley; AP = Appalachian Plateau. The symbol * identifies Mt. 

fauna and flora assemblages of the North 
Temperate Zone. Immatures of these species 
usually occur in cooler streams or springs 
of the Appalachian Plateau, Ridge and Val- 
ley, Blue Ridge, and Piedmont Plateau 
provinces. Some of these species are pre- 
dominantly southeastern Appalachian, with 
southern Virginia being their northern limit 
of distribution. Included in this group are 
Allocapniafumosa Ross, A. stannardi Ross, 
Alloperla nanina Banks, Megaleuctra wil- 
liamsae Hanson, Strophopteryx limata (Pri- 
son), Sweltsa mediana (Banks), S. urticae 
(Ricker), and Yugus arinus (Prison). The 
northern border of the southern Appala- 
chians is generally considered to be where 
the Roanoke River slices through the Blue 
Ridge Mountains (Fig. 1). Most the the 
species listed above may be collected from 
streams draining Virginia's highest peaks, 
Mt. Rogers-White Top area in Grayson, 
Smyth, and Washington counties (Fig. 1). 

Several Appalachian species have strong 
northeastern affinities, and Virginia in- 
cludes their southern distribution limits. 
These species may have survived the ice 
ages in the extensive unglaciated areas of 

Canada. These likely northern emigrants in- 
clude: Allocapnia maria Hanson (most 
southern record, Smyth County), Alloperla 
banksi Prison, all four of the eastern species 
of Ostwcerca, and Sweltsa naica (Pro- 

Many species, however, are widespread, 
occurring all along the Appalachian Moun- 
tains from Canada to Georgia: Acroneuria 
cawlinensis (Banks), Bolotoperla rossi (Pri- 
son), Hansonoperla applachia Nelson, Ma- 
lirekus hastatus (Banks), and Sweltsa onkos 

About 57 species may be considered to 
have widespread boreal distributions. These 
generally occur east of the Rocky Mountain 
region. In Virginia, these species are found 
in all five physiographic provinces. Good 
examples are Acroneuria abnormis (New- 
man), Allocapnia granulata (Claassen), A. 
recta Claassen, A. rickeri Prison, Alloperla 
caudata Prison, Amphinemura nigritta 
(Provancher), Attaneuria ruralis (Hagen), 
Clioperla clio (Newman), Haploperla brevis 
(Banks), Isoperla bilineata (Say), Leuctra 
ferruginea (Walker), Perlinella drymo 
(Newman), P. ephyre (Newman), Prostoia 



completa (Walker), P. similis (Hagen), Neo- 
perla clymene (Newman), and Strophopte- 
ryxfasciata (Burmeister). 

Six species are generally restricted to the 
Coastal Plain physiographic province, and 
as Stark (1979) stated, occupy "a boomer- 
ang shaped range that extends northward 
along the Atlantic Coast and to the west 
along the Gulf Coast." Acroneuria arenosa 
(Banks), Neoperla carlsoni Stark and Bau- 
mann, and Taeniopteryx lonicera Ricker and 
Ross are examples of this group. Nymphs 
of these species are most commonly col- 
lected on submerged woody substrate and 
among accumulated organic debris in the 
many shifting sand streams of the area. 
Adults of the perlids are often only collected 
in numbers using light traps. Several species, 
including Allocapnia virginiana Prison, A. 
wrayi Ross, Eccoptura xanthenes (New- 
man), Helopicus subvarians (Banks), and 
Paragnetinafumosa (Banks) range through- 
out the Coastal Plain but also occur in the 

There are apparently several regional en- 
demic species whose affinities are clearly 
with the Appalachian fauna. These include 
Diploperla morgani Kondratieff and Vosh- 
ell, D. kanawholensis Kirchner and Kon- 
dratieff, Alloperla biserrata Nelson and 
Kondratieff, Taeniopteryx nelsoni Kondra- 
tieff and Kirchner, Allocapnia harperi 
Kirchner, and Tallaperla lobata Stark. 

Significant range extensions are noted for 
several species. Isogenoides varians (Walsh) 
was collected from Bedford County (Big Ot- 
ter River). This species is known from II- 
hnois, Indiana, Michigan, Mississippi, South 
Carolina, and Tennessee (Stewart and Stark, 
1984). The Big Otter River has a diverse 
stonefly fauna. Diploperla kanawholensis 
was also collected from this river and rep- 
resents a new state record (Kirchner and 
Kondratieff, 1984). The eastern species of 
Ostrocerca have been considered rare. How- 
ever, the four eastern species often occur 
abundantly but locally in the Appalachian 
region of Virginia. 

In the list that follows, the 21 new state 
records are indicated by # and the seven 
species described since 1979 by +. In ad- 
dition, 2 1 other species that probably occur 
in Virginia are indicated by *, and the ad- 
jacent states where they have been reported 
are listed in parentheses. Each species in the 
following list is identified by its continental 
distribution: widespread boreal species 
(WB), Appalachian (AP), and Coastal Plain 
(CP), and Appalachian-Coastal Plain dis- 
tribution (AP-CP). We are following the 
classification of Zwick (1973). The follow- 
ing papers are necessary to supplement those 
Usted by Kondratieff and Voshell (1979) to 
identify the stoneflies of Virginia and sur- 
rounding states: Harper and Stewart ( 1984), 
Kirchner (1980, 1982), Kirchner and Har- 
per (1983), Kirchner and Kondratieff ( 1984, 
1985), Kondratieff and Kirchner (1982a, 
1982b), Kondratieff and Voshell (1981, 
1982), Kondratieff et al. (1981), Nelson 
(1979, 1982), Nelson and Kondratieff 
(1983), Stark (1983, 1985, 1986), Stark and 
Ray (1983), Stark and Stewart (1981), Stark 
and Szczytko (1981), Stewart and Stark 
(1984), Surdick (1985), and Szczytko and 
Stewart (1976). 

List of Stoneflies of Virginia 

Order Plecoptera 

Suborder Arctoperlaria 

Group Euholognatha 


Family Nemouridae 
Subfamily Amphinemurinae 

Amphinemura delosa (Ricker) WB 
A. nigritta (Provancher) WB 
A. wui (Claassen) AP 

Subfamily Nemourinae 

Ostrocerca albidipennis (Walker) AP 

# O. complexa (Claassen) AP 

# O. prolongata (Claassen) AP 
O. truncata (Claassen) AP 
Paranemoura perfecta (Walker) AP 



Prostoia completa (Walker) WB 

+ P. hallasi Kondratieff & Kirchner CP 

P. similis (Hagen) WB 

Shipsa rotunda (Claassen) WB 

Soyedina carolinensis (Claassen) AP 

S. vallicularia (Wu) AP 

* Zapada chila (Ricker) (TN) AP 

Family Taeniopterygidae 
Subfamily Brachypterinae 

Bolotoperla rossi (Prison) AP 
Oemopteryx contorta (Needham & Claas- 
sen) AP 

* O. glacialis (Newport) (WV) WB 
Strophopteryx appalachia Ricker & Ross AP 
S. fasciata (Burmeister) WB 

S. limata (Prison) AP 

Taenionema atlanticum Ricker & Ross AP 

Subfamily Taeniopteryginae 

Taeniopteryx burksi Ricker & Ross WB 

* T. lita Prison CP 

T. lonicera Ricker & Ross WB 

T. mama (Pictet) WB 

T. metequi Ricker & Ross WB 

+ T. nelsoni Kondratieff & Kirchner AP 

T. parvula Banks WB 

T. ugola Ricker & Ross AP 

Pamily Capniidae 

Allocapnia aurora Ricker AP 

* A. brooksi Ross (TN) AP 
A. curiosa Prison AP 

* A. forbesi Prison (KY, TN, WV) WB 
A. frisoni Ross & Ricker AP 

* A. frumi Kirchner (WV) AP 
A. fumosa Ross AP 

A. granulata (Claassen) WB 

+ A. harperi Kirchner AP 

A. illinoensis Prison WB 

A. loshada Ricker AP 

A. maria Hanson AP 

A. mystica Prison AP 

A. nivicola (Pitch) WB 

A. pygmaea (Burmeister) WB 

A. recta (Claassen) WB 

A. rickeri Prison WB 

+ A. simmonsi Kondratieff & Voshell AP 

A. stannardi Ross AP 

A. virginiana Prison AP-CP 

A. vivipara (Claassen) WB 

A. wrayi Ross AP-CP 

A. zola Ricker AP 

Nemocapnia Carolina Banks WB 

Paracapnia angulata Hanson AP 

Pamily Leuctridae 
Subfamily Megaleuctrinae 

Megaleuctra flinti Baumann AP 

# M. williamsae Hanson AP 

Subfamily Leuctrinae 

Leuctra alexanderi Hanson AP 

L. grandis Banks/biloba Claassen AP 

L. carolinensis Claassen AP 

L. duplicata Claassen AP 

L. ferruginea (Walker) WB 

L. mitchellensis Hanson AP 

L. monticola Hanson AP 

# L. nephophila Hanson (NC, TN) AP 

# L. rickeri James WB 
L. sibleyi Claassen AP 

# L. tenella Provancher WB 
L. tenuis (Pictet) WB 

L. triloba Claassen AP 
L. truncata Claassen AP 
L. variabilis Hanson AP 
Paraleuctra sara (Claassen) WB 

Group Systellognatha 


Pamily Pteronarcyidae 

Pteronarcys biloba Newman AP 
P. comstocki Smith AP 
P. proteus Newman AP 
P. scotti Ricker AP 
P. dorsata (Say) WB 

Superfamily Peltoperloidea 
Pamily Peltoperlidae 
Subfamily Peltoperlinae 

Peltoperla arcuata Needham AP 
P. tarteri Stark & Kondratieff AP 
Tallaperla anna (Needham & Smith) AP 

# T. Cornelia (Needham & Smith) AP 

# T. laurie (Ricker) AP 



+ T. lobata Stark AP 

T. maria (Needham & Smith) AP 

* Viehoperla ada (Needham & Smith) (NC, 


Family Perlodidae 
Subfamily Isoperlinae 

Clioperla clio (Newman) WB 
Isoperla bilineata (Say) WB 

* /. bellona Banks (NC, TN) AP 
/. burksi Prison WB 

* /. cotta Ricker (WV) WB 

* /. coushatta Szczytko & Stewart CP 
/. dicala Prison WB 

* /. distincta Nelson (NC, TN) AP 

* /. gibbsae Harper (WV) AP 
/. holochlora (Klapalek) AP 
/. lata Prison WB 

+ /. major Nelson & Kondratieff AP 
/. marlynia Needham & Claassen WB 
/. montana (Banks) AP 
/. orata Prison AP 

* /. richardsoni Prison (WV) WB 
/. signata (Banks) WB 

/. similis (Hagen) AP 
/. slossonae (Banks) WB 

* /. transmarina (Newman) WB 

Subfamily Perlodinae 

Cultus decisus (Walker) WB 
Diploperla duplicata (Banks) AP-CP 

* D. kanawholensis Kirchner & 
Kondratieff AP 

D. morgani Kondratieff & Voshell AP 
D. wbusta Stark & Gaufin WB 
Helopicus subvarians (Banks) AP-CP 
Isogenoides hansoni (Ricker) AP 

* /. varians (Walsh) WB 
Malirekus hastatus (Banks) AP 

* Oconoperla innubila (Needham & Claas- 
sen) (NC, TN) AP 

Remenus bilobatus (Needham & 

Claassen) AP 
Yugus arinus (Prison) AP 
Y. bulbosus (Prison) AP 

Family Chloroperlidae 
Subfamily Paraperlinae 

* Utaperla gaspesiana Harper & Roy 
(WV) AP 

Subfamily Chloroperlinae 

* Alloperla aracoma Harper & Kirchner 
(WV) AP 

A. atlantica Baumann AP 

A. banksi Prison WB 

A. biserrata Nelson and Kondratieff AP 

A. caudata Prison WB 

A. chloris Prison AP 

A. concolor Ricker AP 

A. idei (Ricker) AP 

A. imbecilla (Say) WB 

A. nanina Banks AP 

* A. neglecta Prison (NC, TN) AP 
A. usa Ricker AP 

Haploperla brevis (Banks) WB 
Suwallia marginata (Banks) AP 
Sweltsa lateralis (Banks) AP 
S. mediana (Banks) AP 
S. naica (Provancher) AP 
S. onkos (Ricker) WB 
S. urticae (Ricker) AP 

* Rasvena terna (Prison) (NC, TN, WV) AP 

Family Perlidae 
Subfamily Acroneuriinae 

Acroneuria abnormis (Newman) WB 
A. arenosa (Pictet) CP 

* A. arida (Hagen) (TN) WB 
A. carolinensis (Banks) AP 
A. evoluta Klapalek WB 

A. ft I ids Prison WB 

A. flinti Stark & Gaufin AP 

* A. internata (Walker) WB 
A. lycorias (Newman) WB 

* A. perplexa Prison (TN, WV) WB 

* A. peter si Stark & Gaufin (NC, TN) AP 
Attaneuria ruralis (Hagen) WB 

* Beloneuria stewarti Stark & Szczytko (NC, 

Eccoptura xanthenes (Newman) AP-CP 

* Hansonoperla appalachia Nelson AP 



Perlesta placida (Hagen) "complex" WB 
Perlinella drynno (Newman) WB 
P. ephyre (Newman) WB 

Subfamily Perlinae 

# Neoperla carlsoni Stark & Baumann CP 

# N. catharae Stark & Baumann WB 

# A^. clymene (Newman) WB 

# N. Choctaw Stark & Baumann (WV) WB 
TV. freytagi Stark & Baumann WB 

# A^. Stewart! Stark & Baumann WB 
Paragnetina fumosa (Banks) AP-CP 
P. immarginata (Say) AP 

# P. ichusa Stark & Szczytko AP 
P. media (Walker) AP 

# Agnetina annulipes (Hagen) CP 
A. capital a (Pictet) WB 

# A. flavescens (Walsh) WB 


We thank the following persons for mak- 
ing specimens available for study: M. I. Bass, 
O. S. Flint, Jr., R. L. Hoffman, R. E. Jen- 
kins, M. Kosztarab, and J. R. Voshell, Jr. 
Rebecca F. Surdick kindly provided the 
Hansonoperla record. Stanley W. Szczytko 
allowed us to include several Isoperla rec- 
ords from Virginia. Bill P. Stark verified 
several species determinations. The base 
map of Virginia was kindly provided by R. 
E. Jenkins, Roanoke College. It is a modi- 
fication of the U.S.G.S. state of Virginia, 
scale 1: 500,000 map, 1957 edition. 

Literature Cited 

Harper, P. P. and K. W. Stewart. 1984. Plecoptera. 
Chapter 13, pp. 182-230. In Merritt, R. W. and 
K. W. Cummins, eds.. An introduction to the 
aquatic insects of North America. 2nd Edition. 
Kendall/Hunt Publ. Co. Dubque, Iowa. 

Hoffman, R. L. 1969. The insects of Virginia. No. 1. 
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Kirchner, R. F. 1980. A new Allocapnia from Vir- 
ginia (Plecoptera: Capniidae). Entomol. News 91: 

. 1982. A new Allocapnia from West Virginia 

(Plecoptera: Capniidae). Proc. Entomol. Soc. Wash. 
84: 786-790. 

Kirchner, R. F. and P. P. Harper. 1983. The nymph 
of Bolotoperla rossi (Prison) (Plecoptera: Taenio- 
pterygidae: Brachypterinae). J. Kans. Entomol. Soc. 

Kirchner, R. F. and B. C. Kondratieff. 1984. Anew 
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. 1985. The nymph of Hansonoperla appala- 

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Kondratieff, B. C. and R. F. Kirchner. 1982a. Tae- 
niopteryx nelsoni, a new species of winter stonefly 
from Virginia (Plecoptera: Taeniopterygidae). J. 
Kans. Entomol. Soc. 55: 1-7. 

. 1982b. Notes on the winter stonefly genus 

Allocapnia (Plecoptera: Capniidae). Proc. Ento- 
mol. Soc. Wash. 84: 240-244. 

Kondratieff B. C. and J. R. Voshell, Jr. 1979. A 
checklist of the stoneflies (Plecoptera) of Virginia. 
Entomol. News. 90: 241-246. 

. 1981. Allocapnia simmonsi. a new species of 

winter stonefly (Plecoptera: Capniidae). Ann. 
Entomol. Soc. Am. 74: 58-59. 

. 1982. The PerlodinaeofVirginia, USA (Ple- 
coptera: Perlodidae). Proc. Entomol. Soc. Wash. 
84: 761-774. 

Kondratieff, B. C, R. F. Kirchner, and J. R. Voshell, 
Jr. 1981. Nymphs of D/^/o/Jpr/a. Ann. Entomol. 
Soc. Am. 74: 428-430. 

Nelson, C. H. 1 979. Hansonoperla appalachia, a new 
genus and a new species of eastern nearctic Ac- 
roneuriini (Plecoptera: Perlidae), with a phenetic 
analysis of the genera of the tribe. Ann. Entomol. 
Soc. Am. 72: 735-739. 

. 1982. Notes on the life histories of S/wp/zo- 

pteryx limata (Prison) and Oemopteryx contorta 
(Needham and Claassen) (Plecoptera: Taeniop- 
terygidae) in Tennessee. J. Tenn. Acad. Sci. 57: 

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major, a new species of eastern nearctic Isoperli- 
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Am. 76: 270-273. 

Stark, B. P. 1979. The stoneflies (Plecoptera) of Mis- 
sissippi. J. Miss. Acad. Sci. 24: 109-122. 

. 1983. The Tallaperla maria comxAex of easX- 

em North America (Plecoptera: Peltoperlidae). J. 
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lidae). J. Kans. Entomol. Soc. 59: 437-445. 

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genus Helopicus (Plecoptera: Perlodidae). Fresh- 
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6: 71-79. 


89(1), 1987, pp. 31^2 





Michael E. Schauff 

Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % 
U.S. National Museum, NHB 168, Washington, D.C. 20560. 

Abstract.— The egg parasitoids of citrus weevils are reviewed, and an illustrated key is 
presented. Nine species of Hymenoptera in two superfamilies (Chalcidoidea: Eulophidae, 
Mymaridae, and Trichogrammatidae and Proctotrupoidea: Platygastridae) are now known 
to be primary or secondary parasites in these weevil egg masses. Two new species, Ho- 
rismenus bennetti and Tetrastichus fennahi, both eulophids, are described and figured. 
Additional data on the known hosts and distributions of the parasitoids are given. 

Weevils in the genera Pachneus, Exop- 
thalmus, Artipus, and Diaprepes (Curcu- 
lionidae) feed on a variety of plants. Adults 
are leaf feeders and larvae feed on and bore 
into the roots of the host plants. When these 
weevils attack citrus trees, they are collec- 
tively called citrus weevils and are capable 
of causing substantial economic damage. 
Larvae can even kill citrus by girdling the 

Since 1 964, when specimens oi Diaprepes 
abbreviatus (L.) were discovered in citrus 
groves in the vicinity of Apopka, Florida 
(Woodruff, 1964), several attempts to es- 
tablish effective biological control programs 
have been made (e.g. Sutton et al., 1972; 
Beavers and Selhime, 1975). Unfortunately, 
these efforts have met with limited success. 

The weevil is native to Puerto Rico and 
the West Indies (Woodruff, 1968), but the 
exact route by which it became introduced 
into continental United States is unknown. 
In Puerto Rico, where D. abbreviatus has 
been a pest of sugarcane (commonly known 
as the sugar-cane root-stalk borer weevil), 
it has been effectively suppressed by an egg 

parasite, Tetrastichus haitiensis Gahan 
(Wolcott, 1948). This parasite has been the 
subject of most of the efforts to establish a 
biological control agent in the U.S. How- 
ever, several other parasites have been 
reared from the eggs of citrus weevils and 
the inability to identify and correctly assign 
specific names to these species has ham- 
pered efforts at control (R. Woodruff, pers. 
comm.). In addition, it is now apparent that 
some of these species are undescribed. 

I take this opportunity to clarify the tax- 
onomy of these species and provide a key 
to enable field workers to identify them. 
Much information about the biology and 
interrelationships of these species remains 
to be discovered and a sounder taxonomy 
should enable that work to proceed. Listed 
hosts for the parasites refer only to known 
species of citrus weevils and, in the case of 
hyperparasites, to hosts associated with cit- 
rus weevils. 

The majority of specimens used in this 
study were collected by R. Woodruff, E. E. 
Grissell and F. D. Bennett. They and the 
types are deposited in either the Florida State 



Collection of Arthropods at Gainesville 
(FSCA), the British Museum of Natural 
History (BMNH) or the U.S. National Mu- 
seum, Washington, D.C. (USNM). 

Morphological terminology follows that 
of Graham ( 1959), except that I use the term 
metasoma to refer to that part of the ab- 
domen past the propodeum (often called the 
gaster) and metanotum rather than meta- 
scutellum. Terms for sculpturing follow 
Harris (1979). 

Key TO THE Parasites of Citrus 
Weevil Eggs 

1 . Forewings nearly parallel sided (Fig. 5) and with 
marginal cilia longer than width of wing; fe- 
male antennae with 6 funicle articles and club 
entire (Fig. 14); male antennae with 13 articles 
Cleruchus sp. 

- Forewings not parallel sided (Figs. 1-4) and 
with marginal cilia much shorter than wing 
width; female antennae with fewer than 6 fu- 
nicle articles and club when present with 2 or 
3 articles (Figs. 6-1 1); male antennae with few- 
er than 1 3 articles 2 

2. Tarsi 5-segmented; forewing with only sub- 
marginal vein (Fig. 1); antenna with 4 small 
funicular articles (Fig. 6) and a 3-segmented 
club; scutellum without paired setae (Fig. 1 5) 
Platystasius cith Nixon 

- Tarsi 3 or 4-segmented; forewing with sub- 
marginal, marginal and stigmal veins (Figs. 2- 
4); antennae with fewer than 4 funicular arti- 
cles or club not 3-segmented (Figs. 7-1 1); scu- 
tellum with 1 or two pairs of setae (Figs. 16, 
20) 3 

3. Tarsi 3-segmented; forewings with obvious se- 
tal tracts radiating from stigmal vein and other 
wing veins as in Fig. 2; female antenna as in 

Fig. 10; body length about 0.8 mm 

Brachyufens osborni (Dozier) 

- Tarsi 4-segmented; forewings without setal 
tracts, wing veins as in Figs. 3, 4; antennae as 
in Figs. 7-9,. 1 1; body length greater than 0.9 
mm 4 

4. Scutellum with 2 longitudinal submedian 
grooves and 2 pairs of setae (Figs. 16, 17); ax- 
illae advanced forward of anterior edge of scu- 
tellum; clypeus bilobed (Fig. 24); propodeum 
narrowed medially, only about as wide as 
metanotum, with a simple median carina which 
may be incomplete (Fig. 23); metasoma broad- 
ly joined to propodeum, petiole not apparent 

- Scutellum without grooves or with only a single 

median groove (Figs. 1 8-20), with only a single 
pair of setae; axillae not advanced past anterior 
edge of scutellum, clypeal margin straight; pro- 
podeum wider than metanotum; metasoma 
petiolate 5 

5. Scutellum with median longitudinal groove 
(Figs. 18, 19); propodeum with median raised 
area flanked by submedian carinae (Fig. 22) . . 6 

- Scutellum without median groove (Fig. 20), 
propodeum with a pair of parallel submedian 
carinae (Fig. 21) ... Pediobius irregularis Kerrich 

6. Notaulices complete, well defined (Fig. 18); first 
metasomal tergum with narrow reticulate band 
of sculpture posteriorly (Fig. 1 2); body metallic 

green; male femora and scape yellow 

Horismenus cupreus (Ashmead) 

- Notaulices faint, incomplete (Fig. 19); first 
metasomal tergum nearly smooth or with faint 
open crack-like sculpture on posterior half (Fig. 
1 3); body black; male femora and scape brown 

to black Horismenus bennetti sp. nov. 

7. Most of head and metasoma yellow, rest of 
body often with extensive areas of yellow, may 
be nearly entirely yellow; male flagellar articles 

with long whorled setae (Fig. 11) 

Tetrastichus gala (Walker) 

- Body dark brown or blackish (except legs yel- 
low); male flagellar articles without long whorled 
setae (Fig. 8) 8 

8. Submarginal vein with a single seta (rarely 2); 
mesoscutum midlobe with single row of setae 
(Fig. 1 7); body without metallic greenish lustre; 
base of metasoma yellow; propodeum smooth 
and without carinae arising from nucha .... 
Tetrastichus haitiensis Gahan 

- Submarginal vein with 3 or 4 setae (Fig. 4); 
mesoscutum with an irregular second row of 
setae (Fig. 16); body with metallic greenish 
lustre; propodeum sculptured and with carinae 

arising from nucha (Fig. 21) 

Tetrastichus fennahi sp. nov. 


Brachyufens osborni (Dozier) 

Figs. 2, 10 

Ufens osborni Dozier, 1932: 36. 

Notes.— This species was described from 
specimens reared from Diaprepes abbrevi- 
atus eggs in Puerto Rico. In his original de- 
scription, Dozier noted that osborni was 
somewhat different than other species of 
Ufens, but decided that it still fell within the 
limits of that genus. Doutt and Viggiani 
(1968) noted several differences between the 



Figs. 1-5. Forewings. 1, Platystasius citri. 2, Brachyufens osborni. 
fennahi. 5, Cleruchus sp. Scale line equals 0.1 mm. 

3, Horismenus bennetti. 4, Tetrastichus 

Other species of Ufens and osborni and 
transferred it to their new genus Brachyu- 
fens as the type species. B. osborni has been 
reared from Pachnaeus lit us eggs as well as 
species of Diaprepes. It has been reported 
to be quite common at times and has parasi- 
tized as many as 8 1 % of the eggs of P. litus 
(Baranowski, 1960). In view of the known 
biology of other trichogrammatids, it is 

doubtful that reports of this species as a 
possible secondary parasite (Burks, 1979) 
are correct. 

Diagnosis.— As the only trichogramma- 
tid reared from citrus weevil eggs, this 
species is quite easily separated from the 
other parasites by the following characters: 
tarsi 3 -segmented (4 or 5 segmented in oth- 
ers), forewings with distinct setal tracts (no 



Figs. 6-14. 6-11, Antennae. 6, Platystasius citri, female. 7, Horismenus bennetti, female. 8, Tetrastichus 
haitiensis, male. 9, Tetrastichus fennahi female. 10, Brachyufens osbomi, female. 11, Tetrastichus gala, male. 
12, 13, Metasomas. 12, Horismenus cupreus. 13, Horismenus bennetti. 14, Cleruchus sp., female antenna. Scale 
line equals 0.1 mm. 

tracts in other species) and venation as in 
Fig. 2, length about 0.8 mm and female an- 
tennae as in Fig. 10 (other antennae as in 
Figs. 6-9). 

WosXs.—Pachneus litus (Germar), P. opa- 
lus (Olivier), Diaprepes abbreviatus (L.). 

Specimens examined. — Dominican Re- 
public: San Cristobal Prov., San Cristobal. 
Puerto Rico: Mayaguez, University Cam- 
pus; Maricao. Andros Island, San Andros. 
Montserrat, Plymouth. United States. Ror- 
ida: Dade Co., Homestead; Palm Beach Co., 
West Palm Beach; Indian River Co., Vero 

Beach; St. Lucie Co.; Ft. Pierce, Hardee Co., 
Ft. Green. Collection dates range from 14 
June through 3 October. 


Tetrastichus haitiensis Gahan 

Figs. 8, 17 

Tetrastichus haitiensis Gahan, 1929: 17. 

Notes.— This species was described from 
a series of specimens reared from Exop- 
thalmus quadrivittatus eggs at Port-au- 
Prince, Haiti. It is a primary parasite of the 



Figs. 15-20. SEMs. Thorax, dorsal view. 15, Platystasius citri. 16, Tetrastichus fennahi. 17, Tetrastichus 
haitiemis. 18, Horismenus cupreus. 19, Horismenus bennetti. 20, Pediobius irregularis. 

eggs and has also been reared from Dia- 
prepres abbreviatus and Pachnaeus litus. It 
is apparently the commonest parasite of cit- 
rus weevil eggs in this area and at times may 
kill up to 100% of the eggs and egg masses 
(Van Whervin, 1968). As a result, it has 
received the most attention from biological 
control workers and has been imported and 
released into Florida from the West Indies 

on several occasions (Sutton et al., 1972; 
Beavers and Selheime, 1975). A few spec- 
imens have recently been recovered from 
citrus weevil eggs in Central Florida (C. Tar- 
rant, pers. comm.). 

Diagnosis. — This species is most easily 
confused with Tetrastichus gala or T. fen- 
nahi. It differs in the following: the body is 
generally dark brown without a greenish 



tinge and with the legs and base of the 
metasoma yellow (body extensively yellow 
in gala and body with greenish tinge and 
base of metasoma not yellow in fennahi); 
submarginal vein usually with only 1 seta 
(submarginal with 3 or more in others); se- 
tae on male funicles about 2 x as long as 
width of segment (Fig. 8) (shorter in fennahi 
and much longer and whorled in gala. Fig. 
1 1); propodeum smooth and without para- 
spiracular carinae (propodeum sculptured 
in others and with paraspiracular carinae in 
fennahi); mid lobe of scutum with only a 
single row of setae laterally (Fig. 1 7) {T. fen- 
nahi with partial second row, Fig. 16). 

Variability. — Females may occasionally 
have some yellow or very light brown areas 
around the scrobes, occelli, and notauli. 
There may also be some light brown on the 
femora. A few males have been observed 
with much shorter setae on the funicular 
articles than is typical for this species and 
the expanded ridge on the anterior surface 
of the scape is present although not as no- 
ticeable as in males with elongated funicular 

WosXs.—Diaprepes abbreviatus (L.), Ex- 
opthalmus quadrivittatus (Olivier), Pach- 
neus litus (Germar), and P. opalus (Olivier). 

Specimens examined. — Dominican Re- 
public: La Romana Prov., Cacata; San Cris- 
tobal Prov., San Cristobal; Republic Prov., 
Monte Cristi, 4 Km. N. Villa Vasquez. 
Puerto Rico: Isabela; Mayaguez, University 
Campus; Ponce, Fortuna Fruit Experiment 
Station. Andros Island, San Andros. Ja- 
maica: St. Catherine Parish, Charlton nr. 
Ewarton; Red Hill. Cuba, Santiago, D. I. 
Vegas. Haiti, Port-au-Prince. United States. 
Florida, Hardee Co., Ft. Green. Collection 
dates range from March through October. 

Tetrastichus gala (Walker) 

Figs. 11, 24 

Tetrastichus gala Walker, 1847: 28. 

Notes.— This species has been misiden- 
tified as T. marylandensis Girault ( 1 9 1 6) (Z. 

Boucek, pers. comm.) and the exact rela- 
tionship of these two species remains in 
doubt. I can find little difference between 
specimens of the two species, but the host 
ranges indicate that two species may indeed 
be involved. T. marylandensis is recorded 
as a parasite of lepidopterous larvae, aphids, 
and midges as well as from the eggs of citrus 
weevils. When associated with citrus wee- 
vils, T. gala (cited as marylandensis) has 
been cited as an"egg predator" feeding ex- 
ternally in the weevil egg masses (Van 
Whervin, 1968). I have studied specimens 
identified as both marylandensis and gala 
reared from various hosts, but many are in 
such poor condition that definitive identi- 
fication to species is not possible. Since this 
paper is not revisionary in scope, I am re- 
luctant to propose the synonymy of mary- 
landensis with gala. Until further specimens 
from the various hosts can be obtained or 
a comprehensive revision is undertaken, I 
believe it better to leave the two names as 
they are. 

Diagnosis.— This species is most easily 
confused with Tetrastichus fennahi and T. 
haitiensis. It can be separated by the follow- 
ing: submarginal vein with 3-5 setae (sub- 
marginal with only one seta or rarely 2 in 
T. haitiensis); body with at least some yel- 
low markings on the face, venter of thorax, 
and metasoma and often nearly entirely yel- 
low (body entirely dark brown or blackish 
except for the legs or base of the metasoma 
in other Tetrastichus species); male funic- 
ular articles with greatly lengthened, whorled 
setae (Fig. 1 1) (funicular setae much shorter 
in other Tetrastichus as in Fig. 8). 

Variability.— This species shows marked 
variation in coloration. Some specimens are 
almost entirely yellow with only a little 
brown on the edges of the metasomal terga, 
posterior margin of the propodeum, ante- 
rior edge of the pronotum, and around the 
foramen of the head. At the other extreme 
a series of specimens from Jamaica has al- 
most the entire dorsum of the thorax dark 
brown, and has large brown areas on the 



Figs. 2 1-24. SEMs. 2 1-23, Propodeums. 2 1 , Pediobius irregularis. 22, Horismenus bennetti. 23, Tetrastichus 
fennahi. 24, Face of Tetrastichus gala. 

metasoma, side of the thorax, and head. 
Variation in color ranges almost the whole 
spectrum between these two extremes. The 
face and frons are apparently always light 
colored, as is the venter of the thorax. 

Hosts.— Diaprepes abbreviatus (L.), D. 
famelicus (Olivier). 

Specimens examined. — Jamaica, St. 
Catherine Parish, Charlton nr. Ewarton. 
Montserrat, Plymouth. Guadeloupe, Do- 
main Duclos. Puerto Rico, Isabela, Maya- 
quez. Dominica. Collection dates range from 
December through June. 

Tetrastichus fennahi SchauflF, New Species 

Figs. 4, 9, 16, 23 

Holotype female. — Length 1.2 mm. Col- 
or as follows: body dark brown with me- 
tallic green reflections; basal half of hind 
coxae light brown; rest of legs, tegulae, and 

antennae yellow. Head slightly wider than 
thorax, about as high as wide; face, frons, 
occiput with scattered silvery setae; frons 
with scattered small setigerous punctures 
near eye margins; occiput lightly imbricate; 
POL 2.5 X OOL; malar space V2 eye height; 
malar suture complete, curved; antenna in- 
serted on line with ventral margin of eye; 
scape not quite reaching level of tip of eye; 
antenna as in Fig. 9; scutum (Fig. 16) im- 
bricate, median longitudinal groove com- 
plete; midlobe with 2 irregular lines of setae; 
slightly longer than scutellum (18:15); scu- 
tellum as long as wide, propodeum (Fig. 23) 
imbricate, with several irregular small cari- 
nae projecting anteriorly from nucha, me- 
dially only about as wide as metanotum, 
paraspiracular carinae complete, with only 
a single seta laterad of spiracle; metasoma 
slightly longer than thorax; first tergum 



smooth medially, becoming imbricate lat- 
erally, other terga imbricate; ovipositor 
sheaths cylindrical, barely protruding past 
tip of metasoma; forewing slightly more than 
2x as long as wide (43:20) (Fig. 4); sub- 
marginal vein with 3 setae; ratio submar- 
ginal: marginal : stigmal 15:25:7; tip of 
hindwing rounded, marginal fringe V3 width 
of hindwing at hamulus. 

Male.— The only available male speci- 
men is very badly shriveled and no obvious 
differences from the female (except for the 
genitalia) can be discerned. 

Diagnosis.— This species is most easily 
confused with T. haitiensis and gala. It can 
be differentiated from them and from other 
Tetrastich us species by the following: body 
dark brown with greenish tinge (at least face 
and venter of thorax yellow in T. gala, no 
greenish tinge in T. haitiensis or base of 
metasoma yellowish); mid lobe of scutum 
with partial second row of setae (Fig. 1 6) 
(mid lobe with only a single row of setae in 
other species (as in Fig. 17); submarginal 
vein with 3-5 setae (submarginal with only 
1 or rarely 2 setae in T. haitiensis); propo- 
deum imbricate and with carinae arising 
from nucha (Fig. 23), paraspiracular carina 
present (other species with propodeum 
mostly smooth, without carinae, paraspi- 
racular carina absent); male flagellum with 
setae only about as long as width of segment 
(male flagellar setae about 2x as long as 
width of segment in haitiensis (Fig. 8), much 
longer and whorled in T. gala (Fig. 1 1). 

Variability. — Very little variation was 
observed in the specimens available for 
study. The body length ranges from 1 .0-1 .3 
mm. Eye color varies from bright red to 
silver or grayish. In a few specimens, the 
hind coxae are nearly entirely yellow. The 
majority of the specimens are badly shriv- 
eled and the propodeum is collapsed mak- 
ing it difficult to see if the paraspiracular 
carinae are as evident as in the type female 
and in a specimen that was photographed 
with the scanning electron microscope. In 
some of these poor specimens it appears 

that the carina is absent and caution should 
be used when assessing this character. 

Types. — Holotype $ on point with data: 
St. Lucia, B.W.I., 1937, R. G. Fennah, Ex. 
Diaprepes abbreviatus eggs. Forewing and 
antenna slide mounted. Deposited in U.S. 
National Museum of Natural History. Para- 
types: 1 7 9, 1 5 same data as holotype; 1 9 
Barbados, W. I., ex. eggs of^ Diaprepes ab- 
breviatus on citrus; 5 9 Machourie, Domin- 
ica, B.W.I., June, 1954, Coll. F. D. Bennett, 
ex. ova Diaprepes on legume; 1 9 same data 
as above except collected at Grand Savan- 
nah [Grande Savanne], July, 1954; 2 2 Ja- 
maica, St. Catherine Parish, Charlton nr. 
Ewarton, 19-VI-1975, em. 29-VI-1975. E. 
E. Grissell, R. E. Woodruff, ex. Exopthal- 
mus or Pachneus eggs. Paratypes deposited 
in the USNM except for 2 9 each deposited 
in British Museum, Canadian National Col- 
lection, and Horida State collection of Ar- 

Hosts.— Diaprepes abbreviatus (L.). 

Etymology.— This species is named in 
honor of the collector of the holotype, R. 
G. Fennah. 

Pediobius irregularis Kerrich 

Figs. 20, 21 

Pediobius irregularis Kervich, 1973: 190. 

Notes.— This species was described from 
specimens reared from the egg mass ofEx- 
opthalmus viticollis Champion on citrus by 
L. W. Van Whervin taken in Belize (British 
Honduras). Little else is known of this 
species and it apparently has not been col- 
lected in the West Indies where most of the 
collecting by biological control workers has 
taken place. Kerrich listed this species as a 
primary parasite of the weevil eggs. 

Diagnosis.— This species is most easily 
confused with the two species of Horisme- 
nus. It can be separated by the following: 
scutellum without median groove (Fig. 20) 
(groove present in Horismenus, Fig. 19); 
propodeum with a pair of parallel subme- 
dian carinae (Fig. 21) (propodeum medially 



with a shiny raised area in Horismenus, Fig. 

Hosts.— Exopthalmus vitticollis Cham- 

Specimens examined. — Paratypes of P. 
irregularis and an additional short series of 
specimens collected a year earlier by the 
same collector (L. W. Van Whervin) who 
collected the types series (from the same 
locality). This species is known only from 
the type locality (Belize). 

Horismenus cupreus (Ashmead) 

Figs. 12, 18 

Holcopelte cupreus Ashmead, 1894: 171. 

Notes. — The types of this species were 
collected on St. Vincent from unknown host 
(specimens deposited in BMNH). There are 
additional specimens in the USNM labelled 
as reared from eggs of Diaprepes famelicus 
esuriens on Montserrat. Whether it is a pri- 
mary or secondary parasite of the eggs is 

Diagnosis.— This species is most easily 
confused with H. bennetti or Pediobius ir- 
regularis. It is easily separated from Pe- 
diobius by the presence of a longitudinal 
groove on the scutellum and the raised 
smooth median area of the propodeum (as 
in Figs. 19, 22) (scutellar groove absent in 
Pediobius and median propodeum with 
paired carinae, Figs. 20, 21). //. cupreus d\{- 
fers from bennetti by having complete and 
well defined notauli (Fig. 1 8), the first meta- 
somal tergum with a narrow sculptured band 
(Fig. 12), the body is black, and the male 
scape and femora are yellow (in bennetti, 
the notauli are not well defined (Fig. 19), 
the first tergum usually has only minute 
cracklike sculpture (Fig. 13), the body is 
black, and the male femora and scape are 
brown or black). 

WosXs. — Diaprepes famelicus esuriens 

Specimens examined.— Type of H. cu- 
preus and other specimens from St. Vincent, 
and Montserrat. 

Horismenus bennetti Schauff, New Species 

Figs. 3, 7, 13, 19, 22 

Holotype female. — Length 1.6 mm. Col- 
or black except the following: scape, legs 
past coxae yellow; last tarsomere brownish 
apically. Antennae as in Fig. 7; apex of scape 
even with arms of frontal forks at margin 
of eye; face below toruli lightly imbricate; 
area laterad of scrobes and below frontal 
grooves more strongly sculptured, nearly al- 
veolate; between scrobes smooth; frons me- 
dially above frontal forks very lightly im- 
bricate, nearly smooth; vertex imbricate; 
genae smooth to very faintly strigate, oc- 
ciput alveolate; POL 3 x OOL; pronotum 
imbricate except smooth along posterior 
margin; scutum and scutellum imbricate as 
in Fig. 19; notaulices fading anteriorly; me- 
dian scutellar groove nearly reaching pos- 
terior margin of scutellum; with row of small 
alveolae extending posteriorly from scutel- 
lar setae and curving inwards near margin; 
metanotum smooth; propodeum (Fig. 22) 
smooth except at posterior edge and laterad 
of nucha; petiole in dorsal view slightly 
longer than wide (13:10), rugulose; prepec- 
tus imbricate, mesopleuron smooth; meta- 
soma equal in length to thorax, nearly 2 x 
as long as wide (60:35); first tergum covering 
-h length, smooth except for small postero- 
medial patch of very fine crack-like acicu- 
lations (Fig. 13); forewing as in Fig. 3. 

Male. — Similar to female except the fol- 
lowing: length about 1.1 mm; scape and 
femora brown to black; basal half of tibiae 
occasionally light brown; scape 3 x as long 
as wide, funicular articles covered by nu- 
merous white setae; petiole 2 x as long as 
wide; metasoma ovate, only about as wide 
as long. 

Diagnosis.— This species is most easily 
confused with Pediobius irregularis and H. 
cupreus. It can be separated from Pediobius 
by the presence of a longitudinal groove on 
the scutellum and the raised smooth median 
area of the propodeum (as in Figs. 19, 22) 
(scutellar groove absent in Pediobius and 



median propodeum with paired carinae, 
Figs. 20, 21). It differs from H. cupreus by: 
the black body color {cupreus and many oth- 
er species are metallic green); the notauilces 
are faint and incomplete (Fig. 19) (notau- 
lices well defined in cupreus, Fig. 18); the 
first metasomal tergum nearly smooth, with 
only faint crack-like sculpture (Fig. 1 3) (nar- 
row reticulate band in cupreus, Fig. 1 2); male 
femora and scape brown to black (male scape 
and femora of cupreus yellow). Additional 
characters which help to separate this species 
from other Horismenus are: propodeum 
mostly smooth and without reticulation near 
nucha (many other species have some re- 
ticulate sculpture on the propodeum); legs 
of the female yellow past coxae (several 
species have the femora and/or tibiae dark 
colored); small row of alveolae adjacent to 
scutellar setae and scutellar surface nearly 
completely smooth (alveolae lacking and 
scutellar surface sculptured in some other 

Variability.— There is some variation in 
the appearance of the sculptured area on the 
first metasomal tergum. In one specimen, it 
appears as more of a reticulated pattern 
somewhat similar in appearance to that in 
cupreus. However, in this specimen, the 
band of sculpture was quite wide (about half 
as wide as long), while in cupreus the band 
is very narrow (only about a sixth as wide 
as long). 

Types. — Holotype 9 on point with data: 
Jamaica, Red Hill, May 1956. Curculionid 
eggs on citrus. Coll. by F. D. Bennett. De- 
posited in the U.S. National Museum of 
Natural History. Paratypes: 3 9; Puerto Rico, 
Isabela, VI- 1 6- 1932, G. N. WoUcott, ex. eggs 
Diaprepes abbreviatus; 1 9, 1 3, Jamaica, 
B.W.I., July 1954, Ova Prepodes, coll. by 
Simmonds; 1 9, Jamaica, Mona, Dec. 1, 
1967, ex. eggs of Tetrastichus parasite on 
citrus weevil eggs, coll. by Van Whervin; 5 
$ and 4 9, Dominican Republic, San Cris- 
tobal, 23-VI-1976, on citrus, emerged 24- 
VI- 1976; 8 9 and 3 5, Jamaica, Parish of St. 
Catherine, Charlton. Exp. Sta., 19-VI-1975. 

Grissell & Woodruff. Ex. Exopthalmus eggs 
on citrus, em. 25-VI-1975. One 5 and 1 9 
paratype deposited in BMNH and FSCA, 
rest in USNM. 

Hosts.— The species is apparently a hy- 
perparasite. Its most likely host is Tetra- 
stichus haitiensis, although it may also para- 
sitize other Tetrastichus. 

Etymology.— This species is named in 
honor of F. D. Bennett who collected many 
of the specimens used in this study. 


Platystasius citri Nixon 

Figs. 1,6, 15 

Platystasius citri Nixon, 1969: 447. 

Notes.— This species is a primary endo- 
parasite (Van Whervin, 1968) of citrus wee- 
vil eggs {Exopthalmus or Pachneus sp. ac- 
cording to label data). It is known from 
Jamaica, although it is possible that it oc- 
curs on other islands in the West Indies. 
Although originally described in the genus 
Platystasius, this species is now considered 
better placed in Fidiobia (L. Masner, pers. 
comm.). A formal change in generic place- 
ment is being proposed in a manuscript on 
platygastrid taxonomy that is currently in 
preparation but has not yet been published. 
Therefore, I have used the currently pub- 
lished combination. 

Diagnosis.— This species is the only proc- 
totrupoid that has been associated with cit- 
rus weevil eggs. It can be identified by the 
following: tarsi 5 -segmented (3 or 4 seg- 
mented in other species); forewing with only 
a submarginal vein (Fig. 1) (obvious sub- 
marginal and marginal and stigmal veins in 
other species, except Cleruchus sp. (see Figs. 
2-4); antenna with four small funicles and 
large 3-segmented club (Fig. 6) (antennae of 
other species as in Figs. 7-11, 14); scutellum 
without obvious paired setae (Fig. 1 5) (scu- 
tellum with one or two pairs of large setae 
in others, Figs. 16, 17,20,22). 

Hosts.— Pachneus or Exopthalmus sp. 

Specimens examined. — Jamaica, St. 



Catherine Parish, Worth Park and Charlton 
nr. Ewarton; Manchester. Mona Island. 
Collected in June, August, and September. 

Clemchus sp. 

Notes.— This species was recently reared 
from eggs of Artipus floridanus Horn in the 
vicinity of Wabasso, Florida (Indian River 
Co.). Unfortunately, only 4 specimens have 
been collected, and these are in poor con- 
dition. I would place them in the genus Cle- 
mchus (sensu Schauff, 1984). They are very 
similar to C brevipennis Ogloblin (1940). 
This species is almost certainly unde- 
scribed, but without additional specimens I 
am reluctant to name it at this time. 

Diagnosis. — This species is the only 
member of the family Mymaridae yet re- 
corded from citrus weevil eggs. It can be 
identified by the following characters: tarsi 
4-segmented; forewings nearly parallel-sid- 
ed and with marginal cilia much longer than 
wing width (Fig. 5) (other species with fore- 
wings much broader and not parallel-sided 
and marginal cilia much shorter than wing 
width); female antenna with 6 funicular ar- 
ticles and a single segmented club (Fig. 14); 
male antennae with 13 segments (other 
species with male antennae with fewer than 
13 segments). 


I give special thanks to E. E. Grissell whose 
preliminary work on the taxonomy of these 
parasites made possible the timely comple- 
tion of this study. I thank also R. Woodruff, 
who together with Dr. Grissell collected 
much of the material on which this study 
has been based. R. E. White, E. E. Grissell, 
D. Wahl, and F. D. Bennett made many 
valuable comments on the manuscript. I am 
grateful to J. S. Noyes and Z. Boucek for 
the loan of specimens from the British Mu- 
seum and L. Stange for material from the 
Florida State collection of Arthropods. I am 
grateful to L. Masner (Biosystematic Re- 

search Institute, Agriculture Canada, Ot- 
tawa, Ontario), for his help with determin- 
ing the platygastrid and D. Whitehead 
(Systematic Entomology Lab, USDA, 
Washington, D.C.) for help with the weevil 
names. C. Tarrent (University of Florida) 
provided valuable specimens and infor- 
mation on Brachyufens osborni and the 
species of Cleruchus. 

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Baranowski, R. M. 1960. Notes on a parasite of the 
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Beavers, J. B. and A. G. Selhime. 1975. Further at- 
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89(1), 1987, pp. 43^6 




Robert G. Bellinger and Robert L. Pienkowski 

Department of Entomology, Virginia Polytechnic Institute and State University, Blacks- 
burg, Virginia 2406 1 . 

Abstract.— Tht brachytperous melanopline grasshopper, Appalachia hebardi Rehn and 
Rehn, is a late season, univoltine species that occurs in the Appalachian ridges of West 
Virginia, Virginia, and Pennsylvania, U.S.A. Data on this species were taken from a 
population near Mathias, West Virginia. A. hebardi feeds on understory vegetation and 
prefers older foliage. Both sexes went through five instars to become adults. Body size 
was within previously published ranges. Mean population ovariole number was 1 1 .44 ± 
1.13 (Bellinger and Pienkowski, 1985), and mean pod size was 9.67 ± 2.08. Maximum 
pod size is estimated to be 14. Some females can lay at least two pods during the season. 
Our observations corroborate previous reports and extend the life history information on 
A. hebardi. 

Appalachia hebardi Rehn and Rehn is a 
brachypterous grasshopper from the Ap- 
palachian Mountains of West Virginia, Vir- 
ginia, and Pennsylvania (Rehn and Rehn, 
1936, 1939). The genus contains one other 
species, A. arcana Hubbell and Cantrall, 
known from Michigan (Hubbell and Can- 
trail, 1938). Little is known of their biology. 
This paper provides new information, in- 
cluding food plants in the field and labo- 
ratory, development, fecundity, and sea- 
sonal occurrence of A. hebardi. We 
previously published femur lengths and 
ovariole numbers for this species (Bellinger 
and Pienkowski, 1985), but include these 
data here for completeness. 

Materials and Methods 

Grasshoppers were collected by the senior 
author (RGB) from the undergrowth around 
a cabin on the eastern slope of Big Ridge, 
east of the boundary of Lost River State 
Park, Mathias, Hardy Co., West Virginia 

(Bellinger and Pienkowski, 1985). The ele- 
vation of the site is ca. 700 meters. The 
species had been seen feeding in small num- 
bers on native and cultivated plants during 
1981 and 1982, but were less abundant in 
1983 (F. E. Wood, Dept. of Entomology, 
University of Maryland, personal commu- 
nication). The collection site was charac- 
terized by an overstory dominated by chest- 
nut oak, with an understory of black gum 
{Nyssa syhatica Marsh.), dogwood, and 
sucker growth of American chestnut {Cas- 
tanea dentata (Marsh.) Borkh.). Under- 
growth was predominately mountain laurel 
{Kalmia latifolia L.), and Vaccinium spp. 
Grasshoppers were all on the ground when 
collected. Collecting was difficult because of 
the scarcity of individuals and because when 
disturbed they jumped long distances, even 
in the dense undergrowth. One immature 
and one adult male, and one immature and 
two adult females were collected on 1 2 Sep- 
tember, 1983. One adult male and six adult 


females were collected in mid-October, 
1983. Individuals from each collection were 
returned to the laboratory, at Blacksburg, 
VA. The initial intent of the collections was 
to identify the species and determine ovar- 
iole number in the females. The species was 
tentatively identified by RGB as A. hebardi. 

Grasshoppers were maintained at 30°C in 
a rearing chamber, with 2-3 individuals per 
0.45 L paper carton with screened top and 
bottom. Initially, grasshoppers were pro- 
vided with fresh romaine lettuce {Lactuca 
sativa L. cv. 'Romana'), ground dry dog food 
(25% crude protein), and misted with water 
twice daily. Because the grasshoppers did 
not noticeably feed on the lettuce, other 
plants were tried as food, including fescue 
grass {Poa pretensis L.), leaves of dandelion, 
{Taraxacum officinale Weber), Crimson 
clover (Trifoliiim incarnatum L.), three 
species of dogwood {Cornus florida L. and 
C spp.), and red, white, and chestnut oak 
(Quercus rubra L., Q. alba L., and Q. prinus 
L.). Chestnut oak leaves were brought from 
the collection site. All other plants were col- 
lected near the laboratory. 

Grasshoppers from each collection were 
held for three to four days, or until all in- 
dividuals had become adults. All individ- 
uals were sacrificed at that time. Females 
were dissected to determine ovariole num- 
ber (Bellinger and Pienkowski, 1985) and 
their ovipositional status (Launois-Luong, 
1978). Measurements of total body length, 
hind femur, and tegmen were made on three 
males and nine females, and of the length 
and width of the pronotum of the females. 
Because development of the grasshoppers 
was not observed in the field or in the lab- 
oratory, the number of antennal segments 
in the adults were counted to determine 
number of instars required to reach the adult 
(Shotwell, 1941). 


The identification of this species as Ap- 
palachia hebardi Rehn and Rehn was con- 
firmed by Irving J. Cantrall, Museum of Zo- 

ology, University of Michigan, Ann Arbor 

Habitat and food plants. — Feeding pre- 
viously had been noted on native mountain 
laurel, sucker growth on cut chestnut oak 
stumps, several herbaceous weeds, and on 
cultivated rose of sharon {Hibiscus sp.). In 
the laboratory the grasshoppers ate ground 
dry dog food but not romaine lettuce. Grass- 
hoppers ate old leaves from lower parts of 
the three dogwood species but not newer 
leaves from branch ends. Similarly, only 
older leaves of dandelion were eaten. Grass- 
hoppers did not feed on foliage from the 
three species of oak, or the grass, or the 

Body size. — Body measurements (mean ± 
SD (mm)) were: total body length, males— 
20.87 ± 1.16, females-25.27 ± 2.06, hind 
femur, males— 10.9 ± 0.35, females — 
12.28 ± 0.52, tegmen, males-4.33 ± 0.57, 
females— 4.59 ± 0.38, pronotal length, fe- 
males— 5.09 ± 0.43, pronotal width — 
4.10 ± 0.38. 

Antennal segments and instar number.— 
Two of the males had 23 antennal segments, 
and one had 24 segments. Three of the fe- 
males had 23 segments, and the remaining 
six had 24. There was no apparent difference 
in segment number because of collection 
dates. Antennal segment counts showed that 
all individuals in each sex had gone through 
five developmental stadia to reach the adult. 
Antennae were long for the number of seg- 
ments. Individual segments were visibly 
longer than in the genus Melanoplus Stal. 

¥QC\xnd\\y. —Appalachia hebardi mated 
readily in the laboratory. No eggs were laid 
in the laboratory, probably because a suit- 
able ovipositional media was not provided. 
Mean ovariole number per female was 
11.44 ± 1.13 SD, and ranged from 10 to 13 
(Bellinger and Pienkowski, 1985). There 
were five to seven ovarioles per ovary, so 
there could be up to 1 4 ovarioles per female. 
Dissections of females, collected as adults 
in the field, showed that the two adults col- 
lected in September each had 1 2 ovarioles. 



One showed little development of the ova- 
ries, and the other had laid one pod with 1 2 
eggs but had not yet begun development of 
a second pod. Adult females collected in 
October showed a range of ovarian devel- 
opment. One female had oviposited twice 
and was developing a third pod. This female 
had 1 1 ovarioles and had laid eight eggs in 
the first pod and nine eggs in the second 
pod. Seven oocytes were being developed, 
indicating a maximum of seven eggs in the 
potential third pod. None of the others had 
oviposited, nor had they begun oocyte de- 
velopment, but two had 12 ovarioles and 
12 well developed oocytes each. Average 
pod size was thus 9.67 ± 2.08 (n = 3 pods). 
While color of the chorion in grasshopper 
eggs varies among species, developing oo- 
cytes of this species were bright yellow, the 
same color observed in other Melanoplinae, 
and in most Oedipodinae and Gomphocer- 
inae (unpublished). We did not observe any 
post-ovipositional eggs. No parasites were 
noted in any dissection. 


Observations made on a small population 
of A. hebardi from West Virginia confirm 
some observations previously made on the 
species. The collection site was a habitat 
similar to those described by Rehn and Rehn 
(1936). The elevation in West Virginia was 
within the 550 to 1 220 m range of elevations 
given for the species (Rehn and Rehn, 1 939). 
We found as did Rehn and Rehn ( 1936) that 
the species jumped long distances to escape 
capture. Plant associations but no food 
plants were given by Rehn and Rehn (1936). 
Our field observations and laboratory feed- 
ings, although somewhat limited, suggested 
that the species feeds on older foliage of 
undergrowth species. 

Individuals from the Mathias, WV pop- 
ulation in 1983 went through five stadia. 
This is a common number of stadia for ac- 
ridid grasshoppers. Brachypterous acridid 
grasshoppers frequently go through a re- 
duced number of stadia (four or five), and 

as a result have small body sizes (Mason, 
1954; Uvarov, 1966, 1977). Species from 
woodlands and higher altitudes are fre- 
quently brachytperous. Body size measure- 
ments for the Mathias, WV population fell 
within the range reported by Rehn and Rehn 
(1936), except for the length of the tegmen 
in males, which was longer. 

Appalachia herbardi has small ovaries 
(low number of ovarioles), a condition which 
is related to its small body size (Bellinger 
and Pienkowski, 1985). Our counts of ovar- 
ioles indicate potential pod sizes of 10 to 
1 4 eggs per pod. Our data showed that pod 
sizes in the field averaged 9.67 eggs. Rehn 
(1938) reported "approximately ten eggs" 
from a single pod. Females in the Mathias 
population are capable of laying at least two, 
and possibly three pods per female, based 
on one female. Thus, this species may lay 
up to ca. 42 eggs per female, on the basis of 
these limited data. The oviposition sub- 
strate for this species remains unknown. 
While most acridid grasshoppers oviposit 
in the soil, some woodland species oviposit 
in holes in dead wood (Blatchley, 1920). 

Rehn and Rehn (1936) reported that A. 
hebardi occurred as early as the first week 
of July, to as late as early September, over 
its range. Our collections were made in mid- 
September, when we collected immatures 
and adults, and mid-October. The repro- 
ductive status of adult females showed that 
the population began oviposition before 
mid-September, but that as late as mid-Oc- 
tober some females had yet to oviposit, or 
even begin to develop their first pod. At the 
elevation of the Mathias site, the growing 
season is over by mid-October, and unless 
this species is adapted to cooler tempera- 
tures and senescent food plants, some fe- 
males in the population may never oviposit. 


We thank F. E. Wood for informing us 
of the study population and allowing us ac- 
cess to it. We thank Irving J. Cantrall for 
confirming the identification of Appalachia 



hebardi and encouraging our studies of 
grasshopper biology. We thank also David 
A. Nickle, Systematic Entomology Labo- 
ratory, Agricultural Research Service, c/o 
U.S. National Museum, Washington, D.C. 
for providing us with literature on Appa- 

Literature Cited 

Bellinger, R. G. and R. L. Pienkowski. 1985. Inter- 
specific variation in ovariole number in melanop- 
line grasshoppers (Orthoptera: Acrididae). Ann. 
Entomol. Soc. Am. 78: 127-130. 

Blatchley, W. S. 1920. Orthoptera of northeastern 
America. The Nature Publishing Co. Indianapolis. 
784 pp. 

Hubbell, T. H. and I. J. Cantrall. 1938. A new species 
of Appalachia from Michigan (Orthoptera, Acri- 
didae, Cyrtacanthacridinae). Univ. Michigan Mus. 
Zool. Occ. Paper No. 389: 1-21. 

Launois-Luong, M. H. 1978. Methode pratique d'in- 
terpretation de Tetat des ovaires des acridiens du 
Sahel. Ann. Zool. Ecol. Anim. 10: 569-587. 

Mason, J. B. 1954. Number of antennal segments in 
adult Acrididae (Orthoptera). Proc. R. Entomol. 
Soc. Lond. B 23: 228-238. 

Rehn, J. A. G. and J. W. H. Rehn. 1936. On new or 
redefined genera of nearctic Melanopli (Orthop- 
tera: Acrididae, Cyrtacanthacridinae). Trans. Am. 
Entomol. Soc. 62: 1-56. 

. 1939. Studies of certain cyrtacanthacridoid 

genera (Orthoptera: Acrididae). Part I. The Podis- 
ma complex. Trans. Am. Entomol. Soc. 65: 61- 

Rehn, J. W. H. 1938. Notes on the eggpods of Ap- 
palachia hebardi and Dendrotettix quercus (Or- 
thopter: Acridiae; Cyrtacanthacridinae). Entomol. 
News 49: 259-260. 

Shotwell, R. L. 1941. Life histories and habits of 
some grasshoppers of economic importance on the 
Great Plains. U.S. Dept. Agric. Tech. Bull. No. 

Uvarov, B. 1966. Grasshoppers and locusts: a hand- 
book of general acridology, vol. 1 . Cambridge Uni- 
versity Press, Cambridge. 481 pp. 

. 1977. Grasshoppers and locusts: a handbook 

of general acridology, vol. 2. Centre for Overseas 
Pest Research, London. 613 pp. 


89(1), 1987, pp. 47-50 





Michael J. Sharkey 

Biosystematics Research Centre, Agriculture Canada, Ottawa, Ontario Kl A 0C6, Canada. 

Abstract.— \d\x\Xs of Agathis thompsoni n. sp. (Braconidae: Agathidinae) from north- 
western U.S.A. are described and illustrated. Larvae are parasitic on Greya subalba (Braun), 
which feeds on the schizocarps of Lomatium spp. (Umbelliferae). Females of the new 
species are differentiated from those of both European and North American species. 

Among the many undescribed species of 
Agathis Latreille in North America is one 
that has been studied by J. N. Thompson 
(Thompson, 1986). It is to compliment his 
research on oviposition behaviour and 
searching efficiency that I describe this new 

Agathis thompsoni Sharkey, New Species 

Diagnosis. —Agathis thompsoni is distin- 
guished from other species oi Agathis by the 
following combination of character states: 
basal flagellomere 1.6 x longer than follow- 
ing flagellomere; malar space 0.6 x greatest 
diameter of eye; ovipositor 1.2 x as long as 
metasoma when fully extended. 

Description, holotype $.— (Intraspecific 
variation is given in parentheses). Color: 
Black except metasoma slightly paler lat- 
erally and yellowish-orange as follows: 
mandible, all femora over distal 0.7, fore 
and middle tibiae, hind tibia except for bas- 
al and apical melanic bands, basal 0.2 of all 
basitarsomeres; fore wing hyaline. Head 
(Fig. 1): Antenna with 26 (23-26) flagello- 
meres; basal flagellomere 1.6 x longer than 
following flagellomere; head subrostiform, 
malar space 0.6 x greatest diameter of eye; 
weak, V-shaped depression anterior to me- 
dian ocellus; smooth, longitudinal ridge 

from near median ocellus to level of anten- 
nal insertion; galea 2.2 x longer than max- 
imum width. Mesosoma (Figs. 2, 3): Notauli 
deeply impressed, pitted, scutellar groove 
with numerous longitudinal ridges; prono- 
tum smooth except for crenulae along pos- 
terior margin; stemaulus 0.7 x length of 
mesopleuron and complete to posterior 
margin; metapleuron rugose over ventral 0.2 
(0.2-0.3); propodeum with transverse, an- 
terolateral ridge and with 3 longitudinal 
ridges, medial ridge weak; (propodeum may 
be somewhat rougher than in Fig. 2); mid 
tibia with 3 (2-3) preapical spines; hind tib- 
ia with 6 (4-6) apical spines; hind tarsal claw 
with strong basal tooth. Metasoma (Fig. 4): 
First tergum as long as wide, with pair of 
weak longitudinal ridges and weak striae 
over anterior 0.5; tergum 2 + 3 mostly 
smooth with basal swelling; ovipositor 1.2 x 
longer than metasoma when fully extended; 
ovipositor sheaths slightly shorter (0.9 x) 
than metasoma. Length: 3.7 (3.5-4.3) mm. 

Allotype (5.— As for the holotype except 
antenna with 23 flagellomeres. (Left hind 
leg missing after coxa). 

This species is named after John N. 

Material examined. — Holotype 9, U.S.A., 
Washington, Whitman Co., Smoot Hill Biol. 



Figs. 1, 2. Agathis thompsoni. 1, Head, lateral. 2, Metanotum and propodeum, dorsal 

Pres. nr. Albion, ex. Greya subalba (Braun), 
5.VI.1985, J. N. Thompson, (United States 
National Museum). Allotype $, same data 
as holotype except date, 17. VI. 80. Para- 
types: 38 9, same data as holotype, (Cana- 
dian National Collection, United States Na- 
tional Museum). I 6, same data as allotype, 
(Canadian National Collection). 

Discussion. — In a recent paper (Sharkey, 
1985) I defined my concept oi Agathis La- 
treille. The following Nearctic and Holarc- 
tic species belong to this genus: A. brevicor- 
nis (Muesebeck), A. cupressi Muesebeck and 
Walkley, A. gibbosa (Say), A. malvacearum 
Latr., A. pumila (Ratzburg), A. rubripes 
Cresson, A. terminata Cresson, A. thomp- 
soni Sharkey, A. tibiator Provancher. 

Females of ^. thompsoni differ from those 
of most other North American species of 
Agathis by their short ovipositor, which is 
only 1.2 X as long as the metasoma when 

fully extended. Other species have the ovi- 
positor fully as long as the body except A. 
pumila, a Holarctic species. Unlike A. 
thompsoni, A. pumila has the two most bas- 
al antennal flagellomeres subequal in length. 
In Nixon's (1986) key to the European 
females of Agathis, A. thompsoni keys to 
couplet number 30. Females differ from 
those of five of the six species that key 
through this couplet by their short ovipos- 
itor. Females of A. melpomene Nixon, which 
also have a short ovipositor, differ in that 
the first tergum of the metasoma is mostly 
smooth, though sometimes with weak ru- 
gosity medially. Females of ^. thompsoni 
have striae in the anterior 0.6 of the first 
metasomal tergum. 


I thank G. Gibson and J. Huber for their 
reviews, B. Jinkinson for the scanning elec- 



Figs. 3, 4. Agathis thompsoni. 3, Mesosoma, lateral. 4, Metasoma, dorsal. 

tron photomicrographs and the Electron America north of Mexico. Proc. U.S. Natl. Mus. 

Microscope Centre, Agricuhure Canada, for 69: 1-73. 

the use of its facilities. Nixon, G. E. J. 1986. A revision of the European 

Agathidinae (Hymenoptera: Braconidae). Bull. Br. 

Literature Cited Mus. (Nat. Hist.) Entomoi. 52: 183-242. 

Muesebeck, C. F. W. 1927. A revision of the parasitic Sharkey, M. J. 1985. Notes on the genera Bassus 

wasps of the subfamily Braconinae occurring in Fabricius and Agathis Latreille, with a description 


of Bassus arthurellus n. sp. Can. Entomol. 117: searching efficiency in a natural population of a 

1 497-1502. braconid parasitoid. J. Anim. Ecol. 55: 351-360. 

Thompson, J. N. 1986. Oviposition behaviour and 


With this issue the Proceedings begins a two-column format not including titles and 
abstracts. This is a cost-saving measure that the Publications and Executive Committees 
believe does not sacrifice utility or beauty. The new format should allow for some growth 
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89(1), 1987, pp. 51-56 




C. Riley Nelson and Richard W. Baumann 

Department of Zoology, Brigham Young University, Provo, Utah 84602. 

Abstract. —Scdinmng electron micrographs are produced for nine species of Cap ni a: C. 
barbed, C. cheama, C. coloradensis, C. decepta, C. elevata, C. fibula, C. manitoba, C. 
melia, and C. uintahi. Structural details not previously reported such as spines and sensilla 
may prove useful in constructing future phylogenies. The value of scanning electron 
microscopy for species identification and systematic work is discussed. 

Scanning electron microscopy (SEM) has 
been used to examine structural details of 
Plecoptera eggs (Baumann, 1973; Stark and 
Baumann, 1978; Szczytko and Stewart, 
1979; Stark and Stewart, 1981; Stark and 
Szczytko, 1981a, b; Stark and Stewart, 
1982c; Stark and Ray, 1983; Stark, 1983; 
Stark and Szczytko, 1982; Stark and Szczyt- 
ko, 1984), mouthparts (Baumann, 1973; 
Stark and Stewart, 1981; Stark and Stewart, 
1982a-c; Stark and Ray, 1983), and sensilla 
(Stark and Stewart, 1981; Stark and Stewart, 
1982a; Stark and Ray, 1983; Stark, 1983; 
and Kapoor and Zachariah, 1984). These 
studies have helped in constructing phylog- 
enies such as those of Nelson (1984). The 
use of SEM to illustrate male genitalia of 
stoneflies has been limited to the species 
description of Prostoia hallasi (Kondratieff 
and Kirchner, 1984). A study of the male 
genitalia of Capnia was undertaken to ex- 
amine the feasibility of using SEM as a tool 
to differentiate species and species groups 
in this large genus. 

Materials and Methods 

Specimens of North American Capnia 
were examined and compared to holotypes 
(except that of C. fibula Claassen, which was 

missing). The term genitalia refers to the 
epiproct of the male and associated sclerites 
while terminalia refers to all the appendages 
of the abdominal segments including tergal 
knobs, genitalia, and cerci. Collection data 
for the specimens used in this study are: 

Capnia barberi Claassen — CALIFOR- 
NIA, Nevada County, Sagehen Creek, 26 
February 1965, Sheldon and Hawthorne. 

Capnia cheama Ricker— MONTANA, 
Lincoln County, Kootenai River, 1 9 March 
1970, R. L. Newell. 

Capnia coloradensis Claassen— COLO- 
RADO, Routt County, Fish Creek, Hwy 40, 
Steamboat Springs, 6 April 1963, A. R. 

Capnia decepta Banks — ARIZONA, 
Cochise County, Huachuca Mountains, 
Ramsey Canyon, 17 January 1984, Bau- 
mann and Nelson. 

Capnia elevata Frison— OREGON, Was- 
co County, creek, Hwy 30, 4 mi E Rowena, 
3 March 1984, Baumann and Nelson. 

Capnia fibula Claassen— ARIZONA, Co- 
conino County, West Fork Oak Creek, 6 
February 1984, M. W. Sanderson. 

Capnia manitoba Claassen— MAINE, sta. 
2 Rt. 5 Saco, 3 April 1964, C. H. Nelson. 

Capnia melia Frison— OREGON, Clack- 



amas County, Salmon River at headwaters, 
jet. Hwys 26 & 35, 2 March 1984, Bau- 
mann. Nelson, Fiala. 

Capnia uintahi Gaufin — UTAH, Davis 
County, Farmington Canyon at Fork Bridge, 
5 April 1966, R. W. Baumann. 

In preparation for study with the scanning 
electron microscope, specimens were seri- 
ally dehydrated from their storage fluid of 
70% alcohol for more than 10 minutes in 
95% ethanol, 20 minutes in absolute (100%) 
ethanol over calcium sulfate, and stored for 
more than 20 minutes in acetone previously 
kept over calcium sulfate. The specimens 
were then transferred to acetone-immersed 
holding baskets in a Sorvall critical point 
drying apparatus attached to a carbon diox- 
ide tank. The dried abdomens of the spec- 
imens were then placed on aluminum stubs 
covered with double-sided tape. The spec- 
imen-bearing stubs were gold coated using 
a Polaron gold-coating apparatus. Coated 
specimens were then examined using an 
AMRay 1000 scanning electron microscope 
at 20 kV. 


This study reveals that male terminalia 
of Capnia are well suited for examination 
using SEM. The epiproct, which is used in 
species determinations, is heavily sclero- 
tized and thus resistant to deformation dur- 
ing drying and other preparation procedures 
necessary for viewing with a scanning elec- 
tron microscope. This structural stability in 
the Capnia is uncommon in stoneflies since 
important taxonomic characters in genera 
in other families are usually on softer body 
parts that are susceptible to deformation 
during specimen preparation. 

Detailed structures not previously noted 
in species descriptions were seen with the 
SEM. A few of these features that indicate 
phylogenetic relationships are outlined be- 

In C. cheama (Fig. 1) and C. fibula (Figs. 
4 and 1 2) details of the dorsal abdominal 

hump are shown. Both trichoid and cam- 
paniform sensilla are present. The paucity 
of trichoid sensilla on the knob of tergum 
seven in C. fibula compared with the nu- 
merous sensilla of the knob of tergum eight 
in C. cheama distinguishes two groups of 
species in the genus. In C. excavata and C. 
uintahi the setation of the tergal knob re- 
sembles that of C cheama and the setation 
of C fibula is similar to that of C. venosa, 
C. elevata, C wanica, and C. manitoba. 

In C. fibula (Fig. 3) spines are asymmet- 
rical along the lateral margins of the epi- 
proct when seen in dorsal view. These spines 
are directed cephalad and would seemingly 
hinder insertion of the epiproct into the fe- 
male. Similar spines occur in several species 
groups of Capnia and take on a diversity of 
placements including the stout, tightly 
bunched configuration of C. decepta (Figs. 
11,13 and 1 5) and a scattered pattern along 
the anterior margin of the epiproctal bulb 
of C. melia (Figs. 14 and 16). Other species 
such as C. barberi (Fig. 9) show no indica- 
tion of these spines. 

The two-limbed epiproct of C manitoba 
(Fig. 8) probably represents a plesiomorphic 
condition for at least one group of Capnia. 
The presence of a lower epiproctal limb was 
first noted using SEM in C elevata (Fig. 10) 
and later noted using a light microscope in 
C. fibula, C venosa, and C. wanica. These 
species show a reduced lower process and a 
cline featuring a gradual reduction in the 
length of this character as one moves away 
from the range of C. manitoba. No other 
named species of Capnia in North America 
has a similar two-limbed epiproct or hint 
of a reduced lower limb. 

A relatively simple tube epiproct such as 
that of C coloradensis (Fig. 7) is shared by 
several species in the genus, but only one 
other, C. petila, has the apex of the epiproct 
drooping ventrally. The tube epiproct of 
some Capnia represents the most apo- 
morphic condition of the epiproct and is 
shared by several species, including C. con- 



Figs. 1-6. Male terminalia of Capnia spp. 1, C. cheama Ricker. epiproct and tergal knob, 225 x (dorsal). 2, 
C. cheama Ricker, tip of epiproct, 650 x (anterolateral). 3, C. fibula Claassen, epiproct, 180x (dorsal). 4, C. 
fibula Claassen, epiproct and tergal knob, 75 x (lateral). 5, C. uintahi Gaufin, epiproct and tergal knob, 105 x 
(dorsal). 6, C. unitahi Gaufin, epiproct and tergal knob, 1 lOx (lateral). 

fusa, C. elongata, C. gracilaria, C. lacustra, 
C. lineata, C. promota, C. vernalis, and C 
Two closely related species, C cheama 

and C. uintahi demonstrate the value of 
SEM in illustrating characters useful for de- 
terminations at the specific level. The dis- 
tribution of the two kinds of sensilla on the 



Figs. 7-12. Male terminalia of Capnia spp. 7, C. coloradensis Claassen, epiproct and tergal knob, 105 x 
(lateral). 8, C manitoba Claassen, epiproct and tergal knob 120 x (lateral). 9, C. barbeh Claassen, epiproct, 90 x 
(lateral). 10, C. elevata Frison, epiproct, 105 x (lateral). 11, C decepta Frison, epiproct, 100 x (lateral). 12, C. 
fibula Claassen, tergal knob, 700 x (dorsal). 

tergal knobs (Figs. 1 and 5) are useful in 
separating these two species. Higher mag- 
nification of the tip of the epiproct (Figs. 2 
and 6) shows a dorsal appendage near the 
apex in C. cheama which is not present in 

C uintahi. These structures are not readily 
apparent using a dissecting microscope and 
phylogenetically important information 
given by these characters has been unavail- 
able using classical techniques. 



Figs. 13-16. Male terminalia of Capnia spp. 13, C. decepta Prison, epiproct and tergal knob, 80 x (dorsal). 
14, C. melia Prison, epiproct, 150x (dorsal). 15, C decepta Prison, epiproct, 85 x (dorsal). 16, C melia Prison, 
epiproct, 800 x (dorsal). 




The higher magnification and greater 
depth of field available make SEM a valu- 
able tool for investigating evolutionary and 
taxonomic relationships within Capnia and 
the potential for greater use is apparent. 
Comparison of minute details between taxa 
can aid systematists in understanding species 
variation and constructing accurate phylog- 
enies. The expense of SEM work and the 
often destructive manipulation of speci- 
mens, though, Hmit usefulness of SEM for 
routine species determination. These con- 
straints could result in overconfidence based 
on the characters observed from a single 
specimen rather than from a series of spec- 
imens. However, once a new character is 
observed using SEM more specimens may 
be examined using a dissecting microscope 
to confirm the character from a series of 

Although relatively low magnification was 
used during this preliminary study, other 
structures of probable taxonomic worth, 
such as epiproctal spines, sensilla, and se- 
tation patterns were observed. Once the mi- 
nute characters are examined from series of 
specimens and relationships established, 
routine species determinations can be per- 
formed using characters visible with an or- 
dinary dissecting microscope. 


We thank Wilford M. Hess, James V. Al- 
len, and Connie Swensen of the Electron 
Optics Laboratory at Brigham Young Uni- 
versity for their help with technical and crit- 
ical aspects of the scanning electron mi- 

Literature Cited 

Baumann, R. W. 1973. Studies on Utah stoneflies 
(Plecoptera). Great Basin Nat. 33: 91-108. 

Kapoor, N. N. and K. Zachariah. 1 984. Scanning and 
transmission electron microscopy of the devel- 
opmental stages of the flower-shape sensillum of 
the stonefly nymph Thaumatoperla alpina Bums 
and Neboiss (Plecoptera: Eustheniidae). J. Insect 
Morphol. Embryol. 13: 177-189. 

Kondratieff, B. C. and R. F. Kirchner. 1984. Anew 
species of Nemouridae (Plecoptera) from the Great 
Dismal Swamp, Virginia, USA. Proc. Entomol. 
Soc. Wash. 86: 578-581. 

Nelson, C. H. 1984. Numerical cladistic analysis of 
phylogenetic relationships in Plecoptera. Ann. 
Entomol. Soc. Am. 77: 466-473. 

Stark, B. P. 1983. A review of the genus Soliperla 
(Plecoptera: Peltoperlidae). Great Basin Nat. 43: 

Stark, B. P. and R. W. Baumann. 1978. New species 
of nearctic Neoperla (Plecoptera: Perlidae), with 
notes on the genus. Great Basin Nat. 38: 97-1 14. 

Stark, B. P. and D. H. Ray. 1983. A revision of the 
genus Helopicus (Plecoptera: Perlodidae). Fresh- 
wat. Invertebr. Biol. 2: 16-27. 

Stark, B. P. and K. W. Stewart. 1981. The nearctic 
genera of Peltoperlidae (Plecoptera). J. Kans. 
Entomol. Soc. 54: 285-311. 

. 1982a. Oconoperla, a new genus of North 

American Perlodinae (Plecoptera: Perlodidae). 
Proc. Entomol. Soc. Wash. 84: 746-752. 

. 1982b. T\it nymx)\\ o{ Viehoperla ada {?\t- 

coptera: Peltoperlidae). J. Kans. Entomol. Soc. 55: 

. 1982c. Notes ofPer/mo^e^aMrra (Plecoptera: 

Perlodidae). Ann. Entomol. Soc. Am. 75: 84-88. 

Stark, B. P. and S. W. Szczytko. 1981a. Contributions 
to the systematics of Paragnetina (Plecoptera: Per- 
lidae). J. Kans. Entomol. Soc. 54: 625-648. 

. 1981b. Skwala brevis (Kopenen) from Japan 

(Plecoptera: Perlodidae). Aquat. Insects 3: 61-63. 

. 1982. Egg morphology and phylogeny in 

Pteronarcyidae (Plecoptera). Ann. Entomol. Soc. 
Am. 75: 519-529. 

. 1984. Egg morphology and classification of 

Perlodinae (Plecoptera: Perlodidae). Ann. Limnol. 
20: 99-104. 

Szczytko, S. W. and K. W. Stewart. 1979. The genus 
Isoperla (Plecoptera) of western North America; 
Holomorphology and systematics, and a new 
stonefly genus Cascadoperla. Mem. Am. Entomol. 
Soc. 32: 1-120. 


89(1), 1987, pp. 57-60 


J. E. LowRY AND H. Derrick Blocker 

(JEL) Insect Pathology Laboratory, Agricultural Research Service, USDA, Beltsville, 
Maryland 20705; (HDB) Department of Entomology, Kansas State University, Manhat- 
tan, Kansas 66506. 

Abstract.— Two new species of Flexamia from perennial chloridoid grasses in the Ne- 
braska Sand Hills are described and illustrated. These are F. celata and F. arenicola. The 
previously unknown female of F. dakota is also described. 

The genus Flexamia is one of the most 
important and conspicuous groups of grass- 
land cicadellids. In addition to species rec- 
ognized by Young and Beime ( 1 9 5 8) in their 
revision, Hamilton and Ross (1975) de- 
scribed F. satilla, and Ross and Cooley 
(1969) added F. delongi. Although the latter 
form may be a geographic variation of San- 
ders i, its description raises to 36 the number 
of described species. In this report we de- 
scribed two new species and the female of 

The new species, F. arenicola and F. ce- 
lata, were discovered in sand blowouts in 
the Nebraska Sand Hills; they are closely 
related to F. flexulosa and F. stylata re- 
spectively, but differ in male genitalic struc- 
tures. F. dakota, whose female had been 
previously unknown, was collected from 
Texas to North Dakota and on west-facing 
slopes in the Loess hills of western Iowa. 
All collections of dakota were from Schi- 
zachyrium scoparium (Michx.) Nash. The 
cicadellid species we describe are named 
from material collected by R. F. Whitcomb. 

' Contribution No. 85-535-J, Department of Ento- 
mology, Kansas Agricultural Experiment Station, Kan- 
sas State University, Manhattan, Kansas 66506. 

Type material is deposited in the U.S. Na- 
tional Museum of Natural History (USNM) 
and Kansas State University (KSU). Para- 
types are deposited also in the Grassland 
Cicadellid Collection of the Insect Pathol- 
ogy Laboratory, ARS, USDA, Beltsville Ag- 
ricultural Research Center-East (BARC-E), 
Beltsville, Maryland. 

Flexamia celata Lowry and Blocker, 

New Species 

Fig. 1 

Description. — Length of <5 4.4 mm, 9 4.4- 
4.7 mm; head width of 3 1.3 mm, 9 1 .4 mm; 
pronotal width of 3 1.2 mm, 9 1.3 mm; in- 
terocular width of 6 0.6 mm, 9 0.7 mm; 
vertex length of 5 0.7 mm, 9 0.8 mm; pro- 
notal length of (5 0.5 mm, 9 0.6 mm. Vertex 
not produced; median length of vertex ap- 
proximately Vi head width and slightly long- 
er than interocular width. 

Color pale stramineous with irregular dark 
markings on dorsum and forewings; vertex 
with black interocular line; head and prono- 
tum with faint tan markings as stylata; ven- 
ter and legs with irregular fuscous markings; 
9 sternum VII with pair of conspicuous fus- 
cous stripes. 

Male pygofer with central margin strongly 



Fig. 1 . Flexamia celata. A, aedeagus and connective, lateral aspect. B, same, ventral aspect. C, right style, 
dorsal aspect. D, male plates. E, male pygofer, lateral aspect. F, female seventh sternum. G, bases of first valvulae 
of female. 

produced to form spinelike process as sty- 
lata; plates extend to approximately % length 
of pygofer, fused mesally for Vi length, nar- 
rowed apically to rounded lateral lobe, api- 
ces of the two plates meeting in v-shaped 
notch; connective in lateral view with dorsal 
keels broad, approximately Vi height of dor- 
sal apodeme; apodemal processes straight, 
unmodified, in dorsal aspect parallel to ae- 
deagal shaft; style digitate in apical Vi, 
preapical lobe pronounced; aedeagus sym- 
metrical, shaft slender, not conspicuously 
curved, not expanded apically, central and 
paired apical processes approximately equal 
in length and less than 'A length of shaft, 
gonopore apical on caudodorsal surface. 

Female sternum VII with posterior mar- 
gin produced medially, with slight medial 
notch; ovipositor with basal processes of first 

valvulae recurved and extending laterad be- 
yond lateral margin. 

Types. — Holotype S: Garden Co., Ne- 
braska, Crescent Lake National Wildlife 
Refuge, 27-VI-1984, R. F. Whitcomb. De- 
posited in USNM. Paratypes: 1 2, Sheridan 
Co., Nebraska, Lakeside, 28-VI-1984; 2 $, 
holotype locality, 28-VI-1984. Deposited at 
KSU and BARC-E. 

Remarks.— Flexamia celata keys to cou- 
plet 9 in Young and Beime's key to males 
and is apparently closely related to stylata. 
It can be distinguished by the shaft of the 
aedeagus which is slender, not conspicu- 
ously curved, not expanded apically, and 
which has shorter apical appendages. The 
processes at the base of the first valvulae of 
the ovipositor are diagnostic. 

Additional records: Most specimens were 



Fig. 2. Flexamia arenicola. A, aedeagus and connective, lateral aspect. B, same, ventral aspect. C, right style, 
dorsal aspect. D, male plates. E, male pygofer, lateral aspect. F, female seventh sternum. G, bases of first valvulae 
of female. 

taken from Redfieldia fJexuosa (Thurb.) Va- 
sey. NEBRASKA: Box Butte, Cherry, and 
Hooker counties June-August. 

The name celata is an adjective meaning 

Flexamia arenicola Lowry and Blocker, 

New Species 

Fig. 2 

Description.— Length of 3 3.7-3.8 mm, 2 
3.8-4.1 mm; head width of 3 1.1-1.2 mm, 
2 1.2-1.3 mm; pronotal width of 3 1.0-1.1 
mm, 9 1.1-1.2 mm; interocular width of 3 
0.5-0.6 mm, 9 0.6-0.7 mm; vertex length 
of 3 0.6-0.7 mm, 9 0.7-0.8 mm; pronotal 
length of 3 0.4-0.5 mm, 9 0.5-0.6 mm. Me- 
dian length of vertex approximately % head 
width and 1 'A times interocular width. 

Color pale stramineous with irregular dark 
brown markings on forewings and apex of 
vertex; faint stripes on vertex and prono- 

tum; basal interocular lines fuscous to black; 
venter and legs with irregular fuscous mark- 
ings; 9 sternum VII with pair of fuscous spots 
on hind margin. 

Male pygofer with posterior lobe pro- 
duced anteroventrally, caudal margin 
fuscous, with or without a small acute 
projection apically; plates extending to ap- 
proximately % length of pygofer, mesal mar- 
gins touching for Vi length then forming con- 
spicuous V-shaped notch apically; style 
digitate apically; connective in lateral aspect 
with dorsal keels Vi height of dorsal apo- 
deme; aedeagus symmetrical, apodemal 
processes divergent apically, nearly attain- 
ing tip of shaft, gonopore apical on caudo- 
ventral surface, with pair of lateral apical 
processes approximately Vs length of shaft 
and curved laterodorsally, with unpaired 
ventral process of approximate same length 
directed anteriorly. 



Fig. 3. Flexamia dakota, female. A, seventh ster- 
num. B, bases of first valvulae. 

Female sternum VII with posterior mar- 
gin produced medially, with slight median 
notch; ovipositor with recurved processes 
at base of first valvula not exceeding lateral 
margin, processes extending caudad, sin- 
uate and digitate apically. 

Types. — Holotype 6: Garden Co., Ne- 
braska, Crescent Lake National Wildlife 
Refuge, 27-VI-1984. Deposited at USNM. 
Paratypes: 40 6, 1 5 9, same data as holotype; 
1 9, Sheridan Co., Nebraska, 28-VI-1984; 
1 3, Cherry Co., Nebraska, Brownlee, 9- VIII- 
1977. Deposited at KSU, BARC-E. 

Flexamia arenicola is apparently closely 
related Xoflexulosa but can be distinguished 
by the expanded dorsal keels of the con- 
nective, the more robust aedeagal shaft and 
the longer aedeagal processes that are pres- 
ent in arenicola. It keys to couplet 17 in 
Young and Beirne's key to males. It can be 
distinguished from modica and texana by 
its larger size, and its nearly symmetrical 
aedeagus with lateral apical processes that 
are dorsally curved. The processes at the 
base of the first valvulae of the ovipositor 
are diagnostic. 

Additional records.— All specimens were 
taken from Muhlenbergia pungens Thurb. 
NEBRASKA: Lincoln, McPherson, and Box 
Butte counties; COLORADO: Morgan Co., 

The name arenicola is a noun in appo- 
sition meaning "sand inhabitant." 

Flexamia dakota Young and Beirne 

Fig. 3 

Description of 9.— Length 3.1-3.2 mm; 
head width 0.9-1 .0 mm; pronotal width 0.8- 
0.9 mm; interocular width 0.4-0.5 mm; ver- 
tex length 0.6-0.7 mm; pronotal length 0.3- 
0.4 mm. Median length of vertex approxi- 
mately % head width and IV2 times inter- 
ocular width. Sternum VII with posterior 
margin produced with slight medial con- 
cavity; ovipositor without basal processes. 

Remarks.— Young and Beirne did not 
separate the female of dakota from that of 
sandersi (see also delongi of Ross and Coo- 
ley, 1969). The females described here were 
found in association with males of dakota. 

Additional records.— All specimens tak- 
en from Schizachyrium scoparius. IOWA: 
Harrison, Monona, Pottawattomie, and 
Woodbury counties; NEBRASKA: Cherry, 
Lincoln and McPherson counties; WYO- 
MING: Campbell, Platte, and Weston 
counties; TEXAS: Roberts and Travis 
counties; OKLAHOMA: Alfalfa Co., June- 


The authors acknowledge the assistance 
of R. F. Whitcomb who collected the ma- 
terial used in this paper and initiated this 
collaboration. We also thank W. E. Steiner 
and A. L. Hicks for advice and assistance. 

Literature Cited 

Hamilton, K. G. A. and H. H. Ross. 1975. New 
species of grass-feeding deltocephaline leafhoppers 
with keys to the Nearctic species of Palus and 
Rosenus (Rhynchota: Homoptera: Cicadellidae). 
Can. Entomol. 107:601-611. 

Ross, H. H. and T. A. Cooley. 1969. A new Nearctic 
leafhopper of the genus Flexamia (Hemiptera: Ci- 
cadellidae). Entomol. News 80: 246-248. 

Young, D. A. and B. P. Beirne. 1958. A taxonomic 
revision of the leafhopper genus Flexamia and a 
new related genus (Homoptera: Cicadellidae). 
USDA Tech. Bull. 1 173, 53 pp. 


89(1), 1987, pp. 61-73 


Robert A. Wharton 

Department of Entomology, Texas A&M University, College Station, Texas 77843. 

Abstract.— LQcXoXy pes are designated for 13 species of opiine Braconidae and a neotype 
is designated for Bracon carbonarius Nees von Esenbeck, 1834. Three genera are trans- 
ferred to the Opiinae: Baeocentrum Schulz and Coeloreuteus Roman from the Rogadinae, 
and Neodiospilus Szepligeti from the Helconinae. All are synonyms of Opius Wesmael. 
Diachasmimorpha Viereck and Psyttalia Walker are elevated to generic rank in the Opi- 
inae, and Austroopius Szepligeti is placed as a subgenus of Psyttalia. Rhynchosteres (Fopius), 
New Subgenus, is described for several Ethiopian species previously referred to Biosteres. 
Four species of Opius are renamed, one new species, Rhynchosteres silvestrii, parasitic 
on Tephritidae is described, and 1 2 taxa are newly synonymized. Repositories are listed 
for the types of three species previously treated as nomina nuda or nomina dubia. 

Fischer (1963a, 1966, 1972, 1977) re- 
vised the opiine Braconidae from all major 
geographic regions of the World, and pub- 
lished a catalog of the known species (Fisch- 
er, 1971). Over the past 30 years, Fischer 
has also redescribed almost all of the older 
opiine species, thus providing a uniform 
format for comparison. His works are of 
particular value in pinpointing repositories 
for type specimens, and in clarifying the sta- 
tus of many uncertain species. A few prob- 
lems still remain, however (e. g. Fischer, 
1971, pp. 130-132), and the purpose of the 
present work is to solve some of these. 

The following abbreviations are used for 
museums: IRSN, Institut Royal des Sci- 
ences Naturelles de Belgique, Bruxelles 
(Wesmael Collection); TAMU, Texas A&M 
University, College Station; ZMHB, Zoo- 
logisches Museum der Humboldt Univer- 
sitaet. East Berlin (Foerster Collection); 

' Approved as TA #2 1 733 by the Texas Agricultural 
Experiment Station. 

BMNH, British Museum (Natural History), 
London; CNC, Canadian National Collec- 
tion, Ottawa; NRS, Naturhistoriska Riks- 
museet, Stockholm; WIEN, Naturhistorish- 
es Museum, Vienna; USNM, U.S. National 
Museum of Natural History, Washington, 
D.C. All of the lectotypes designated herein 
have been so labelled by me. 

Baeocentrum Schulz, 1911 

Szepligeti (1907) established the mono- 
typic genus Brachycentrus for a species col- 
lected by M. Rothschild in British East Af- 
rica. Szepligeti placed this genus in the 
Rogadinae. Schulz (1911), noting that Bra- 
chycentrus was preoccupied, renamed Sze- 
pligeti's genus as Baeocentrum. The type 
species, minutus Szepligeti, 1907, was de- 
scribed from two specimens. One of these, 
bearing Rothschild's blue locality label, a 
printed type label, and Szepligeti's hand- 
written type label, is in the Museum Na- 
tional d'Histoire Naturelle, Paris. I hereby 
designate this specimen as lectotype. The 
specimen matches Szepligeti's original de- 



scription, except that the specimen is a male 
and Szephgeti's description is of a female. 
The male genitaha are strongly protruding 
in this specimen, however, and this feature 
apparently misled Szepligeti. The paralec- 
totype (with metasoma missing) is conspe- 
cific, and located in the Hungarian Natural 
History Museum, Budapest. This species is 
clearly a member of the genus Opius s. /.. 
as characterized by Fischer (1972). I there- 
fore consider Bacocentrum to be a junior 
synonym of Opius, New Synonym. Opius 
niinulus (Szepligeti), New Combination, (not 
miuutus Granger, 1949, see below) fits the 
characterization of the subgenus Phlebose- 
ma Fischer, 1972. In Fischer's (1972) clas- 
sification, Fhlebosema thus becomes a ju- 
nior subjective synonym of Bacocentrum. 

Coeloreuteus Roman, 1910 

This genus was established by Roman 
(1910) for Atoreutcus africanus Szepligeti, 
1908, and placed in the Exothecinae. Shenc- 
felt (1975) retained it in the Rogadinac. I 
have examined the holotype oC africanus (in 
the NRS), and it belongs in Opius s. I. (as 
characterized by Fischer, 1972). The ab- 
sence of the mid-dorsal portion of the oc- 
cipital carina, the flat, exposed labrum, and 
the fore wing venation all support this place- 
ment. 1 therefore consider Coeloreuteus to 
be a junior synonym of Opius, New Syn- 
onym. I have not examined the other species 
listed under Coeloreuteus (Shenefelt, 1975) 
to determine their placement. 

transferred several species to it, and treated 
some of those included by Fullaway (1951) 
as synonyms of longicaudatus Ashmead. 
Wharton and Gilstrap (1983) further char- 
acterized this group on the basis of the api- 
cally sinuate ovipositor. Diachasmimorpha 
tryoni (Cameron), New Combination, is 
clearly a member of this group based on the 
shape of the ovipositor, wing venation, body 
sculpture, and shape of the clypeus, despite 
the almost complete absence of an occipital 
carina. Fischer (1967a) used the absence of 
an occipital carina as the basis for his genus 
Parasteres. Since the type species of Par- 
astcres is a synonym o{ tryoni (Wharton and 
Marsh, 1978), however, ParastercshtcornQS 
a junior subjective synonym of Diachas- 
mimorpha, New Synonym. The genus Di- 
achasmimorpha is very similar to the Neo- 
tropical genus Doryctobracon Enderlein, 
1920, based on similarities in clypeal mor- 
phology (lower margin of clypeus more or 
less sinuate, ventral-lateral margin of clyp- 
eus completely separated from face ventrad 
of the anterior tentorial pit). Doryctobracon 
lacks the sinuate ovipositor of Diachasmi- 
morpha, and generally has the recurrent vein 
antefurcal to interstitial rather than the 
postfurcal insertion of Diachasmimorpha. 
The clypeal morphology and fore wing ve- 
nation are, in my opinion, sufficient for re- 
moving Diachasmimorpha from Biosteres, 
where it has been placed by all recent au- 
thors (e.g. Fischer, 1972; Wharton and 
Marsh, 1978). 

Diachasmimorpha Viereck, 1913 

Gahan (1915) synonymized DiachastJii- 
morpha with Opius. Fullaway (1951) was 
the first to subsequently recognize this dis- 
tinctive group of tephritid parasitoids, which 
he called the Opius loni^icaudatus group (or 
group 1 ). Fullaway (1951) characterized this 
group by the presence of a postnervellus in 
the hind wing and the deep but unsculptured 
notauli. Wharton and Gilstrap (1983) re- 
ferred to this as the tryoni species group. 

Hexaulax Cameron, 1910 

Fischer ( 1 97 1 , p. 1 30) was unable to place 
this monotypic genus, stating only that it 
belonged "to the Opiinae according to 
Muesebeck." Muesebeck (1967, p. 48), 
however, treated He.xaula.x as a synonym 
of Opius on the authority of Fischer (1964). 
A clearly labelled specimen o( He.xaula.x ruf- 
iceps Cameron, 1910 (type species by 
monotypy of Hexaulax),. collected by 
Fruhstorfer and from the type locality in 



Java, and bearing a co-type label, is now in 
the BMNH. This specimen belongs to Opius 
s. /.. as interpreted by Fischer (1972), but 
cannot readily be assigned to a subgenus 
because a pin through the mesonotum makes 
it impossible to determine whether or not 
a midpit is present. Cameron stated in his 
original description that the occiput was not 
margined. The apparent absence of an oc- 
cipital carina is an artifact, however, due to 
the head being pinched in the occipital re- 
gion. Traces of the carina can still be seen 
on close inspection. Because of this occipital 
feature, the matching locality label, and the 
lack of evidence in the original description 
that more than one specimen was involved, 
this specimen may be the holotype. How- 
ever, in order to alleviate any confusion in 
the future, I am treating it as a lectotype, 
and hereby designate it as such. 

Lytacra Foerster, 1862 

Fischer (1959) stated that the type of Lv- 
tacra stygia Foerster, the type species of Lv- 
tacra, was lost. Fischer (1971) repeated this 
statement and placed stygia in Ademon Hal- 
iday, 1833 based on Foerster's original de- 
scription. A single female from Foerster's 
collection, labelled Lytacra stygia, is now 
present in the ZMHB; and I believe this to 
be the specimen on which Foerster (1862) 
based his description. The tips of the wings 
are missing and Foerster's characterization 
of the radial cell incompletely closed (and 
thus similar to the condition in Ademon) is 
therefore misleading (and in fact incorrect). 
Foerster's specimen is identical to the lec- 
totype and paralectotype of Diachasma caf- 
fer (Wesmael) in the IRSN. I therefore con- 
sider stygia to be a junior synonym of caffer, 
New Synonym, and Lytacra thus becomes 
a synonym of Diachasma. 

Neodiospilus Szepligeti, 1911 

Szepligeti (1911) included two species 
(/^r£'/?ra/or Szepligeti, 1911 and//av/";7^5 Sze- 
pligeti, 1911) in his original description of 
Neodiospilus. Brues (1926) subsequently 

designated fJavipes as the type species. 
Shenefelt ( 1 970), following Szepligeti (1911), 
placed Neodiospilus in the Diospilini, and 
listed the holotype of flavipes as being in the 
NRS. A search for this specimen in Stock- 
holm in 1984 was unsuccessful. However, 
a female collected by Fuellebom from the 
type locality in Langenburg on Lake Nyasa 
is now in the ZMHB. It bears a red type 
label, and a determination label in Szepli- 
geti's handwriting which reads "Neodio- 
spilus luteipes m"; and it matches Szepli- 
geti's original description of flavipes. In the 
drawer near this specimen is an old note 
saying the label = luteipes but the MS = 
flavipes. In the same drawer are the male 
and female syntypes of Neodiospilus tere- 
hrator, both with Szepligeti determination 
labels. I have no doubt that these are the 
specimens used by Szepligeti as the basis for 
his description of Neodiospilus, and consid- 
er the specimen labelled as luteipes to be the 
holotype of flavipes. Both flavipes and ter- 
ebrator are opiines, and belong in Opius s. 
I. as interpreted by Fischer (1972). Neo- 
diospilus thus becomes a junior synonym of 
Opius, New Synonym. There are pins ob- 
literating the mesonotal regions of the fla- 
vipes and terehrator specimens. The two 
species thus cannot readily be assigned to a 
subgenus, though it is clear from the clypeal 
morphology and wing venation that they 
should belong to different subgenera in 
Fischer's (1972) classification. 

Szepligeti (1914), in an apparent lapsus, 
described another genus with the name Neo- 
diospilus. The type series for the two in- 
cluded species, haumanni Szepligeti and 
zenkeri Szepligeti, are also in the ZMHB. 
Shenefelt ( 1 970) renamed Neodiospilus Sze- 
pligeti, 1914 as Repetiodiospilus. It is cor- 
rectly placed in the Diospilini (as presently 

Psyttalia Walker, 1860 

Muesebeck (1931) synonymized Psyttalia 
with Opius, and renamed the type species 
walkeri Muesebeck. Fischer (1971) listed 



Psyttalia under Opiiis, but inadvertently 
omitted walkeri from his species catalog. 
Fischer (1972) treated Psyttalia as a sub- 
genus of Opius. Psyttalia walkeri belongs to 
the concolor species group as characterized 
by Wharton and Gilstrap (1983). As noted 
by Wharton and Gilstrap (1983), the con- 
color and fletcheri species groups share a 
number of characters suggestive of a close 
relationship. In addition to characters noted 
by Fischer (1972), these include reduced 
mesonotal sculpture, large, attenuate hy- 
popygium, short clypeus, exposed labrum 
with reduced setation, and identical hind 
wing venation. I believe these characters are 
sufficient to elevate Psyttalia to generic rank 
in the Opiinae, with Austroopius Szepligeti, 
1900 included as a subgenus. Members of 
the subgenus Psyttalia (= concolor species 
group) generally have the gaster weakly 
sculptured, at least basally, while in the sub- 
genus Austroopius (= the fletcheri species 
group) the recurrent vein is somewhat 
bowed. I do not consider the thickening of 
the first cubital cross- vein adequate for sep- 
arating the species formerly included in 
Austroopius (e.g. Fischer, 1963b, 1966) from 
closely related species, such a.s fletcheri Sil- 
vestri and incisi Silvestri, which lack this 

Opius africanus Szepligeti, 1910 

With the transfer ofCoeloreuteus to Opius 
(see above), Opius africanus (Szepligeti, 
1908) becomes a senior secondary hom- 
onym oi Opius africanus Szepligeti, 1910. 
The latter is therefore renamed Opius teph- 
ritivorus, New Name. Opius tephritivorus is 
a member of the subgenus Utetes Foerster, 
as characterized by Fischer (1972), and is 
at least subgenerically different from afri- 

Pachythecus albobalteatus Cameron, 1912 

This species was transferred to Biosteres 
by Fischer (1967b). The sculpture and gen- 
eral color pattern of the male holotype in 

the BMNH (BM type # Hym. 3.C.71 1) are 
the same as that ofangaleti Fullaway, 1952. 
Although the mid coxae tend to be darker 
in most specimens of angaleti that I have 
seen, I can find no other differences, and 
therefore treat angaleti as a junior subjec- 
tive synonym of albobalteatus, New Syn- 
onym. The New Combination is Diachas- 
mimorpha albobalteatus (Cameron). The 
type locality of albobalteatus is "Borneo," 
that of angaleti North Borneo (= Sabah). 

Diachasma brevistyli Paoli, 1934 

Fischer (1971) treated brevistyli as a no- 
men nudum, referring only to a subsequent 
note on brevistyli by Martelli (1937). 
Through the courtesy of the librarians at the 
Museum "La Specola" in Firenze, Italy, I 
was able to locate Paoli's (1934) original 
description, and therefore consider it (as well 
as Phanerotoma somalica Paoli, 1934) to 
be a validly described species. The type se- 
ries Q){ brevistyli is in the Museum "La Spe- 
cola." This species is almost identical to 
Biosteres carinatus Szepligeti, 1910; and the 
generic placement of these two species is 
currently under investigation. 

Opius cajfer Wesmael, 1835 

Wesmael (1835) described caffer on the 
basis of two specimens, a male and a female. 
Both are in excellent condition in the IRSN. 
The female is hereby designated as lecto- 
type. It bears the following labels: top label: 
Coll. Wesmael (printed); 2nd label: 1865 
(printed); 3rd label: Opius caffer mihi (hand- 
written) det. C. Wesmael (printed); 4th la- 
bel: Type (printed in red). 

Bracon carbonarius Nees von Esenbeck, 

See procerus Wesmael (below). 

Biosteres carinatus Szepligeti, 1910 

I have examined the holotype male in the 
NRS and am unable to separate this species 
from Hedylus gijfardii Silvestri (as charac- 



terized by Wharton and Gilstrap, 1983). I 
therefore consider gijfardii to be a junior 
subjective synonym ofcarinatus, New Syn- 
onym. See additional comments under 
brevistyli (above). 

Opius cingulatus Wesmael, 1835 

This is the type species (by original des- 
ignation) of Nosopoea Foerster, 1862. Wes- 
mael (1835) described cingulatus on the ba- 
sis of 14 specimens. Eleven specimens, 
bearing identical type and determination la- 
bels, but representing at least three species, 
are now present in the IRSN. I hereby des- 
ignate as lectotype one of four females most 
closely matching the original description, 
and bearing the following labels: top label: 
1832 (printed); 2nd label: Coll. Wesmael 
(printed); 3rd label: Type (printed in red); 
4th label: S Opius 9 cingulatus. mihi (hand- 
written) det. C. Wesmael (printed). The lec- 
totype agrees with Fischer's (1972) defini- 
tion of cingulatus; and also fits the concept 
of cingulatus used by Haliday (1837), Foers- 
ter (1862), Marshall (1891) and Thomson 
(1895). The lectotype has the hypostomal 
and occipital carinae widely separated at the 
mandible, and the mandible bears a carinate 
ridge basally on its ventral border. Paralec- 
totypes which are not conspecific have the 
oral and occipital carinae meeting near the 
base of the mandible, the stemaulus at least 
weakly sculptured, and/or a broad basal lobe 
ventrally on the mandible (as in O. pallipes 
Wesmael or O. crassipes Wesmael). 

Opius cojfeae Fischer, 1962 

I have been unable to locate the holotype 
of cojfeae, but a paratype in the Naturhis- 
torisches Museum, Basel is identical to the 
lectotype of Biosteres caudatus Szepligeti, 
1913. I therefore treat cojfeae as a junior 
subjective synonym of caudatus. New Syn- 
onym. If the holotype is ever located, it 
should be examined to confirm this syn- 
onymy. This species is characterized by the 
narrow patch of deep punctures between eye 

and ocelli (Wharton and Gilstrap, 1983, Fig. 
12). Generic placement is discussed below 
under silvestrii, n. sp. 

Opius comatus Wesmael, 1835 

This is the type species (by original des- 
ignation) of Holconotus Foerster, 1862. 
Ashmead (1900) later proposed the name 
Aulonotus for Foerster's Holconotus, which 
was preoccupied. Wesmael (1835) de- 
scribed comatus on the basis of 1 1 speci- 
mens. There are now eight specimens in 
IRSN, two of which fit the description of 
Wesmael's var. 1 . 1 hereby designate as lec- 
totype of comatus the female bearing R. 
Koenig's 1968 lectotype label. Koenig un- 
fortunately never published a lectotype des- 
ignation for this species. The specimen has 
the following additional labels: top label: 
Coll. Wesmael (printed); 2nd label: 1860 
(printed); 3rd label: <5 Opius 2 comatus. mihi 
(hand-written) det. C. Wesmael (printed); 
4th label: Type (printed in red); 5th label: 
R. I. Sc. N. B. I. g. 3.317; 6th label: LEC- 
TOTYPE Opius comatus Wesmael 9 de- 
signe par R. Konig 1968. 

There are two more females bearing type 
and comatus determination labels (includ- 
ing the additional specimen of var. 1) than 
hsted by Wesmael (1835) in the original de- 
scription. It is not possible at present to 
determine whether specimens were added 
after the original description, or Wesmael 
erred in listing the sexes. Labelling of para- 
lectotypes has therefore been conservative. 

Opius crassipes Wesmael, 1835 

This is the type species (by original des- 
ignation) of Hypocynodus Foerster, 1862. 
Wesmael (1835) described crassipes on the 
basis of a single female, and also indicated 
that he had a male which possibly belonged 
to this species. In the Wesmael Collection 
there are now two females plus a third spec- 
imen lacking an abdomen. All bear Opius 
crassipes type labels. The two intact females 
are not conspecific, however, and I have 



labelled the one that fits Wesmael's original 
description as the holotype. 

Neodiospilus flavipes Szepligeti, 1911 

^incQ flavipes is preoccupied in Opius by 
flavipes Szepligeti, 1898, it becomes nec- 
essary to rename the junior homonym fol- 
lowing transfer of the type species of Neo- 
diospilus to Opius (see above). It is therefore 
renamed Opius sepivalfus, New Name. The 
specific epithet is an arbitrary combination 
of letters. 

Opius irregularis Wesmael, 1835 

This is the type species ofAllotypus Foers- 
ter, 1862. Wesmael (1835) described Opius 
irregularis on the basis of one male and five 
females. Six specimens (2 6, 4 9) matching 
his original description and bearing iden- 
tical type and determination labels are in 
the IRSN. These specimens also bear the 
lectotype and paralectotype labels of R. 
Koenig. Unfortunately, Koenig's lectotype 
designation has never been published. Koe- 
nig labelled a male as lectotype, but I believe 
the female offers better characters for species 
discrimination. I therefore designate as lec- 
totype a female with the following labels: 
top label: Coll. Wesmael (printed); 2nd la- 
bel: 1847 (printed); 3rd label: S Opius 9 ir- 
regularis mihi (hand-written) det. C. Wes- 
mael (printed); 4th label: Type (printed in 
red); 5th label: PARALECTOTYPE Opius 
irregularis Wesmael 9 designe par R. Konig 

Opius lev is Wesmael, 1835 

This is the type species of the subgenus 
Opiothorax Fischer, 1972. Wesmael (1835) 
described Opius levis on the basis of seven 
specimens. There are now six specimens in 
the IRSN. I hereby designate as lectotype 
the female labelled by Koenig in 1968 as 
lectotype (but never published). The spec- 
imen has the following additional labels: top 
label: Coll. Wesmael (printed); 2nd label: 
1835 (printed); 3rd label: Opius levis mihi. 
3. 9 (hand-written) det. C. Wesmael (print- 

ed); 4th label: Type (printed in red); 5th 
label: R. I. Sc. N. B. I. G. 3.317; 6th label: 
LECTOTYPE Opius levis Wesmael 9 de- 
signe par R. Konig 1 968. Fischer (1972) rec- 
ords this as a color-variable species, but 
specimens with dark hind coxae will not run 
to levis in his key (Fischer 1972, p. 441). 
Wesmael (1835) specifically mentions the 
dark coxae in his original description; all 
members of the type series have this feature. 

Rhogadopsis miniacea Brethes, 1913 

This is the type species (by monotypy) of 
Rhogadopsis Brethes, 1913. It was trans- 
ferred to Opius by de Santis (1967). I des- 
ignate as lectotype a female in the Museo 
"Bernardino Rivadavia" (Buenos Aires) 
with a single label in Brethes' handwriting 
which reads Rhogadopsis miniacea Br. The 
species runs to the tucumanus -group ofLis- 
sosema in Fischer (1977). Rhogadopsis thus 
becomes a senior synonym of the subgenus 
Lissosema Fischer, 1972, New Synonym. 

Opius minutus Granger, 1949 

It becomes necessary to rename the junior 
homonym following transfer of Brachycen- 
trus minutus Szepligeti, 1907 to Opius (see 
above). Opius minutus Granger is therfore 
renamed Opius gregnar. The specific epithet 
is an arbitrary rearrangement of the name 

Opius pallipes Wesmael, 1835 

This is the type species (by subsequent 
designation of Muesebeck and Walkley, 
1951), of Opius Wesmael, 1835. Wesmael 
(1835) described this species on the basis of 
15 females and five males. There are now 
19 specimens in the IRSN. I designate as 
lectotype a female with abdomen mounted 
beneath the rest of the specimen on a sep- 
arate card, and characterized by an excep- 
tionally smooth petiole and propodeum, 24 
segmented antenna (right side, left side bro- 
ken), and pale clypeus. This characteriza- 
tion fits just within the range of variation 
given for this species by Wesmael. The lee- 



totype was selected specifically on the basis 
of the sculptural characters of the petiole 
and propodeum so as to more clearly define 
this species. It should be pointed out, how- 
ever, that studies on intraspecific variation 
in sculptural features have not been con- 
ducted for this species and are desperately 
needed. The diverse array of host records 
for pal I ipes (e. g. Fischer, 1971) suggests that 
some confusion has existed in the past re- 
garding the application of this name. It is 
hoped that restriction of the definition of 
paUipes by means of this lectotype desig- 
nation will enable future workers to more 
accurately identify this species; and that this 
in turn will lead to a re-examination of pub- 
lished host records. The species is of poten- 
tial use in biological control of Liriomyza 
(Hendrikse, 1980). The lectotype bears the 
following labels: top label: Coll. Wesmael 
(printed); 2nd label: 1831 (printed); 3rd la- 
bel: <5 Opius 9 pallipes mihi (hand-written) 
det. C. Wesmael (printed); 4th label: Type 
(printed in red). 

Opius procerus Wesmael, 1835 

Wesmael (1835) described this species 
from two individuals. There are still two 
specimens (both male) of this species in the 
IRSN labelled as types. I hereby designate 
as lectotype the male bearing R. Koenig's 
Opius carbonarius (Nees) determination la- 
bel. It has the following additional labels: 
top label: Coll. Wesmael (printed); 2nd la- 
bel: 1872 (printed); 3rd label: Opius pro- 
cerus, mihi 6 (hand-written) det. C. Wes- 
mael (printed); 4th label: Type (printed in 
red). I designate this specimen as also the 
neotype of Bracon carbonarius Nees von 
Esenbeck, 1834. The vast majority of Nees 
von Esenbeck's collection, including the 
type(s) o{ carbonarius, have been destroyed 
(e. g. Shenefeh, 1970; Papp, 1985; and Van 
Achterberg, pers. comm.). Haliday (1837) 
treated procerus Wesmael as a synonym of 
carbonarius Nees, and I consider Haliday 
as the first reviser relative to the establish- 
ment of a fixed concept for carbonarius. Al- 

though this concept is in agreement with the 
latest treatment of carbonarius (Fischer, 
1977), at least two major authors (Wesmael, 
1835; Thomson, 1895) have interpreted 
carbonarius differently. Bracon carbonarius 
is the type species of Biosteres Foerster, 
1862. The subgeneric concepts used by 
Fischer (1972) for Biosteres are based on 
Haliday's interpretation o{ carbonarius, and 
are therefore in conflict with the interpre- 
tations of Wesmael and Thomson. A neo- 
type designation for carbonarius will resolve 
this conflict. Fischer (1972, 1977) has pre- 
sented keys for the separation of carbon- 
arius from closely related species. 

Opius ruficeps Wesmael, 1835 

This is the type species (by original des- 
ignation) of Therobolus Foerster, 1862. 
Wesmael (1835) described this species from 
two specimens. There are now three speci- 
mens in the Wesmael Collection, all bearing 
the identical type and determination labels 
noted above for other Wesmael species. I 
hereby designate as lectotype a female hav- 
ing an additional label with the hand-writ- 
ten number 3 on it. 

Hexaulax ruficeps Cameron, 1910 

Transfer of the type species of Hexaulax 
to Opius results in a secondary homonym. 
I therefore rename ruficeps Cameron (not 
ruficeps Wesmael, 1835) as Opius indenta- 
tus. New Name, in reference to the damaged 
lectotype and misleading original descrip- 
tion (Cameron, 1910) resuhing therefrom. 

Celiestiella testaceipes Cameron, 1903 

This is the type species (by monotypy) of 
Celiestiella Cameron, 1 903. Cameron ( 1 903) 
mentioned only the male sex in his ex- 
tremely brief description of this species. 
However, the clearly labelled type series in 
the BMNH (BM type # Hym 3.C.713) con- 
sists of two males and one female. Of the 
two males, one is damaged by the pin. I 
therefore designate the other male as lec- 
totype. Fischer (1967b) redescribed this 



species from the female, and transferred it 
to Biosteres. I agree with this placement. 

Opius testaceus Wesmael, 1838 

This is the type species (by original des- 
ignation) of Utetes Foerster, 1862. Wesmael 
(1838) described this species from four fe- 
males and one male, but there are now only 
three specimens in the IRSN. I hereby des- 
ignate as lectotype the remaining intact fe- 
male. It bears the following labels: top label: 
119. (hand-written); 2nd label: Coll. Wes- 
mael (printed); 3rd label: Type (printed in 
red); 4th label: O. testaceus. mihi (hand- 
written) det. C. Wesmael (printed); 5th la- 
bel: cf. Opius (hand-written) Rev. J. A. Mar- 
shall (printed) 1887 (hand-written). 

Biosteres testaceus Szepligeti, 1914 

I have examined the holotype female in 
the ZMHB, and am unable to separate this 
species form Biosteres fullawayi Silvestri, 
1913. I therefore treat testaceus Szepligeti 
as a junior subjective synonym o^ fullawayi, 
New Synonym. This species was recently 
diagnosed by Wharton and Gilstrap (1983). 
It's generic placement is currently under in- 

Description of New Taxa 

Most of the Ethiopian and Indo-Pacific 
species currently placed in Biosteres are not 
readily accommodated by the existing opiine 
classification. Most authors, following 
Fischer (1963a, 1966), have placed the in- 
cluded species in Biosteres on the basis of 
the short second cubital cell. However, this 
feature is inadequate for characterizing 
opiine genera. Closer examination of the 
wing venation shows that there are signifi- 
cant differences between Biosteres s. s. and 
these Ethiopian and Indo-Pacific species. 
The most obvious difference is in the shape 
of the stigma. It is longer and narrower in 
Biosteres s. s., with the first radial segment 
arising basad of the midpoint. The post- 
nervellus is also greatly reduced in Biosteres 
s. s., and is often only barely visible as a 

weakly infumate crease. Other major dif- 
ferences are in the shape of the mandible, 
which has a basal tooth in Biosteres s. s.; 
and in the host preferences. 

Some of the Indo-Pacific species formerly 
placed in Biosteres have now been trans- 
ferred to Diachasmimorpha (see above). 
Based on the differences enumerated in the 
preceding paragraph, I believe most of the 
remaining Indo-Pacific and Ethiopian 
species are more closely related to Diachas- 
mimorpha and Rhynchosteres Fischer, 1 965 
than to Biosteres s. s. I prefer to retain Di- 
achasmimorpha as restricted by the defi- 
nition given earlier in this paper. Since the 
new species described below does not fit this 
restricted definition, I have placed it in 

Rhynchosteres Fischer, 1965 
Fopius Wharton, New Subgenus 

Type species: Rhynchosteres silvestrii, new 

Occipital carina absent dorsally, but well- 
developed laterally; oral and hypostomal 
carinae widely spearated at mandible. La- 
bral setae sparse, confined to ventral and 
lateral margins; labrum concealed by large, 
somewhat hemispherical clypeus with con- 
vex to nearly truncate lower margin; margin 
of clypeus extending ventrally from anterior 
tentorial pit somewhat reflected, and sepa- 
rated from gena by a sharp groove. Man- 
dibles gradually and evenly tapering distal- 
ly, without basal tooth or lobe, not deflected 
ventrally; ventral tooth shorter than dorsal 
tooth. Apical antennal segment with spine- 
like tip. Genal sulcus shallow, often ob- 
scured by facial sculpture. Second radial 
segment shorter than 1 st cubital cross- vein; 
recurrent vein antefurcal to interstitial; first 
radial segment arising from or distad of mid- 
stigma; stigma discrete, not merging im- 
perceptibly with metacarpus distally; post- 
nervellus long, distinctly sclerotized and 
pigmented; radiella absent at base, often in- 
dicated by fold or infumate trace distally. 



Pronotum largely unsculptured and without 
median pit. Stemaulus crenulate; notauli 
complete to midpit and crenulate through- 
out. Postpectal carina usually present an- 
teriorad mid coxae. Claws simple, with 
broad base. Petiole with deep glymma; dor- 
sope present or absent. Spiracles of terga 
2+3 near lateral margins (tergum 3 spi- 
racle often not visible in dorsal view due to 
postmotem collapse of tergum). Hypopyg- 
ium attenuate. Known species are parasit- 
oids of tephritids, and have the ovipositor 
at least 1.5 times length of mesosoma. 

Diagnosis.— Rhynchosteres differs from 
other large tropical opiines with short sec- 
ond cubital cell and broad, discrete stigma 
as follows: the occipital carina is absent in 
Doryctobracon\ the ovipositor is sinuate in 
Diachasmimorpha, and the mandible has a 
well-developed flange along the ventral bor- 
der; the pronotum has a large median pit in 
Pseudorhinoplus Fischer, 1972. Rhynchos- 
teres was described for species with a dis- 
tinctively protruding and medially reflected 
clypeus (Fischer, 1965; Van Achterberg, 
1983). In Rhynchosteres (Fopius), the clyp- 
eus does not protrude in this fashion. 
Nevertheless, some of the described species 
have the clypeus thickened ventral-medi- 
ally, and I believe this is the initial step in 
the transformation to the condition found 
in Rhynchosteres s. s. Once the Afrotropical 
fauna becomes better known, I suspect the 
distinction between Rhynchosteres s. s. and 
R. (Fopius) will be less obvious. The per- 
sulcatus species group (FuUaway, 1951; 
Wharton and Gilstrap, 1983) is somewhat 
intermediate between Diachasmimorpha 
and R. (Fopius); and needs more detailed 
study before it can be accurately placed. 

I include the following species in Rhyn- 
chosteres (Fopius): bevisi (Brues), New 
Combination, caudatus (Szepligeti), New 
Combination, desideratus (Bridwell), New 
Combination, niger (Szepligeti), New Com- 
bination, ottotomoanus (FuUaway), New 
Combination, and pyknothorax (Fischer), 
New Combination. At least three species 

groups are evident, based on clypeal mor- 
phology and sculpture of the petiole and 
frons. I transfer also clypeatus (Bridwell) to 
Rhynchosteres s. s., New Combination. 

The subgeneric name is masculine and is 
a contraction formed from the words Fisch- 
er and Opius. 

The type species has been misidentified 
in the past as Biosteres caudatus, and has 
recently been referred to as caudatus auct. 
(Wharton and Gilstrap, 1983; Steck et al., 
1986). I have been unable to find an avail- 
able name for this species, and it is therefore 
described as new. 

Rhynchosteres (Fopius) silvestrii 
Wharton, New Species 

Figs. 1,2 

¥Qma\Q. — Head: 1.66 ± 0.06 times 
broader than long; 1.35 ± 0.05 times broad- 
er than mesonotum; face with deep punc- 
tures and well-developed midridge, the lat- 
ter extending between antennal bases as a 
low, sharp ridge; frons varying from nearly 
unsculptured in small individuals to exten- 
sively rugosopunctate (variation as in 
Wharton and Gilstrap, 1983; Figs. 10, 11); 
ocellar triangle margined, at least in part, 
and often completely, by a crenulate sulcus. 
Ventral margin of clypeus weakly convex, 
not noticeably thickened in middle. Eyes 
bare or apparently so, large, 2.36 ± 0.43 
times longer than temples; temples receding 
in dorsal view. Antenna roughly 3.4 times 
longer than mesosoma; 1st flagellomere 
subequal to 2nd. 

Mesosoma: 1.28 ± 0.05 times longer than 
high, 1.67 ± 0.04 times longer than broad. 
Median mesonotal lobe with 2 parallel, ru- 
gosopunctate grooves anteriorly; lateral me- 
sonotal lobes hairy, and usually with scat- 
tered, deep punctures; with setae confined 
to margins in smaller species; notauli meet- 
ing in a narrow midpit, midpit varying from 
discrete furrow bordered anteriorly by striae 
to a small rugosostriate patch of sculpture. 
Propodeum rugose, sculpture variable, but 
often with obvious transverse elements on 



Fig. 1 . Fore and hind wing of Rhynchosteres (Fopius) silvestrii. 

anterior face, sculpture on posterior, decli- 
vous face more uniform and slightly weak- 
er; flange-like midridge present anteriorly. 
Stemaulus broad, deep, crenulate, but usu- 
ally absent over posterior one-third; cren- 
ulate sculpture extending dorsally along an- 
terior border of mesopleuron through 
subalar depression; posterior margin of me- 
sopleuron noticeably crenulate ventrad of 
speculum, suture unsculptured or nearly so 
dorsad of speculum; mesopleural disc hairy; 
postpectal carina usually well-developed 

Wing: 2nd radial segment 2.23 ± 0.29 
times longer than 1st; 1st cubital cross- vein 
1.22 ± 0.09 times longer than 2nd radial 
segment; 3rd radial segment extending near- 
ly to wing tip; cubitus arising from about 
anterior 0. 1 5 of basal vein, with 1 st segment 

sinuate; nervulus inclivous, usually post- 
furcal by less than its length, more rarely 
interstitial; junction between nervulus and 
basal vein thickened; recurrent vein ante- 
furcal to interstitial; parallel vein arising be- 
low middle of brachial cell. Postnervellus 
reclivous, though usually weakly recurved 
posteriorly, extending nearly to posterior 
margin as a well-developed, deeply im- 
pressed crease which is usually weakly pig- 

Metasoma: Petiole 1.00 ± 0.05 times 
longer than apical width, apex 2.04 ±0.14 
times wider than base; finely striate, with a 
stronger median longitudinal carina some- 
times evident; dorsal carinae well-devel- 
oped basally, weak over posterior half, but 
usually extending to posterior margin, very 
weakly converging (at posterior margin, dis- 



tance between carinae usually equal to or 
slightly less than distance to lateral margin); 
dorsope well-developed. Metasoma un- 
sculptured beyond petiole. Ovipositor tip 
not narrowed, with barely developed dorsal 
notch and weak ventral serrations; 2.56 ± 
0. 1 5 times longer than mesosoma; ovipos- 
itor sheath dorsally with 2 rows of setae, 
each row with about 35 setae, setae shorter 
and about twice the density ventrally. 

Color: Quite variable, generally reddish- 
brown to black; palps white to pale yellow; 
scape, pedicel, mandibles (except tips), all 
legs, and terga 2 + 3 yellow (remaining terga 
usually darker); clypeus, gena, orbits, 
pronotum, and antenna variable, but usu- 
ally at least partly yellow; wings hyaline. 

Male. — Somewhat darker than female, 
with all metasomal terga brown to black 
(rarely pale at extreme base of tergum 2). 
Otherwise essentially as in female, but with 
mesonotum slightly narrower, midpit often 
weakly developed to more or less absent, 
and petiole 1.21 ± 0.07 times longer than 
apical width (and thus distinctly longer and 
narrower than in female). 

Length (exclusive of ovipositor): 2.0-3.5 

Material examined. — Holotype 9, CAM- 
EROON: Centre-Sud Province, Nkolbis- 
son, 22.VIII.1982, G. Steck, reared ex ripe 
coffee berries (USNM). Paratypes (TAMU, 
CNC, WIEN): Same locality, host plant, and 
collector as holotype: IV. 18. 1982 (1 9), 
V.1982 (1 (5), 22.VIII.1982 (9 9, 4 <5), 
17.IX.1982 (2 9, 1?), 26.IX.1982 (2 9, 1 <5), 
IX.1982 (5 9, 2 $), 3.X.1982 (4 9, 2 <5), 
10.X.1982 (6 9, 6 3), 17.X.1982 (6 3), 
26.X.1982 (4 9, 1 3), X.1982 (10 9, 46 <5), 
1.XI.1982 (2 9, 3 3), 8.XI.1982 (2 9, 1 3), 
15.XI.1982 (4 9, 10 3), 22.XI.1982 (4 9, 13 
3), XI. 1 982 (3 9, 2 3); CAMEROON: Centre- 
Sud Prov., Akonolinga, VI. 1982 (1 9, 1 3); 
24. VII. 1982 (1 9); 28. VIII. 1982 (1 3), 
VIII.1982 (5 9, 13 3); IX.1982 (1 9); X.1982 
(2 9), G. Steck, reared from Coffea robusta 
berries, with Trirhithrum cojfeae (Bezzi) 
(Tephritidae) as probable host; Ouest Prov., 

Fig. 2. Ovipositor tips. A, Rhynchosteres (Fopius) 
silvestrii. B, R. (F.) caudatus (Szepligeti). 

Bafoussam, 30.VI.1982 (1 9), 13.VII.1982 
(1 9), G. Steck, reared from Coffea arabica 
berries; Centre-Sud Prov., Etoug-Ebe, 
26.V.1982, G. Steck, reared from Dacus on 
squash (1 9); Yaounde, Agr. Res. Inst., 
2.VI.1980, F. E. Gilstrap, W. G. Hart, & D. 
Perkins, from coffee berries (12 9). COSTA 
RICA: Turrialba, 12.XI.1982, M. Fischel, 
recovered from release site in coffee plan- 
tation (1 3); shipped to Costa Rica from 
Cameroon, via Texas, IX.23.1982 (3 9), 
XI. 1982 (3 9). TOGO: Kpime-Seva, 
16.XII.1981, G. Steck, from Coffea robusta 
berries (1 9). 

The material collected in 1982 was 
shipped through the Texas A&M Univer- 
sity (TAMU) quarantine as #s T82001, 
T82009, T82013, T82031, T82033, 
T82034, T82036, T82039, T82040, 
T^2041, T82044, T82046, and T82047. 
Further details are provided by Steck et al. 

Diagnosis. — This species has frequently 
been reared with R. (F.) caudatus from a 
variety of tephritids in West Africa; and has 
often been confused with the latter (Whar- 
ton and Gilstrap, 1983; Steck et al., 1986). 
In caudatus, the dorsope is not developed, 
the clypeus tends to be thickened medially 
along the ventral margin, the ovipositor is 
slightly shorter (usually less than twice length 
of mesosoma), the frons bears a very dis- 
tinctive transverse row of deep, close-set 
punctures on an otherwise unsculptured 
backgorund (frons weakly to heavily rugose 
in silvestrii), and the median mesonotal lobe 
has only a single longitudinal groove me- 
dially. The two species probably attack dif- 
ferent immature stages of their host tephri- 
tids since the ovipositor is exceptionally 



narrow at the tip in caudatus (Fig. 2). A 
similar situation was described in detail by 
Van den Bosch and Haramoto (1951) for 
members of the persulcatus species group. 

A much closer relationship exists between 
silvestrii and pyknothorax, and these two 
species are difficult to separate. However, 
the latter, a South African species, lacks 
punctures and setae on the lateral lobes of 
the mesonotum, and the clypeus is slightly 
thickened mid-ventrally. 

Discussion.— This species is named in 
honor of the Italian entomologist Filippo 
Silvestri, who did much to further our 
knowledge of opiine biology through his bi- 
ological control efforts. The above mea- 
surements (given as a mean ± standard de- 
viation) are based on 20 specimens each. 


I am most grateful to the following cu- 
rators and assistants, without whose help 
this work could not have been accom- 
plished: A. Bachmann (Buenos Aires), P. 
Dessart (IRSN), E. Haeselbarth (Munich), 
T. Huddleston (BMNH), F. Koch (ZMHB), 
P. Marsh (USDA, USNM), S. Mascherini 
(Firenze), J. Papp (Budapest) and A. Sten- 
mark (NRS). I thank also M. Fischer (Wien), 
C. Van Achterberg (Leiden), J. Schaffiier 
(College Station), and F. Bin (Perugia) for 
assistance and fruitful discussions, and W. 
Hart, F. Gilstrap, and G. Steck for making 
available the material on which the new 
species was based. Finally, I thank R. Koe- 
nig for graciously allowing me to publish 
the lectotype designations for Wesmael's 
Opius species. 

Literature Cited 

Ashmead,W. H. 1900. Some changes in generic names 
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Brues, C. T. 1926. Studies on Ethiopian Braconidae 
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Cameron, P. 1903. Descriptions of new genera and 
species of Hymenoptera from India (continua- 
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. 1910. On some Asiatic species of the subfam- 

ilies Braconinae and Exothecinae in the Royal Ber- 
lin Museum. Soc. Entomol. 25: 1 1-12, 14-16, 19- 
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De Santis, L. 1967. Catalogo de los Himenopteros 
argentinos de la serie Parasitica, incluyendo Be- 
thyloidea. Com. Invest. Cient. Prov. Buenos Aires, 
La Plata, pp. 33-34. 

Fischer, M. 1959. Die europaischen Opiinae (Hy- 
menoptera, Braconidae). Acta Entomol. Mus. Nat. 
Pragae 33: 241-263. 

. 1963a. Die Opiinae derathiopischen Region. 

Beitr. Entomol. 13: 194-221, 662-747. 

. 1963b. Das Genus Austroopius Szepligeti 

(Hymenoptera, Braconidae, Opiinae). Mitt. Zool. 
Mus. Beriin 39: 173-186. 

. 1964. Die Opiinae der nearktischen Region 

(Hymenoptera, Braconidae). I. Teil. Polski Pismo 
Entomol. 34: 197-530. 

. 1965. Zwei neue Opiinen — Gattungen aus 

dem Kongo ( 1 . Beitrag ueber die Opiinae des Mu- 
see Royal de I'Afrique Centrale) (Hymenoptera, 
Braconidae). Rev. Zool. Bot. Afr. 71: 309-323. 

. 1966. Revision der indo-australischen Opi- 
inae. Dr. W. Junk B. V., The Hague. 167 pp. 

. 1967a. Zusammenfassung der neotropischen 

Opiinae mit Ausschluss der Gattung Opius Wesm. 
(Hymenoptera, Braconidae). Beitr. Neotrop. Fau- 
na. 5: 1-21. 

. 1967b. Redeskriptionen von einigen Opiinen 

(Hymenoptera, Braconidae). Arbeitsgemeinschaft 
Oesterr. Entomol. 19: 59-69. 

. 1971. Hym. Braconidae. Index of Worid Opi- 

inae. Index of entomophagous insects. Le Fran- 
cois, Paris. 189 pp. 

. 1972. Hymenoptera Braconidae (Opiinae I). 

DasTierreich 91: 1-620. 

. 1 977. Hymenoptera Braconidae (Opiinae II — 

Amerika). Das Tierreich 96: 1-1001. 

Foerster, A. 1862. Synopsis der Familien und Gat- 
tungen der Braconen. Verh. naturh. Ver. preuss. 
Rheinl. 19: 225-288. 

Fullaway, D. T. 1951. Review of the Indo- Austra- 
lasian parasites of the fruit flies (Tephritidae). Proc. 
Haw. Entomol. Soc. 14: 243-250. 

Gahan, A. B. 1915. A revision ofthe North American 
ichneumon-flies of the subfamily Opiinae. Proc. 
U.S. Natl. Mus. 49: 63-95. 

Haliday, A. H. 1837. Essay on Parasitic Hymenop- 
tera. Entomol. Mag. 4: 203-221. 

Hendrikse, A. 1980. A method for mass rearing two 
braconid parasites {Dacnusa sibihca and Opius 
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roniae). Meded Fac. Landbou. Rijksuniv. Gent 
45: 563-571. 

Marshall, T. A. 1891. A monograph of the British 
Braconidae. Part IV. XXIII. Opiides. Trans. Ento- 
mol. Soc. Lond. 1891: 7-61. 

Martelli, G. M. 1937. Contributo alia conoscenza 



biologica del Dacus oleae Rossi e dei suoi parassiti 
in Tripolitania. Agric. Colon. 31(A): 149-155. 

Muesebeck, C. F. W. 1931. Descriptions of a new 
genus and eight new species of ichneumon-flies 
with taxonomic notes. Proc. U.S. Natl. Mus. 79: 

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Krombein, K. V. and B. D. Burks. Hymenoptera 
of America north of Mexico, synoptic catalog. Sec- 
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Muesebeck. C. F. W. and L. M. Walkley. 1951. Fam- 
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W., K. V. Krombein, and H. K. Townes. Hyme- 
noptera of America north of Mexico, synoptic cat- 
alog. USDA Agric. Mon. 2. 1420 pp. 

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Papp, J. 1985. Taxonomical and faunistical novelties 
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Roman, A. 1910. Notizen zur Schlupfwespensamm- 
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Schulz, W. A. 1911. Zweihundert alte Hymenopter- 
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Shenefelt, R. D. 1970. Pars 5. Braconidae 2. Hel- 
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In Ferriere, Ch. and J. van der Vecht, eds., Hy- 
menopterorum catalogus (nova editio). Dr. W. Junk 
B. v., The Hague, pp. 177-306. 

. 1975. Pars 12. Braconidae 8. Exothecinae 

Rogadinae. In van der Vecht, J. and R. D. Shene- 
felt, eds., Hymenopterorum catalogus (nova edi- 
tio). Dr. W. Junk B. V., The Hague, pp. 1115- 

Steck, G. J., F. E. Gilstrap, R. A. Wharton, and W. G. 
Hart. 1986. Braconid parasitoids of Tephritidae 
(Diptera) infesting coffee and other fruits in West- 
Central Africa. Entomophaga 31: 59-67. 

Szepligeti, G. 1907. Collections faites par M. le Baron 
Maurice de Rothschild dans TAfrique Orientale. 
Bull. Mus. Hist. Nat. Paris 1907: 34-36. 

. 1911. Braconidae der I. Zentral-Afrika-Ex- 

pedition. Wiss. Ergebn. dtsch. Zentr. Afr. Exped. 
3: 393-418. 

. 1914. Afrikanische Braconiden des Koenigl. 

Zoologischen Museums in Berlin. Mitt. zool. Mus. 
Beriin 7: 153-230. 

Thomson, C. G. 1895. Lll. Bidrag till Braconidemas 
kannedom. Opusc. Entomol. 20: 2141-2339. 

Van Achterberg, C. 1983. A new species of /?/2v«- 
chosteres Fischer from Zaire (Hymenoptera, Bra- 
conidae). Zool. Meded. 57: 91-95. 

Van den Bosch, R. and F. H. Haramoto. 1951. Opius 
oophilus Fullaway, an egg-larval parasite of the 
Oriental Fruit Fly discovered in Hawaii. Proc. Haw. 
Entomol. Soc. 14: 251-255. 

Wesmael, C. 1835. Monographic des Braconides de 
Belgique. Nouv. Mem. Acad. R. Bruxelles 9: 1- 

. 1838. Monographic des Braconides de Bel- 
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Wharton, R. A. and F. E. Gilstrap. 1983. Key to and 
status of opiine braconid (Hymenoptera) parasit- 
oids used in biological control of Ceratitis and 
Dacus s. I. (Diptera: Tephritidae). Ann. Entomol. 
Soc. Am. 76: 721-742. 

Wharton, R. A. and P. M. Marsh. 1978. New World 
Opiinae (Hymenoptera: Braconidae) parasitic on 
Tephritidae (Diptera). J. Wash. Acad. Sci. 68: 147- 


89(1), 1987, pp. 74-76 




S. C. Harris 

Aquatic Biology Program, Department of Biology, University of Alabama, Tuscaloosa, 
Alabama 35487. 

Abstract.— \ new species, Agarodes alabamensis, from coastal Alabama is described 
and illustrated. The genus now totals ten species, all restricted to eastern North America. 

The genus Agarodes is restricted to and 
widespread throughout eastern North 
America from Canada to Florida (Wiggins, 
1977). Nine species are recognized in the 
genus, with all but Agarodes distinctus Ul- 
mer and Agarodes griseus Banks restricted 
to southeastern United States (Ross and 
Scott, 1974). In Alabama, four species oc- 
cur: Agarodes crassicornis (Walker) and 
Agarodes libalis Ross and Scott on the 
Coastal Plain; A. griseus Banks at scattered 
localities in northern Alabama; and Aga- 
rodes stannardi which is restricted to a small 
area in the northwestern portion of the state. 
To this list is added an apparently rare new 
species, Agarodes alabamensis, known only 
from the type locality on the Coastal Plain. 
As is typical for the genus, this new species 
was collected with a black-light along a small 
sand-bottom stream. 

Type material will be deposited at the Na- 
tional Museum of Natural History, Smith- 
sonian Institution. Terminology follows that 
of Schmid (1980) and Ross and Scott ( 1 974). 

Agarodes alabamensis Harris, 

New Species 

Fig. 1 

Diagnosis. — In many respects this species 
resembles Agarodes tetron (Ross) but differs 

' Contribution No. 93 from the Aquatic Biology Pro- 
gram, The University of Alabama. 

in the rounded apex of segment X and in 
the elongate mesal branch on the basomesal 
process of the inferior appendage. In addi- 
tion, A. alabamensis is a Coastal Plain in- 
habitant, while A. tetron occurs in the mon- 
tane regions of northern Georgia and North 

Male.— Length 17-19 mm. Body, legs and 
head light brown to yellow. Antennae brown, 
with 43 segments, the scape small and trap- 
ezoidal. Labial palpus 1.3 mm long; max- 
illary palpus 0.8 mm in length, oblong, bear- 
ing 2 narrow inner processes. Wings light 
brown; forewing with extensive peg-like se- 
tae in anal region and posteriorly along M 
and Cu veins. Abdominal segment IX an- 
nular. Segment X elongate and narrow in 
dorsal view, divided into lateral plates di- 
verging apically; in lateral view sinuate dor- 
sally and lightly sclerotized, distally round- 
ed to blunt apex, lacking ventral projection. 
Preanal appendage narrow and elongate, ex- 
tending nearly half length tergum X, setose 
distal portion curving dorsad in lateral view; 
in dorsal view somewhat spatulate distally 
and curving mesad. Inferior appendages 
slender anteriorly, widening posteriorly in 
lateral view; in ventral view united basally, 
slender with irregular margins, basomesal 
processes divided basally. Mesal processes 
over % length of inferior appendages, slen- 
der and narrowing to acute apex, distally 
serrate and diverging; lateral processes thin 



Agarodes alabamensis 

Fig. 1. Agarodes alabamensis n. sp., male genitalia. A, Lateral view. B, Dorsal 

view. C, Ventral view. 



and nearly equal in length to mesal pro- 
cesses. Phallus long and generally cylindri- 
cal, membranous apically with acute apex, 
ventral notch subapically. 

Female. — Unknown. 

Immatures. — Unknown. 

Etymology. — Latin: of Alabama. 

Holotype. — Alabama, Clarke County, 
Fishers Creek at unmarked Co. Rd., 1 mile 
SW Whatley (R3E T8N S25), 27 April 1 985, 
S. C. Harris. 

Paratype.— Alabama, same as above, 1 6. 

Disiribuhon.—Agarodes alabamensis is 
known only from the type locality and ap- 
pears to be rare in coastal Alabama. Exten- 
sive black-light trapping throughout Clarke 
County and adjacent counties has not yield- 
ed any additional specimens. Repeated 
black-light collecting along Fishers Creek at 
several locations in May 1985, early and 
late April 1986, and May 1986 found no 
additional material. As well, on four occa- 
sions an extensive search for Agawdes lar- 
vae in Fishers Creek and its tributaries was 

"Discussion.— Agawdes alabamensis is a 
member of the subgenus Agawdes Banks 
(Ross and Wallace, 1 974) based on the small 
antennal scape and slender mesal lobe of 
the maxillary palp. The key to Agawdes 
males in Ross and Scott (1974) is amended 
as follows to accommodate the new species. 

7. Mesal branch of basomesal process short, sit- 
uated apically, much as in Fig. 3 griseus 

Mesal branch of basomesal process long, sit- 
uated basally 7A 

7A. Mesal branch of basomesal process about half 
as long as lateral branch, apex of seqment X 
with ventral projection (Ross, 1948, Fig. 4) 


Mesal branch of basomesal process over half 
as long as lateral branch, apex of segment X 
without ventral projection alabamensis 


The Geological Survey of Alabama pro- 
vided equipment and facilities during the 
study and is gratefully acknowledged. John 
Unzicker of the Illinois Natural History 
Survey provided specimens o^ Agawdes te- 
twn for comparison. I thank also Patrick 
O'Neil for his help in the collections, Ruth 
Turner for photographing the figures, and 
Irene Thompson for typing the manuscript. 

Literature Cited 

Ross, H. H. and D. C. Scott. 1974. A review of the 
caddisfly genus Agawdes, with descriptions of new 
species (Trichoptera: Sericostomatidae). J. Ga. 
Entomol. Soc. 9: 147-155. 

Ross, H. H. and J. B. Wallace. 1974. The North 
American genera of the family Sericostomatidae 
(Trichoptera). J. Ga. Entomol. Soc. 9: 42-48. 

Schmid,F. 1980. Genera desTrichoptdresdu Canada 
et des Etats adjacents, pt. 7, 296 pp. In Les insectes 
et arachnides du Canada. Agric. Canada, Ottawa. 

Wiggins, G. B. 1977. Larvae of the North American 
caddisfly genera (Trichoptera). Univ. Toronto 
Press, Toronto. 40 1 pp. 


89(1), 1987, pp. 77-94 




James B. Woolley and Harold W. Browning 

(JBW) Department of Entomology, Texas A&M University, College Station, Texas 
77843; (HWB) Department of Entomology, Agricultural Research and Extension Center, 
Texas A&M University, 2415 East Highway 83, Weslaco, Texas 78596. 

Abstract.— JhQ literature on parasites of chaff scale, Parlatoha pergandii Comstock, is 
briefly reviewed with emphasis on chaff' scale in Texas. A survey of the natural enemies 
of chaff scale in Texas citrus showed two thelytokous and closely related (cryptic) species, 
Aphytis hispanicus (Mercet) and Aphytis comperei DeBach and Rosen, to be the most 
common parasites. Since these species are reported in the literature to be sympatric in 
many localities, and since individuals with an apparently intermediate morphology were 
found, we tested the hypothesis that the concepts of .4. hispanicus and A. comperei rep- 
resent two points in a continuous distribution of phenotypes. A morphometric study of 
the material was conducted to determine if two distinct morphs corresponding to A. 
hispanicus and A. comperei occur in Texas citrus, and if so, to identify useful morphological 
characters to distinguish between them. Sixteen measurements of anatomical structures, 
six meristic characters, and two qualitative characters were scored for 146 specimens 
reared from isolated chaff scale. The measurement data were analyzed using principal 
component and canonical variates analyses. Principal component analysis of the raw and 
log-transformed data showed that two distinct morphs exist which correspond to A. 
hispanicus and A. comperei. In addition, a third group of individuals, designated as A. 
^hispanicus, was found. These individuals are close to, but somewhat distinct from, A. 
hispanicus. Principal component analysis and canonical variates analysis suggest that the 
A. ^hispanicus group consisted of small specimens of A. hispanicus. Canonical variates 
analysis also showed that 6 of the 1 7 characters used were useful in discriminating between 
A. comperei and A. hispanicus. Two meristic characters showed strong discontinuities 
between A. comperei and A. hispanicus. We conclude that two species, A. comperei and 
A. hispanicus, are the common parasites of chaff scale in Texas citrus. 

This paper is the first in a series reporting in Texas (Dean, 1955; Dean et al., 1 983), 

the results of a survey of the parasites of Israel (Gerson, 1977; Harpaz, 1961), Spain 

armored scale on citrus in south Texas. Here (Limon et al., 1977; Carrero, 1 980), and in 

we discuss the status of uniparental (thely- other locations worldwide (Talhouk, 1975) 

tokous) Aphytis (Hymenoptera: Aphelini- Parlatoria pergandii occurs sympairicaWy in 

dae) reared from chaff scale, Parlatori a per- Israel with Parlatoria cinerea Doane and 

^an^/V Comstock (Homoptera: Diaspididae). Hadden, the tropical grey chaff scale (Ger- 

Chaff scale is a recurring problem on citrus son, 1967a, b). 



The most common parasites we encoun- 
tered on chaff scale were referable to one of 
two species in the proclia group ofAphytis: 
A. hispanicus (Mercet) and A. comperei 
DeBach and Rosen. Both species are uni- 
parental (Rosen and DeBach, 1979) and 
males referable to this species group were 
quite rare in our collections. The similarities 
between these two species and the occur- 
rence of apparent intermediate forms 
prompted this study. 

Prior to the work of DeBach and Rosen 
(summarized in Rosen and DeBach, 1979) 
species identification in Aphytis was ex- 
tremely difficult and often controversial. 
Dean (1965) expressed some frustration at 
the different species names various special- 
ists provided for Aphytis reared from chaff 
scale on Texas citrus— variously A. proclia 
(Walker), A. diaspidis Howard, and A. his- 
panicus. All of these species are in the pro- 
clia group, as are ^. maculicornis (Masi) and 
A. paramaculicornis DeBach and Rosen 
which were imported and released against 
the olive scale, Parlatoria oleae (Colvee), in 
California and elsewhere (Rosen and 
DeBach, 1979). Dean (1965), Dean and 
Hoelscher (1967) and Dean and Bailey 
(1960) refer to an Aphytis "complex" as the 
dominant parasites of chaff scale in Texas. 
Later, DeBach and Rosen (1976) deter- 
mined that Dean's chaff scale parasite ma- 
terial contained two very similar Aphytis 
species: A. hispanicus and a second species 
they described as A. comperei. 

Our examination of early correspondence 
and unpublished reports indicate that chaff 
scale was present early in the history of the 
citrus industry in Texas. A complete review 
of all parasites of chaff scale in Texas is 
forthcoming. Introduction of other Aphytis 
species have been made for chaff scale con- 
trol in Texas, but establishments of exotic 
species have not been documented (Dean 
and Bailey, 1960). In 1968 a strain of .4. 
paramaculicornis originally obtained from 
chaff scale on citrus at Escondido, Califor- 

nia was shipped from Riverside to Texas 
and approximately 35 adults were released 
on chaff scale at Weslaco. This somewhat 
enigmatic Aphytis was originally thought to 
be the "Iran" strain of .4. paramaculicornis, 
but Rosen and DeBach (1979) regard it as 
distinct and apparently indigenous to Cal- 
ifornia. No recoveries of this Aphytis have 
been made. In fact, we have not reared ma- 
terial from chaff scale in Texas referable to 
any introduced species. All Aphytis found 
on chaff scale in Texas can therefore be con- 
sidered to be indigenous, or exotic species 
which moved with chaff scale when it was 
introduced to Texas. 

The holotype o^ Aphytis hispanicus Mer- 
cet (1912) was from material reared from 
chaff scale on citrus at Valencia, Spain. The 
species was most recently redescribed by 
Rosen and DeBach (1979), whose hst of lo- 
calities for this species includes Spain, Italy, 
Turkey, Israel, the Caucasus, Morocco, Tai- 
wan, Brazil, Trinidad, Mexico, Rorida, Cal- 
ifornia, and notably, Texas. Most of the 
specimens were reportedly reared from P. 
pergandii, and to a less extent from P. ci- 
nerea, P. oleae, Aspidiotus nerii Bouche, In- 
sulaspis pallida (Green), Lopholeucaspis 
japonica (Cockerell), and Mytilaspis conch- 
iformis (Gmelin). Records from Acutaspis 
scutiformis (Cockerell), Aonidiella aurantii 
(Maskell) and Chrysomphalus dictyospermi 
(Morgan) were regarded as questionable. 
Aphytis hispanicus has also been reported 
attacking chaff scale in Morocco (Abassi, 
1975), Spain (Limonetal., 1976, 1977) and 
Rorida (Muma, 1971). Crouzel (1973) list- 
ed A. hispanicus and Aphytis argentinus 
Brethes as parasites of P. cinerea and P. 
pergandii, respectively, in Argentina. How- 
ever, Gerson (1977) stated that the chaff 
scale parasite which she listed as A. argen- 
tinus is probably a synonym of /I. comperei 
or A. hispanicus. Rosen and DeBach (1979) 
describe A. hispanicus as a uniparental, sol- 
itary parasite of Parlatoria species which 
attacks second instars, male scale, and adult 



Table 1. Diagnostic characters for Aphytis comperei and A. hispanicus from Rosen and DeBach (1979). 
Information taken from key to species and species redescriptions. 

.1. hispanicus 

Genal sutures 

Antennal club 

Pedicel, funicle, base of club 

Antennal club 


Ratio of lengths of ovipositor/ 
middle tibia 

Forewing, length/width 

Setae in delta region of fore- 

LMC forewing/width forewing 

heavily sclerotized, infuscate from 
oral margin to about % distance 
to eye 

length/width = 2.5-3.0 

uniformly and strongly infuscate 
apical 'A blackish (infuscate) 
6-8 per side, elongate, non-over- 
lapping, faintly infuscated 
less than or = 1.33 


59-161 in 9-12 rows 


less heavily sclerotized, faintly in- 

shorter, thicker, length/width about 


tip with conspicuous black spot 

3-5 per side, wider, distinctly black- 

longer, up to 2.00 


51-96 in 7-9 rows 


female scale, preferring the latter. Gerson 
(1967b, 1968), Rosen (1965, 1967, 1969) 
and Rivnay (1968) provide information on 
the biology of .4. hispanicus in Israel. 

DeBach and Rosen (1976) described A. 
comperei and provided diagnostic charac- 
ters to distinguish it from A. hispanicus and 
A. proclia. The holotype female was reared 
from "Aonidiella aurantii material" on cit- 
rus in McAllen, Texas; however, Rosen and 
DeBach (1979) regarded the California red 
scale record for the holotype as question- 
able. They point out that California red scale, 
chaff scale, and other scale species are often 
found mixed together on citrus and cross 
contamination of rearing samples is com- 
mon. Most of the records listed for .4. com- 
perei are from chaff scale, and records from 
A. aurantii, Chrysomphalus aonidum (L.), 
and Cornuaspis beckii (Newman) were re- 
garded as questionable (Rosen and DeBach, 
1979). The distribution oi A. comperei in- 
cludes Texas, Mexico, Florida, Jamaica, 
South Africa, Hong Kong and Canton, 
China. Little additional information is 
available on the biology oi A. comperei be- 
yond the observation of Rosen and DeBach 
(1979) that this species is uniparental. 

Rosen and DeBach (1979) provide a 
number of characters to distinguish com- 
perei and hispanicus in their key to Aphytis 
species and in the redescriptions of the two 
species. We list their criteria in Table 1 and 
provide figures of typical character states for 
each species. Of particular interest are the 
shape and coloration of the antennal seg- 
ments, the conformation of the crenulae, 
and aspects of the forewings. In the next few 
paragraphs, all discussion of the diagnostic 
characters for the two species refers to the 
criteria of Rosen and DeBach (1979). 

The antennal club oi A. hispanicus (Fig. 
1, Table 1) is characterized by a blackish, 
infuscate region in about the distal third. 
The antennal club of A. comperei (Fig. 2, 
Table 1 ) is shorter, thicker, and the infuscate 
area is confined to a black spot at the tip. 

The propodeum o{ Aphytis typically bears 
several posterior, lamellate projections 
called crenulae. Variations in the size, shape, 
number, and color of the crenulae have been 
used to distinguish between many Aphytis 
species (Rosen and DeBach, 1979). These 
authors state that 12-16 crenulae are found 
on A. hispanicus (Table 1), and further, that 
the crenulae in hispanicus are elongate, pale 






^^te^-«^- ^ 


X. c r 







Figs. 1-5. Photomicrographs of Aphytis species. I, Antennae of A. hispanicus, medial aspect. 2, Antennae 
of^. comperei, medial aspect. 3, Propodeum of .4. hispanicus. 4, Propodeum of A. comperei. 5, Eighth abdominal 
tergum of .4. hispanicus. (cr = crenulae, Is = lateral setae, ms = medial setae). 

in color, and do not overlap one another 
(Fig. 3). In contrast, A. comperei is char- 
acterized by 6-10 crenulae (Table 1), and 
the crenulae are wider, overlapping, and 
distinctly darker in color (Fig. 4). 

The forewing of A. hispanicus is longer 
and narrower than that of ^. comperei (Ta- 
ble 1 , Figs. 6 and 7), but the differences are 
not striking. The oblique, bare streak in the 

aphelinid wing is properly termed the linea 
calva (Hayat, 1983). In Aphytis the region 
of the forewing basal to the linea calva is 
called the delta region (Rosen and DeBach, 
1979). As noted in Table 1, specimens ofv4. 
hispanicus are said to commonly bear more 
setae (microtrichiae) in the delta region (59- 
161) than specimens oiA. comperei (5 1-96 
setae). Finally, the ratio of the longest mar- 



Figs. 6, 7. 6, Forewing of A. hispanicus. 7, Forewing of A. comperei. 

ginal cilia (LMC) on the forewing to the 
greatest width of the forewing in hispanicus 
specimens is usually longer (ratio of '/^-'/j) 
than in comperei specimens ('/j) (Table 1). 
Several of the characters in Table 1 are 
extremely subtle, and require some subjec- 
tive interpretation by the observer. The 
ranges given for several other characters are 
strongly overlapping (e.g. the number of se- 
tae in the delta region of the forewing). Fur- 
ther, some individuals in our material show 
an intermediate morphology, suggesting that 
Rosen and DeBach's (1979) concepts of A. 
comperei and A. hispanicus might represent 
two ends of a continuous distribution of 
phenotypes. The two species are often sym- 
patric (in the sense that both are often reared 

from the same collection ofParlatoria scale) 
which is consistent with this contention. As 
both species are uniparental, a purely phe- 
netic species concept is appropriate. Differ- 
ences in biology, behavior, or ecology, were 
they known, would support the recognition 
of two species. 

The objective of this study was to deter- 
mine if two distinct morphs, corresponding 
to A. comperei and A. hispanicus, occur in 
south Texas citrus, and if so, to determine 
by what morphological attributes they can 
best be distinguished. The null hypothesis, 
in a sense, was that the morphs correspond- 
ing to these species are not distinct and that 
they intergrade. We analyzed the variation 
in a set of morphological characters in our 



populations in order to resolve these alter- 


Parlatoria pergandii material was col- 
lected from 28 citrus groves in Hidalgo and 
Cameron counties in south Texas during 
March through November, 1983. Groves 
were generally unsprayed, and formed an 
east-west transect of the citrus production 
region in Texas. Individual parasitized P. 
pergandii were identified by lifting scale 
covers with a dissecting probe. Individuals 
were then isolated in 0.25 dram glass shell 
vials with cotton stoppers. The parasites 
were allowed to emerge and die in the vials 
and were then slide-mounted in Hoyer's 
medium (Rosen and DeBach, 1 979). A total 
of 146 slide-mounts of specimens referable 
to either A. comperei or hispanicus were 
available for study at the time the morpho- 
metric analyses were begun. 

Data were collected from all available 
specimens using a Zeiss compound micro- 
scope equipped with Nomarski contrast en- 
hancement. Measurements on specimens 
were taken through either a 16 x or 40x 
objective, using a 12.5 x eyepiece contain- 
ing a reticle with 100 divisions. The eye- 
piece reticle was calibrated with a stage mi- 
crometer allowing conversion of eyepiece 
reticle units to microns, the scale used for 
all quantitative measurements. 

At the time the data were taken, a ten- 
tative species determination was made for 
each specimen, using the criteria of Rosen 
and DeBach (1979) (Table 1). In seven cases, 
the character states of a specimen were out- 
side the stated range for either species, but 
close to one of them. Five individuals were 
assigned a tentative determination of A. 
Ihispanicus and two as A. '^comperei. 

The character set used was a mixture of 
quantitative (continuous) measurements 
(Table 2), meristic (counted), and coded 
multistate (qualitative) characters. The 
character set was assembled using various 
criteria. We re-examined the characters of 

Rosen and DeBach (1979), accounting for 
characters 1, 4, 5, and 9-15 in Table 2. We 
also included several additional measure- 
ments (characters 2, 3, 6-8, 16 and 17 in 
Table 2) so that the data would better de- 
scribe differences in shapes between speci- 

Rosen and DeBach (1979) stressed the 
value of the crenulae for species discrimi- 
nation. We coded data for the crenulae as: 
the number of crenulae (meristic), the color 
of the crenulae (dark, some dusky color, or 
pale), and the degree to which the crenulae 
were overlapping (overlapping, contiguous 
but not overlapping, or well separated). In 
addition, we counted the setae present on 
three abdominal terga: the seventh, lateral 
setae on the eighth, medial setae on the 
eighth, and on the syntergum (following Ro- 
sen and DeBach's (1979) numbering of ter- 
ga, in which the propodeum is counted as 
the first tergum). 

All morphometric analyses on the data 
set were performed using the Statistical 
Analysis System (SAS) software (SAS In- 
stitute, 1982a, b) on a VAX 1 1/750 micro- 
computer. Sixteen quantitative characters 
and one meristic character (the number of 
setae in the delta region of the forewing) 
were used for morphometric analyses (Ta- 
ble 2). 

Principal components analysis (PCA) was 
performed on the variance/covariance ma- 
trix computed from the raw data. In all 
multivariate statistical procedures, SAS 
programs remove any observations with 
missing data points for variables used in the 
analysis. Twenty observations had missing 
data for one or more quantitative charac- 
ters, leaving 126 observations available for 
the PCA. Principal components computed 
from a variance/covariance matrix may be 
sensitive to the greater variance associated 
with characters with numerically larger val- 
ues (Neff and Marcus, 1980). For this rea- 
son, PCA was also performed on the vari- 
ance/covariance matrix computed from the 
logarithms (base 10) of the raw data. 



Table 2. Univariate statistics for variables used in the morphometric analyses. Means, 95% confidence 
intervals around the means, and ranges are given in microns. Values are lengths, unless otherwise indicated. An 
asterisk (*) preceding the variable number indicates that the character was discussed by DeBach and Rosen 
(1976) and/or Rosen and DeBach (1979). 


.-1. compere! 

X ± 95% CI. n 


A. Inspamcus 

X ± 95% CI, n 


.-1. Ihispamcus 

X ± 95% CI, n 


*1) Scape 

96.0 ± 1.46, 100 

72.3 to 109.5 

97.8 ± 2.41.37 
78.8 to 111.7 

81.5 ±6.91,5 
72.3 to 92.0 

2) Pedicel 

37.2 ±0.57, 101 
28.5 to 43.8 

36.0 ± 1.58,38 
12.0 to 41.6 

31.1 ± 2.58,5 
26.3 to 32.8 

3) Apical funicle segment 

34.1 ±0.56, 102 
26.3 to 39.4 

33.4 ± 0.72, 37 
26.3 to 37.2 

25.8 ± 1.65,5 
24.1 to 28.5 

*4) Antennal club 

78.8 ± 1.31,98 
61.3 to 94.2 

82.3 ± 1.92,37 
67.9 to 96.4 

68.6 ± 1.92,5 

65.7 to 70.1 

*5) Infuscation on antennal club 

20.2 ± 0.64, 97 
13.1 to 28.5 

35.3 ± 1.35,37 
28.5 to 43.8 

28.9 ± 3.25, 5 
26.3 to 35.0 

6) Mesoscutum 

98.7 ± 1.76, 101 
74.5 to 116.1 

99.8 ± 2.36, 36 
85.4 to 118.3 

81.9 ±4.97,5 
76.6 to 89.8 

7) Scutellum 

80.9 ± 1.52, 102 
59.1 to 94.2 

83.6 ± 1.98,37 

65.7 to 92.0 

67.9 ± 7.79,5 
59.1 to 81.0 

8) Metanotum 

13.6 ±0.30, 103 
11.0 to 17.5 

15.0 ±0.51, 38 

13.1 to 17.5 

12.7 ± 1.65,5 
11.0 to 15.3 

*9) Propodeum 

50.3 ± 0.98, 103 
37.2 to 61.3 

58.1 ± 1.60,38 

48.2 to 65.7 

45.1 ± 7.62,5 
35.0 to 56.9 

*10) Ovipositor 

284.3 ±4.01, 103 

224.4 to 325.4 

261.3 ±4.35, 38 
235.6 to 291.7 

225.5 ± 15.79,5 

207.6 to 252.4 

* 1 1 ) Forewing (length) 

553.0 ±8.51, 102 

415.1 to 628.3 

572.2 ± 10.54, 37 
493.7 to 645.2 

471.2 ± 34.94,5 
432.0 to 527.3 

*12) Forewing (width) 

201.2 ±3.73, 101 
140.2 to 241.2 

192.7 ± 4.13,37 
162.7 to 218.8 

158.2 ± 12.62,5 
145.9 to 179.5 

*13) LMC on forewing 

38.7 ± 1.06, 102 
28.0 to 56.1 

50.6 ±2.13, 37 

33.7 to 61.7 

57.2 ± 8.33,5 
44.9 to 67.3 

*14) Setae in delta region, forewing 

67.4 ± 2.09, 100 
37 to 89 

99.5 ± 4.64, 37 
73 to 129 

62.6 ± 11.88,5 
50 to 80 

*15) Middle tibia 

152.3 ± 2.57, 101 
116.1 to 175.2 

163.4 ± 3.49,38 
138.0 to 179.6 

131.8 ± 11.99,5 
118.3 to 151.1 

16) Basitarsus 

51.0 ± 1.12, 103 
35.0 to 63.5 

56.7 ± 1.62,38 
46.0 to 67.9 

42.9 ± 6.94, 5 
35.0 to 54.8 

17) Apical spur, middle tibia 

50.4 ± 0.82, 103 
37.2 to 59.1 

56.4 ± 1.35,38 
46.0 to 65.7 

45.6 ± 3.25,5 
41.6 to 50.4 

Canonical variates analysis (CVA) was 
used to further evaluate particular variables 
for species discrimination. As discussed be- 
low, the original species determinations were 
supported (in most cases) by the PCA, 
therefore, these determinations were used 
as the class variable for CVA. We expected 
that the first canonical variate would be con- 

structed to optimally discriminate A. his- 
panicus and A. comperei given the presence 
of the third class of .4. Ihispanicus individ- 
uals. In addition, given the hypothesis that 
the A. Ihispanicus individuals were some- 
what distinct from A. hispanicus individu- 
als, we wished to examine which variables 
contribute to the difference. It was hoped 



• comperei 

■ ? comperei 

o hispaniciis 

° ? hispanicus 



i — V 


§0° t°\ 

oP '^o ° 

^ — V 

i h 


-3.0 -- 

ii II (14%) 

I (79%) 

Fig. 8. Observations plotted on the first two principal components computed from the co variance matrix 
fi-om the untransformed data. The first prinicpal component contains 79% of the sample variance, the second 
principal component contains 14%. 

that the second canonical variate would be 
constructed in a way that would provide 
discrimination between A. hispanicus and 
A. ^hispanicus individuals. 

Once the scores for the original obser- 
vations on the first two canonical variates 
were obtained, we constructed 95% predic- 
tion regions around the clusters of points 
for each class using the formulae of Owen 
and Chmielewski (1985), an application of 
standard methods (e.g. Johnson and Wich- 
em, 1982). These ellipses have their centers 
at the group mean for each cluster for each 
canonical variate and enclose a 95% pre- 
diction region in the following sense: if all 

such prediction regions were drawn, 95% of 
them would contain each sample point 
(Owen and Chmielewski, 1985). This tech- 
nique assumes that the scores for each class 
on the first two canonical variates have a 
bivariate normal distribution. We could not 
test for bivariate normality, but we did test 
for univariate normality, and in all cases 
except one, we could not reject the null hy- 
pothesis of univariate normality. The one 
exception was the distribution of scores for 
the first canonical variate for comperei. In- 
spection of the distribution of comperei 
points on the first two canonical variates 
revealed one conspicuous outlier. With this 



outlier removed the null hypothesis (uni- 
variate normality) could not be rejected 
(P > 0.15, Kolmogorov test). For this rea- 
son, the 95% confidence ellipse for comperei 
was constructed without the score for the 
outlier observation. 


Principal component analysis.— A pro- 
jection of the individual specimens on the 
first two principal components computed 
from the variance/covariance matrix from 
the original data is presented as Fig. 8. To- 
gether, the first two principal components 
account for 93.0% of the original variance. 
The remaining principal components rep- 
resent increasingly miniscule proportions of 
the total variance: from 2.1% for the third 
to 0.03% for the seventeenth. The unitized 
eigenvectors associated with the first two 
principal components are shown in Table 
3. The elements of each vector have been 
scaled so that the sum of the squares of all 
the elements in each vector is unity. Thus, 
the elements represent weights, and the val- 
ue for each element squared represents the 
proportion of variance in the principal com- 
ponent which each variable contributes, as- 
suming that PCA has produced uncorrect- 
ed linear transformations of the original 

Two distinct clusters of points represent- 
ing A. comperei and A. hispanicus individ- 
uals were found (Fig. 8). The individual ten- 
tatively determined as A. 1 comperei lies well 
within the cluster formed by A. comperei 
individuals. The clusters formed by A. com- 
perei and A. hispanicus are distinct with re- 
spect to the second principal component 
only. Thus, by examining the weights in Ta- 
ble 3 for the second principal component, 
one can gauge the contribution of individual 
variables to the location of individuals on 
this axis. For example, from Fig. 8 and Ta- 
ble 3 it is noted that A. hispanicus individ- 
uals tend to have longer and narrower fore- 
wings with long marginal cilia, a longer 

Table 3. Eigenvalues and weights for the first two 
principal components, computed from the covariance 
matrix from the untransformed data. The vectors are 
scaled so that the sum of the squares of the elements 
in each vector is unity. The rows have been sorted on 
the elements for the second principal component, from 
numerically highest to lowest. 

Vanable or Quantity 


PC 11 




Proportion of Variance 



14) Setae in delta region. 




5) Length of infuscate area 

on club 



13) Length of LMC on fore- 




1 1 ) Length of forewing 



15) Length of middle tibia 



9) Length of propodeum 



17) Length of midtibial spur 



16) Length of basitarsus 



4) Length of club 



8) Length of metanotum 



7) Length of scutellum 



1 ) Length of scape 



6) Length of mesoscutum 



3) Length of apical funicle 




2) Length of pedicel 



1 2) Width of forewing 



10) Length of ovipositor 



infuscate area on the antennal club and more 
setae in the delta region of the forewing. 
Aphytis comperei individuals tend to have 
wider forewings and longer ovipositors. 

The first principal component in Fig. 8 
fits some of the criteria (Blackith and Rey- 
ment, 1971; Jolicoeur and Mosimann, 1960) 
commonly used to identify a size vector, or 
a principal component which reflects pri- 
marily variation in overall size of the spec- 
imens. The weights for the first principal 
component in Table 3 are all positive except 
for character 13. The weights are not, how- 
ever, of uniform magnitude. The heavily 
weighted characters (14, 11, 15, 12, and 10) 
are measurements of comparatively large 
structures with correspondingly large vari- 



• comperei 

■ ? comperei 

o hispaniciis 

° ? hispaniciis 



I 1 \ h 


□ D 

3.0 -' 

I f- 


□ qO o 
S o 

% o 

^ — I 


o ooo 

-r o 
o o 

A II (28%) 

I (54%) 

Fig. 9. Observations plotted on the first two principal components computed from the covariance matrix 
from the log-transformed data. The first principal component contains 54% of the sample variance, the second 
principal component contains 28%. 

ance. We note also that the clusters of points 
for each species in Fig. 8 are roughly parallel 
to the first principal component axis, lend- 
ing further support to the interpretation of 
this principal component as a size vector. 
If the first principal component simply ap- 
proximates overall size, then the five indi- 
viduals originally determined as A. Ihispan- 
icus would appear to be small specimens of 
A. hispanicus. 

Figure 9 is a projection of the individuals 
on the first two principal components com- 
puted from a variance/covariance matrix 

derived from the log-transformed data. This 
transformation should reduce the overall ef- 
fect of the large variances associated with 
variables with numerically large values. Ta- 
ble 4 shows the eigenvalues and eigenvec- 
tors for the first two principal components, 
as discussed above for Table 3. That the 
tranformation was successful can be seen in 
Table 4, in that the first principal compo- 
nent now accounts for only 53.8% of the 
total variance, with a greater proportion, 
27.7%, now contained in the second prin- 
cipal component. The remaining principal 



components now account for somewhat 
greater, but still relatively small proportions 
of the total variance: 4.7% for the third and 
0.1% for the seventeenth. The weights for 
the first principal component (Table 4) are 
all positive, although not of uniform mag- 
nitude, suggesting that the first principal 
component retains some variance associ- 
ated with overall size of the specimens. 

Again, two distinct clusters of points in 
Fig. 9 correspond with original determina- 
tions of either A. comperei or A. hispanicus 
and A. '^hispanicus. However, the clusters 
in Fig. 9 are at oblique angles to two prin- 
cipal component axes, suggesting that now 
the first principal component expresses 
variance associated with shape differences 
in addition to size differences. Also, the in- 
clusion of a point in one or another cluster 
is now determined by a contribution from 
both principal components. The contribu- 
tion of individual variables to the clustering 
of individuals when projected on the first 
two principal components can be assessed 
by examining the relative weights of vari- 
ables on each component. The length of the 
infuscate portion of the antennal club is the 
only variable which makes a strong contri- 
bution to both the first and second principal 
component, accounting for just over 25% of 
the variance of each. The number of setae 
in the delta region of the forewing accounts 
for slightly over 25% of the variance in the 
first principal component, and the length of 
the marginal cilia on the forewing for slight- 
ly over 25% of the variance represented by 
the second principal component. Other 
variables with high weights on the first prin- 
cipal component are the length of the pro- 
podeum and the length of the basitarsus. 
The width of the forewing, length of the 
pedicel and third funicle segment on the 
antenna, lengths of the mesoscutum and 
scutellum, and length of the ovipositor all 
have relatively high weights on the second 
principal component. 

The A. '^hispanicus individuals are again 
concentrated at one end of the distribution 

Table 4. Eigenvalues and weights for the first two 
principal components, computed from the covariance 
matrix from the log-transformed data. The vectors are 
scaled so that the sum of the squares of the elements 
in each vector is unity. 

Vanable or QuanlUy PC I PC II 

Proportion of Variance 

1 ) Length of scape 

2) Length of pedicel 

3) Length of apical funicle 

4) Length of club 

5) Length of infuscate area 
on club 

6) Length of mesoscutum 

7) Length of scutellum 

8) Length of metanotum 

9) Length of propodeum 

10) Length of ovipositor 

1 1 ) Length of forewing 

1 2) Width of forewing 

13) Length of LMC on fore- 

14) Setae in delta region, 

15) Length of middle tibia 

1 6) Length of basitarsus 

17) Length of midtibial spur 

of y4. hispanicus individuals in Fig. 9. How- 
ever, the effect now appears to be spread 
between the first and second principal com- 
ponents. Furthermore, the points now over- 
lap with A. hispanicus more with respect to 
their location on the second principal com- 
ponent. In this case, more than a simple size 
effect seems to be involved, as the location 
of points with respect to either component 
is not simply size related. It appears that 
these individuals are intermediate with re- 
spect to some aspects of morphology, al- 
though more similar to A. hispanicus than 
to A. comperei. Aphytis '^hispanicus was re- 
tained as a distinct a priori class in the ca- 
nonical variates analysis for this reason. The 
A.'? comperei specimen was treated as a 
member of the A. comperei class for canon- 
ical variates analysis because the point rep- 
resenting this specimen fell in the middle of 









































• comperei 
o hispanicus 
° ? hispanicus 


+ 10 


-10 -L 

Fig. 10. Observations plotted on the two canonical variate axes. The ellipses around each cluster of points 
represent 95% prediction regions (Owen and Chmielewski, 1985). 

the points for the A. comperei specimens 
when projected against either set of prin- 
cipal components (Figs. 8 and 9). 

Canonical variates analysis. — Homoge- 
neity of the CO variance matrices for the three 
classes used in canonical variates analysis 
was tested using the SAS DISCRIM pro- 
cedure. The covariance matrix for the Ihis- 
panicus class was not of full rank due to the 
small number of observations, and there- 
fore could not be tested against the other 
two. However, the tests did show that the 
null hypothesis that the covariance matrices 
of the comperei and hispanicus classes were 

homogeneous could not be rejected {P = 
1.000, likelihood ratio test). We assumed 
that the covariance matrix for the Ihispan- 
icus class was also homogeneous with the 
other two, since we had no reason to assume 

Figure 10 is a plot of the projection of the 
individuals on the first two canonical vari- 
ates. The first canonical variate contains 
97.6% of the between-groups variance, and 
since with three groups only two canonical 
variates can be constructed, the second ca- 
nonical variate contains 2.4% of the be- 
tween-groups variance. One would expect 



Table 5. Standardized coefficients and total canonical structure for the canonical variates analysis. The 
standardized coefficients are the amount that the canonical variate score will change for a change in the original 
variable of one standard deviation. The total canonical structure values are the total-sample correlations between 
the original variables and the canonical structure scores. The rows have been sorted by the elements of the 
vector of coefficients for the first canonical variate. 

Standardized Coefficients 

Total Canonical Structure 



CV 11 

CV 1 

CV 11 

5) Length of club infuscation 





14) Setae in delta region, forewing 





9) Length of propodeum 





13) Length of LMC on forewing 





1 7) Length of midtibial spur 





1 5) Length of middle tibia 





1 1 ) Length of forewing 





8) Length of metanotum 





16) Length of basitarsus 





4) Length of antennal club 





2) Length of pedicel 





6) Length of mesoscutum 





1 ) Length of scape 





12) Width of forewing 





7) Length of scutellum 





3) Length apical funicle segment 





10) Length of ovipositor 





the very tight clusters of points representing 
the comperei and hispanicus individuals (Fig. 
10), because canonical variates are con- 
structed to maximize between-group co- 
variance relative to within-group covari- 
ance. The 95% confidence ellipse for the 
comperei specimens (with one outlier re- 
moved, as discussed above) is well separat- 
ed from the ellipses for both the hispanicus 
and Ihispanicus groups, while the ellipses 
for hispanicus and '? hispanicus are broadly 
overlapping. We note also that the Ihispan- 
icus ellipse differs in size, shape, and ori- 
entation from the other two ellipses, an in- 
dication that the covariance matrix for the 
this class may not be equal (Owen and 
Chmielewski, 1985). 

Clearly, the first canonical variate un- 
ambiguously discriminates the comperei 
group from the hispanicus plus Ihispanicus 
groups (Fig. 10). The standardized canoni- 
cal coefficients for the canonical variates 
(Table 5) are the products of the canonical 
vector coefficients and the pooled within- 
group standard deviations for each variable. 

They represent the amount that the canon- 
ical variate score (e.g. in Fig. 10) will change 
for each change of the original variable by 
one standard deviation. A large absolute 
value for a standardized coefficient gener- 
ally indicates a variable which will be useful 
in discrimination (but see Campbell and 
Atchley, 1981, for a discussion of potential 
problems with this interpretation). The 
strong positive standardized coefficient 
scores for the first canonical variate for vari- 
ables 5, 9, 13, and 14 in Table 5 indicate 
that hispanicus individuals tend to have a 
longer infuscate area on the antennal club, 
a longer propodeum, longer marginal cilia 
on the forewing, and more setae in the delta 
region of the forewing, respectively. The 
strong negative score for variable 1 on the 
first canonical variate indicates that com- 
perei individuals have longer ovipositors 
because the comperei cluster in Fig. 1 is in 
the negative range of the first canonical vari- 
ate. Weaker negative scores on the first ca- 
nonical variate in Table 5 for characters 7 
and 3 indicate that comperei individuals tend 



40 T 



specimens 20 









9 10 11 12 

number of crenulae 

13 14 15 

Fig. 1 1. Number of specimens with observed number of crenulae. (Black bars represent A. comperei, white 
bars represent A. hispaniciis, shaded bars represent A. Ihispanicus specimens.) 

to have a longer scutellum and a longer api- 
cal funicle segment. 

The same pattern is evident in the total 
canonical structure values in Table 5. These 
values represent the total-sample correla- 
tions between the original variables and the 
canonical structure scores. As with the stan- 
dardized coefficients, variables 5,9, 13, and 
14 have strong positive correlations on the 
first canonical variate. Variable 17, the 
length of the midtibial spur, also shows a 
strong positive correlation on the first ca- 
nonical variate, but its standardized coef- 
ficient is relatively low. Variable 10, the 
length of the ovipositor, has a strong neg- 
ative correlation on the first canonical vari- 
ate, with the same implication as discussed 

Meristic characters.— At least two of the 
meristic characters for which we recorded 
data show marked differences between these 
species. As noted by Rosen and DeBach 
(1979), hispanicus individuals tend to have 
more crenulae that comperei individuals. 
Figure 1 1 is a histogram of the number of 

specimens observed with a particular num- 
ber of crenulae. Most comperei specimens 
had 8-10 crenulae (see Fig. 4), most his- 
panicus specimens had 11-15 (see Fig. 3), 
while the Ihispanicus specimens had an in- 
termediate number. Another useful meristic 
character is the number of medial setae on 
the eighth abdominal tergum (Fig. 5). As 
can be seen in Fig. 1 2, most comperei spec- 
imens had two such setae, rarely 1 or 3, 
while most hispanicus specimens had 4, or 
5, rarely 3 or 6. The Ihispanicus specimens 
were again intermediate with 2 or 3 setae 
in this location. 


The ^hispanicus individuals do not ap- 
pear to represent a morph distinct from the 
hispanicus individuals. In all plots (Figs. 8, 
9, 10) the '^hispanicus observations cluster 
at one end or the other of the distributions 
of hispanicus observations. In fact, the 1 his- 
panicus specimens appear to be simply small 
hispanicus individuals. This can be seen 
clearly in Fig. 10 and Table 5. The Ihispan- 




uu ■ 





















1 1 U , c=I , 




2 3 4 5 

medial setae on eighth tergum 

Fig. 12. Number of specimens with observed number of medial setae on the eighth tergum. (Black bars 
represent A. comperei. white bars represent A. hispanicus. shaded bars represent A. Ihispanicus specimens.) 

icus individuals He below the hispanicus 
cluster in Fig. 10, with virtually all discrim- 
ination between these two groups on the 
second canonical variate. In Table 5, the 
total canonical structure values show strong 
positive correlations between the scores on 
the second canonical variate and the origi- 
nal variables for all variables except 5 and 
1 3. Therefore, since 1 hispanicus individuals 
tend to have numerically lower (more neg- 
ative) scores on the second canonical vari- 
ate, they tend to have numerically smaller 
values for all variables except 5 and 1 3. This 
trend is also apparent in Table 2, in which 
the means for "^hispanicus individuals for all 
variables except 1 3 are lower than the means 
for hispanicus individuals. The meristic 
characters are intermediate for the Ihispan- 
icus class (Figs. 1 1 and 12), but again, if the 
Ihispanicus group consists of small hispan- 
icus individuals, they would be expected to 
have fewer crenulae and fewer medial setae 
on the eighth tergum. 

The comperei individuals (except the one 
outlier) are morphologically distinct from 

the hispanicus specimens (including the 
Ihispanicus specimens). In all of the results, 
the following patterns are consistent. Aphy- 
tis comperei individuals tend to have a long- 
er ovipositor than hispanicus individuals. 
Aphytis hispanicus individuals tend to have 
a longer infuscate area on the antennal club, 
more setae in the delta region of the fore- 
wing, longer marginal cilia on the forewing, 
and a longer propodeum than comperei in- 
dividuals, and to some extent, a longer mid- 
dle tibia with a longer apical spur. Of the 
diagnostic characters used by DeBach and 
Rosen (1976) and Rosen and DeBach (1979) 
(Table 1), most are well supported by our 
results. However, we did not find A. com- 
perei individuals to have consistently short- 
er clubs (Tables 3-5) as they stated, nor did 
we find consistently shorter or wider fore- 
wings in A. comperei (Tables 3-5). 

In Table 2 we have tabulated the means, 
95% confidence intervals for the means, and 
observed ranges for the quantitative vari- 
ables. The means for characters 5, 10, 12, 
13, and 14 for comperei and hispanicus are 



well separated (as indicated by the confi- 
dence intervals), but the ranges for these 
variables are strongly overlapping, with the 
exception of character 5 for which the ranges 
are contiguous. Therefore, while there is in- 
formation in many of these characters, any 
particular variable for a single specimen will 
not necessarily be discriminating. In fact, 
the covariation of traits leads to the dis- 
tinctly different morphologies of comperei 
and hispanicus. This points out the dangers 
in comparison of mean values for characters 
(e.g. with /-tests) or of the ranges of char- 
acters in making taxonomic decisions. In 
this case, comparison of means only would 
overstate the differences between comperei 
and hispanicus, while comparison of ranges 
only would not reveal trends which do occur 
in these data. Multivariate techniques such 
as PCA and CVA explicitly represent the 
covariation between many characters, 
thereby providing a method to assess trends 
which occur in several characters simulta- 
neously. For an excellent discussion of this 
general problem, see Albrecht (1980). 


Our results strongly support the conclu- 
sions of Rosen and DeBach (1979) that A. 
comperei and A. hispanicus are two distinct, 
but closely related species. Of course, with 
thelytokous forms such as these, notions of 
reproductive isolation do not apply, and the 
boundaries of species are necessarily arbi- 
trary. We have found rare male specimens 
in our material referable to one of the two 
species. The results of Rossler and DeBach 
(1972, 1973) suggest that the rare males in 
Aphytis species are functional, at least in 
some species. Therefore, the possibility ex- 
ists that mating occurs in the field, and that 
gene flow occurs between clones and even 
between species as we now recognize them. 
More likely, however, is the situation ob- 
served with uniparental strains of A. ma- 
culicornis and biparental strains of /I. par- 
amaculicornis in which laboratory studies 
have indicated that these forms are com- 

pletely reproductively isolated (Rosen and 
DeBach, 1979). The possible role of males 
in comperei and/or hispanicus is a matter 
for further investigation. 

Nevertheless, we have found two distinct 
morphs corresponding almost exactly to the 
concepts of .4. comperei and A. hispanicus 
presented by DeBach and Rosen ( 1976). The 
results from the principal components anal- 
ysis are the strongest evidence for the dis- 
tinctness of the two forms, as this technique 
does not utilize any a priori grouping cri- 
teria in dimension reduction. The meristic 
characters provide further evidence for the 
discontinuity in the morphologies of the two 
species. The A. 1 hispanicus specimens re- 
main somewhat problematic, as both the 
meristic characters and the multivariate 
analyses indicate that these individuals are 
intermediate in form. However, the multi- 
variate analyses also suggest that these spec- 
imens are smaller /i. hispanicus individuals 
and meristic data are consistent with that 
hypothesis. We do not, therefore, support 
recognizing these as a third, distinct morph. 

Given that A. comperei and A. hispanicus 
are morphologically distinct, how can one 
best identify individual specimens? The cri- 
teria of Rosen and DeBach (1979) (Table 1) 
will generally be useful, although we would 
not recommend using the length/width ra- 
tios of either the antennal clubs or the fore- 
wings. One new meristic character, the 
number of medial setae on the eighth ter- 
gum will be useful in most, but not all cases. 
However, as noted above, it is the covaria- 
tion in traits which makes the two species 
morphologically different, and no single 
characteristic will provide a reliable crite- 
rion for identification in all cases. Discrim- 
inant analysis provides a statistical method 
to identify unknowns to known groups in 
just such a situation. Discriminant analysis 
produces a single linear transformation of 
the original variables to optimally discrim- 
inate between a set of predefined groups. 
Canonical variates analysis, which we have 
used, is essentially the multi-group exten- 



sion of discriminant analysis (Albrecht, 
1980), and the special case of CVA of two 
groups is equivalent to discriminant anal- 
ysis (Neff and Marcus, 1980). Once a set of 
diagnostic characters and a discriminant 
function has been developed for a pair (or 
group) of cryptic species, the information 
can be distributed to persons who need to 
make routine identifications, but who do 
not have extensive experience with the 
species and with their subtle diagnostic 
characters. This person could input a series 
of measurements for each unknown speci- 
men. The discriminant function would then 
provide the a posteriori probability of an 
unknown belonging to one of the known 
species. However, one important caveat ex- 
ists, all possible species to which the un- 
known might be referable must be included 
in the discriminant function for the tech- 
nique to be valid. 


We thank Penny Wilkinson for slide- 
mounting the specimens used in this study, 
and for much of the data entry on the com- 
puter. We thank also Willy Melton for his 
assistance with the field work and with many 
other matters. Fred Hendricks, John Her- 
aty, William Smith, and Tom Unruh re- 
viewed the manuscript and each made many 
valuable suggestions and comments. This 
paper is Technical Article No. 2 1 84 1 of the 
Texas Agricultural Experiment Station. The 
specimens discussed above are deposited as 
Voucher Series No. 902 in the Insect Col- 
lection, Department of Entomology, Texas 
A&M University. 

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89(1), 1987, pp. 95-99 


R. D. Waltz and W. P. McCafferty 
Purdue University, Department of Entomology, West Lafayette, Indiana 47907. 

Abstract.— ThvQe new genera of Ephemeroptera are described for Afrotropical species 
of Baetidae with highly distinctive and unusual larvae. They are Pseudopannota Waltz 
and McCafferty, n. gen., including Pseudopannota bertmndi (Demoulin), n. comb, (type 
species) and Pseudopannota vinckei (Demoulin), n. comb.; Ophelmatostoma Waltz and 
McCafferty, n. gen., including Ophelmatostoma kimminsi Waltz and McCafferty, n. sp. 
(= Pseudocloeon sp. A. Kimmins); and Acanthiops Waltz and McCafferty, n. gen., including 
Acanthiops marlieri (Demoulin), n. comb. Pseudopannota and Ophelmatostoma have 
highly modified filtering/sweeping mouthparts. Pseudopannota also has fused mesotho- 
racic wingpads. Acanthiops is one of several genera of baetids with prominent dorsal 

A largely undescribed fauna of mayflies 
in the Afrotropics provides a rich source of 
interesting and phyletically important 
species. We assign the species treated herein 
to three new genera as a part of our con- 
tinuing study of baetid systematics. All three 
of the new genera are striking as larvae be- 
cause of variously exaggerated morphology 
not typically found among Baetidae. 

Pseudopannota Waltz and McCafferty, 

New Genus 

Larva. — Labrum (Demoulin, 1967, Fig. 
2a; 1973, Fig. 3b) notched medially on an- 
terior margin and with labral shelf Man- 
dibles (Demoulin, 1967, Fig. 2b, c; 1973, 
Fig. 3c, d) with incisors fused to apex; bases 
of prosthecae appearing recessed into man- 
dibular margin. Thumb of left mandible 
broad based and slightly elevated above 
plane of incisor bases. Maxillae (Demoulin, 
1967, Fig. 2d; 1973, Fig. 3e) with elongate 
three-segmented palps that exceed length of 
galealacinia; segment 3 of palp much elon- 
gated and swollen relative to two basal seg- 

ments; galealacinia with several apical den- 
ticles and adjoining row of bristles as in 
most baetids. Labium (Demoulin, 1967, Fig. 
2f; 1973, Fig. 3g) with glossae and para- 
glossae subtruncate apically, subparallel, and 
with long, fine setae apically; paraglossae 
exceed glossae in length; palps two seg- 
mented and with second segment swollen 
or expanded laterally and bearing long, fine 

Legs with or without row of long, fine 
setae on foretibiae and foretarsi (Demoulin, 
1967, Fig. la; 1973, Fig. 3h). Ventral fem- 
oral patch absent. Claws (Demoulin, 1967, 
Fig. 2g; 1973, Fig. 3i) with numerically re- 
duced but prominent denticles and without 
subapical bristles. Wingpads of mesothorax 
fused medially and nearly to apices (Fig. 1 
and Demoulin, 1967, Fig. la; 1973, Fig. 3a). 

Abdominal terga with scales and fine se- 
tae. Gills broadly rounded, thickened along 
anterior margin, with distinctly spiculate 
surfaces and spinous margins (Demoulin, 
1967, Fig. lb; 1973, Fig. 3n). Median ter- 
minal filament subequal to cerci. 



Fig. I. Pseudopannota bertrandi larva, mesotho- 

Type spQciQS.—Pseudocloeon bertrandi 
Demoulin, 1967: 227 [= Pseudopannota 
bertrandi (Demoulin), New Combination]. 

Included species.— Pseudopannota ber- 
trandi (Demoulin) and Pseudopannota 
vinckei (Demoulin) n. comb. 

Distribution. — Ivory Coast, Madagascar, 
Senegal (River Gambia). 

Etymology. — Feminine Latin: pseudo — 
false, pan — fused, and nota— back, in ref- 
erence to the fused back condition of this 
schistonote mayfly. It thus appears falsely 
to be a member of the mayfly suborder Pan- 
nota sensu McCafferty and Edmunds ( 1 979). 

Remarks.— Adults are unknown present- 
ly; however, the larvae of Pseudopannota 
are distinct from all other described Baeti- 
dae on the basis of the medially fused meso- 
thoracic wingpads, unique mouthparts, and 
uniquely denticulate claws. The unnamed 
larva described by Crass ( 1 947: 64) as Pseu- 
docloeon sp. appears to approximate very 
closely mouthpart, claw, and gill characters 
described for known species of Pseudopan- 
nota. Although we suspect that this species 
will prove to be a member of Pseudopan- 

nota, we must wait to confirm this because 
we have thus far been unable to study any 
of Crass's (1947) material. 

Ophelmatostoma Waltz and McCafferty, 
New Genus 

Larva. — Labrum (Kimmins, 1955, Fig. 
3b) notched medially on anterior margin 
and with labral shelf. Mandibles (Kimmins, 
1955, Fig. 3c, d) with incisors fused to apex; 
prosthecae prominent and bases appearing 
recessed into mandibular margin; spiculae 
present between bases of incisors and molar 
area. Thumb of left mandible broad based, 
curved, and slightly elevated above plane of 
incisor bases. Maxillae (Kimmins, 1955, Fig. 
3f) with slender two-segmented palps and 
subequal in length to galealacinia; galeala- 
cinia with prominent falcate denticle api- 
cally in addition to two adjoining rows of 
bristles apically. Labium (Kimmins, 1955, 
Fig. 3g) with glossae and paraglossae greatly 
elongated, slender, and apically with long, 
fine setae; paraglossae subequal to glossae 
in length; palps apparently two segmented 
(segments 2 and 3 fused) and swollen api- 
cally, basal segment abruptly angled toward 

Legs with specialized ventral row of long, 
stout bristles on forelegs (Kimmins, 1955, 
Fig. 3h). Ventral femoral patch absent. Claws 
with single row of denticles and without 
subapical bristles. 

Abdominal terga with scales and fine se- 
tae. Gills broadly rounded and with serra- 
tions on distal anterior margins. Median 
terminal filament subequal to cerci. 

Type species. — Ophelmatostoma kim- 
minsi Waltz and McCafferty n. sp. 

Distribution. — Malawi [= Nyasaland 
(Kimmins, 1955)], South Africa (Trans- 
vaal), Zimbabwe [= Southern Rhodesia 
(Agnew, 1963)], and Senegal (River Gam- 

Etymology. — Neuter Greek: ophelma- 
to— broom, stoma — mowXh, in reference to 
the distinctive filtering/sweeping-type 



Remarks.— Adults of Ophelmatostoma 
are presently unknown. This genus, viz. the 
larvae, is clearly distinct from all other gen- 
era of Baetidae on the basis of the mouth- 
part characters (Fig. 2) described above. The 
described genera of baetids known to have 
filtering/sweeping mouthparts, i.e. Guaji- 
rolis Flowers (1985) and now Pseudopan- 
nota, and Ophelmatostoma, have several 
similarly adapted but independently de- 
rived characters of the mouthparts. These 
include the profusion of long bristles and 
fine setae, prosthecae that appear to be re- 
cessed in the mandibular margin, and very 
long, thin apical denticles of the galealaci- 
nia. Specific character states of the mouth- 
parts, tergum, claws, gills, and available 
knowledge of the adult stages indicate that 
these genera represent independently de- 
rived lineages. 

Fig. 2. Ophelmatostoma kimminsi larva, head cap- 
sule (lateral left side). 

Ophelmatostoma kimminsi 
Waltz and McCaflFerty, New Species 

Pseudocloeon sp. A, Kimmins, 1955: 866. 

Larva.— Characterized as per Kimmins 
(1955) and as supplemented by the above 
generic description. 

Type materials. — Holotype, larva, in al- 
cohol, Malawi (=Nyasaland), Tengadzi 
Stream, 22.viii.1952, L. Bemer, deposited 
British Museum (Natural History), London. 
Paratypes: 4 larvae; 3 paratypes in alcohol 
and 1 paratype slidemounted in euparal 
(solvent: absolute alcohol), Senegal (River 
Gambia), Kedougau, 1 98 1 , M. T. Gil- 
lies, deposited Purdue University Ento- 
mological Research Collection, West Lafa- 
yette, Indiana, USA. 

Etymology. — The specific epithet is for D. 
E. Kimmins, the British ephemeropterist 
who first studied the species. 

Acanthiops Waltz and McCafferty, 
New Genus 

Larva. — Labrum (Demoulin, 1967, Fig. 
4a) notched medially on anterior margin and 
with labral shelf Mandibles (Demoulin, 

1967, Fig. 4b, c) with patch of setae between 
base of united incisors and molar region. 
Left mandible with incisors fused to apex; 
prostheca stouter than prostheca of right 
mandible and not appearing recessed into 
mandibular margin; thumb of molar region 
stout and distinctly elevated above plane of 
incisor bases. Right mandibular incisors 
apically separated. Maxillae (Demoulin, 
1967, Fig. 4d) with palps two segmented 
and subequal to or less than length of gal- 
ealacinia. Labium (Demoulin, 1967, Fig. 4f) 
with glossae and paraglossae slightly ta- 
pered apically; paraglossae subequal to glos- 
sae in length; palps three segmented; lateral 
margins of segment 2 subparallel; segment 
3 short and narrower at base than apex of 
segment 2. 

Legs elongate and without highly modi- 
fied setal areas (Demoulin, 1967, Fig. 4a). 
Ventral femoral patch absent. Claws (De- 
moulin, 1967, Fig. 4g) with two rows of 
denticles and one pair of subapical ante- 
riorly directed bristles. Prothorax and me- 
sothorax with more or less prominent dorsal 
tubercles (Demoulin, 1967, Fig. 3a, b). 

Abdominal terga with scales and fine se- 



tae in addition to more or less prominent 
dorsal, medioposterior tubercles (Demou- 
lin, 1 967, Fig. 3a, b; 1 964: 288). Gills asym- 
metric (Demoulin, 1967, Fig. 4i, 1-p), with 
spinous margins anteriorly, and numerous 
fine tracheae. Median terminal filament 
greatly reduced relative to cerci. 

Type species. — Centroptilutn marlieri 
Demoulin, 1967: 230 [= Acanthiops mar- 
lieri (Demoulin), New Combination]. 

Distribution.— Zaire [= Belgian Congo], 

Etymology. — Masculine Greek: akan- 
thos—Xhom, iops—sm2i\\ fish, an allusion to 
this genus being a tuberculate or spiny min- 
now-like mayfly. 

Remarks.— The abdominal tubercles of 
Acanthiops are very highly developed, more 
so than in other baetids. The few other tu- 
bercled baetid genera, e.g. Jubabaetis Miil- 
ler-Liebenau (1980), Neobaetiella Muller- 
Liebenau (1985), and Baetodes Needham 
and Murphy (1924), are clearly separable 
from Acanthiops on the basis of mouthpart, 
tergal, gill, and claw characters. Although 
adults of Acanthiops are unknown at the 
present time, Demoulin (1964) provided il- 
lustrations of a subimago associated with 
the larva he described as Centroptilum No. 
3 from Kenya, a species which is probably 
congeneric with A. marlieri and differing 
from this latter species by lacking gill 1 . The 
hindwing of this subimago possesses a bi- 
furcate median costal process. This peculiar 
character state is thus far only associated 
with certain Afrotropical baetids, i.e. Cen- 
troptiliodes Lestage, and some Centropti- 
lum. We suspect several Afrotropical 
species, which are clearly provisional with 
respect to their present generic placement, 
will eventually prove to belong to Acan- 
thiops. These provisional species, whose lar- 
vae remain unknown at the present time, 
include Centroptilum biarcuatum Kolpelke 
(1980), C. boettgeri Kolpelke (1980), C. di- 
centrum Demoulin (1956), C montanum 
Kimmins (1960), and C. sudafricanum Les- 
tage (1924). 


We thank P. C. Barnard, British Museum 
(Natural History), London; P. Grootaert, 
Institut Royal des Sciences Naturelles de 
Belgique, Brussels; and H. Andre, Musee 
Royale de I'Afrique Centrale, Tervuren, for 
loan of material. We also thank M. T. Gil- 
lies, Sussex, England, for the gift of speci- 
mens now deposited in PERC, and A. V. 
Provonsha for illustrations. This paper has 
been assigned Purdue University Experi- 
ment Station Journal Number 10,676. 

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. 1967. Description de deux larves atypiques 

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R. Entomol. Belgique 103: 226-232. 

. 1973. Ephemeropteres de Madagascar. III. 

Bull. Inst. R. Sci. Nat. Belgique, Entomol. 49: 1- 

Flowers, R. W. 1 985. Guajirolis, a new genus of Neo- 
tropical Baetidae (Ephemeroptera). Stud. Neotr. 
Fauna Envim. 20: 27-31. 

Kimmins, D. E. 1955. Ephemeroptera from Nyasa- 
land, with descriptions of three new species and 
some interesting nymphal forms. Ann. Mag. Nat. 
Hist., Ser. 12 8: 859-880. 

. 1960. Notes on East African Ephemeroptera, 

with descriptions of new species. Bull. Brit. Mus. 
Nat. Hist. (Entomology) II: 23-34. 

Kolpelke, J-P. 1980. Ephemeroptera aus der Emer- 
gen/ des zentralafrikanischen Berbaches Kalengo 
(Zaire). Teil 1 : Baetidae. Entomol. Abh. Staat. Mus. 
f Tierk. Dresden 43: 99-129. 

Lestage, J. A. 1924. Les Ephemeres de I'Afrique du 
Sud. Catalogue critique et systematique des es- 
peces connues et description de trois genres nou- 
veaux et de sept especes nouvelles. Revue Zool. 
Africaine 12: 316-351. 

McCafferty, W. P. and G. F. Edmunds, Jr. 1979. The 
higher classification of the Ephemeroptera and its 



evolutionary basis. Ann. Entomol. Soc. Am. 72: 
Miiller-Liebenau, I. 1980. Jubabaetis gen. n. and 
Platybaetis gen. n., two new genera of the family 
Baetidae from the Oriental Region, pp. 103-1 14. 
In J. F. Flannagan and K. E. Marshall eds.. Ad- 
vances in ephemeroptera biology. Plenum Pub- 
lishing Corporation, New York. 

. 1985. Baetidae from Taiwan with remarks 

on Baetiella Ueno, 1931 (Insecta, Ephemerop- 
tera). Arch, f Hydrobiol. 104: 93-110. 

Needham, J. G. and H. Murphy. 1924. Neotropical 
mayflies. Bull. Lloyd Library, 24, Entomol. Sen 
4: 1-79. 


Erratum: In the article by Wheeler, G. C. and J. Wheeler, 1986, Proc. Entomol. Soc. 
Wash. 88(4), p. 685, couplet 3b, please change ''separate" to "paper." 


89(1), 1987, pp. 100-102 




Charles W. Agnew 

Department of Entomology, Texas A&M University, College Station, Texas 77843- 

Abstract. —Acigona Hubner has been incorrectly used as a generic name in the Cram- 
binae due to an earlier type designation of a noctuid species. Acigona (sensu Bleszynski) 
is composed of genus level taxa which should not be considered congeneric. Friedlanderia 
n. name is proposed as a replacement name for Chiloides Bleszynski 1963, a homonym 
of Chiloides Butler 1881. The type, Tinea cicatricella Hubner, had been incorrectly des- 
ignated the type of Acigona by Bleszynski. Eoreuma morbidella (Dyar) n. comb, and 
Coniesta forsteri (Bleszynski) n. comb, are transferred from Acigona. 

Some unfortunate taxonomic errors have 
resulted in much confusion regarding the 
use of the generic name Acigona within the 
subfamily Crambinae (Lepidoptera: Pyral- 
idae). The incorrect usage of this name for 
crambine species is especially widespread in 
the non-systematic literature as the larvae 
of several species have been reported to in- 
fest agronomically-important Poaceae. Al- 
though many of the problems cannot be 
remedied until a revision o{ Acigona (sensu 
Bleszynski, 1967) is completed, some con- 
tributions toward nomenclatural stability 
can be made at this time. 

The genus Acigona was proposed by Hub- 
ner ([1826]: 342) for two species without a 
type species being designated. Hampson 
( 1 926) selected one of these, Phalaena man- 
to Cramer (Noctuidae), as the type of Aci- 
gona and treated the genus as a synonym of 
Euclystis Hubner. Bleszynski and Collins 
(1962), apparently unaware of this action, 
designated the other included species. Tinea 
cicatricella Hubner (Pyralidae) as the type 
of Acigona in the Crambinae. Bleszynski 
(1963) also designated T. cicatricella as the 

Xy^to{ Chiloides \mst\ 1949. Amsel(1949) 
originally included two species, T. cicatri- 
cella and Chilo hederalis Amsel in Chi- 
loides. Bleszynski (1963) decided Amsel had 
not properly designated a type species as 
required by The International Code of Zoo- 
logical Nomenclature (Article, 1 3b), which 
made the generic name unavailable. Al- 
though Amsel appeared to indicate T. ci- 
catricella was to be the type, the wording is 
somewhat ambiguous and probably fails to 
satisfy the Code. In either case, Bleszynski 
intended Chiloides to become a junior ob- 
jective synonym o^ Acigona. 

Bleszynski (1965, 1967) expanded his 
concept o{ Acigona to encompass species of 
the genera Coniesta Hampson, Eoreuma 
Ely, Haimbachia Dyar, Achilo Amsel, Do- 
nacoscaptes Zeller, Girdhiara Kapur and 
Xubida Schaus, all of which share a con- 
dition of the female genitalia where a nar- 
rowed extension or "bridge" from the an- 
terior margin of the eighth tergite extends 
down onto the ostium bursa. Bleszynski 
(1967) also transferred several species to 
Acigona from Chilo, Erupa, and Eufernal- 



dia and described one species, A. forsteri, 
in Acigona (Bleszynski, 1965). The mono- 
typic genus Occidentalia was included \xn- 
dQv Acigona by Klots (1970, 1983), but not 
by Bleszynski. 

Because Hampson's (1926) type desig- 
nation places Acigona in the Noctuidae, 
there is no name for Bleszynski's concept 
in the Pyralidae. Donacoscaptes Zeller is the 
oldest available name, but Bleszynski's ge- 
neric concept, regardless of the name ap- 
plied to It, is unsatisfactory. Even before the 
problem with the erroneous type species 
designation {or Acigona was brought to light 
(Nye, 1975; Retcher and Nye, 1984), work- 
ers had begun to recognize the validity of 
some of the genera synonymized by Ble- 
szynski (Klots, 1970; Gaskin, 1973). I also 
believe that Acigona (sensu Bleszynski) con- 
tains several good genera as well as species 
for which new genera probably should be 
proposed. For example, Coniesta, Eo- 
reuma, and Haimbachia are closely related, 
but can be separated by genitalic characters 
of both sexes, especially the females. Several 
tropical species of Eoreuma and Haimba- 
chia are incorrectly assigned. After an ex- 
amination of the male type and an associ- 
ated female, I am transferring one former 
member of Acigona, Chilo morbidellus 
Dyar, to Eoreuma as E. morbidella (Dyar) 
n. comb. This South American species has 
the typical uncus and gnathos of Eoreuma 
and most closely resembles E. loftini (Dyar) 
and E. evae Klots in the shape of the costal 
processes of the valvae. The female genitalia 
are also similar to those of E. loftini. 

No single action can correct the problems 
with the name and the concept of Acigona 
(sensu Bleszynski). A replacement name for 
the genus could come from designating the 
type species of the senior synonym, Dona- 
coscaptes validus Zeller, as a new type for 
the generic concept to which Bleszynski ap- 
plied the name Acigona, but this would still 
result in the synonymization of good genera. 
In removing the genera from synonymy, we 
do not resolve the problem of the species 

which do not belong in one of eight valid 
genera once included under Acigona. Be- 
cause T. cicatricella, the 'type' oi Acigona 
(sensu Bleszynski), is also the type of Chi- 
loides, it appears that there is an available 
generic name for this species. However, 
Chiloides Bleszynski 1963, is a junior hom- 
onym of Chiloides Butler 1881, (Lepidop- 
tera: Tortricidae) and thus unavailable. I 
therefore propose the following replace- 
ment name for Chiloides Bleszynski. 

Friedlanderia Agnew, New Name 

Chiloides Amsel 1949, nomen nudum. 

Chiloides Bleszynski 1963, preoccupied by 
ChUoidesBuXXtv 1881 (Lepidoptera: Tor- 

Acigona: Bleszynski and Collins 1962 (not 
Hiibner 1816 [1826]), incorrect type des- 

Type species: Tinea cicatricella Hiibner 
[1823H1824]:pl. 68, fig. 455. 

Diagnosis of genus. — Same as for the 
type cicatricella, provided by Bleszynski 
(1965) who figured both sexes (Plate 5, Figs. 
67-,_3), their genitalia (<5, Plate 44, Fig. 67; 
2, Plate 94, Fig. 67), and the larva (Figs. 
673.4). The heavily sclerotized ovipositor of 
the female is unusually shaped and distin- 
guishes Friedlanderia cicatricella (Hiibner) 
N. Comb, from species in other genera. The 
basally lobed uncus can be used to distin- 
guish the male from species in related gen- 
era. The degree of sexual dimorphism in the 
wing pattern of F. cicatricella is much more 
marked than in most Crambinae. The genus 
is named for Timothy P. Friedlander, whose 
knowledge of Lepidoptera provided me with 
a helpful introduction to the Crambinae. 

At this time, only the type species, F. ci- 
catricella, belongs in the genus as currently 
defined. The other species included by Am- 
sel (1949), Chiloides hederalis (Amsel), is a 
synonym of Thopeutis galleriella (Ragonot) 
(Bleszynski and Collins, 1962). 

No useful purpose would be served by 
transferring from Acigona to Friedlanderia 



those species originally described in Chilo, 
Erupa, and Eufernaldia. For the present, 
these misplaced species should revert to 
those genera until their generic affinities can 
be determined. Species from the genera once 
synonymized under Acigona have valid 
combinations available. One species with- 
out a valid combination, Acigona forsteri 
from China, is transferred to the Old World 
genus Coniesta to become Coniesta forsteri 
(Bleszynski) N. Comb, based on an illustra- 
tion of the male genitalia by Bleszynski 
(1965). Further study may necessitate plac- 
ing it elsewhere, but I feel that the present 
lack of a valid combination should be rec- 
tified for nomenclatural purposes. 


I thank Horace Burke, Timothy Fried- 
lander, Robert Wharton, and James Wool- 
ley of Texas A&M University for taxonom- 
ic and editorial advice on this manuscript; 
Frederick Rindge of the American Museum 
of Natural History for the loan of material; 
Douglas Ferguson of the Systematic Ento- 
mology Laboratory, Agricultural Research 
Service, for the loan of material and com- 
ments on the manuscript; and J. W. Smith, 
Jr. of Texas A&M University for providing 
support for this study. Approved by the 
Texas Agricultural Experiment Station as 
TA #21788. Research supported in part by 
Hatch project 6796 and a grant from the 
Rio Grande Valley Sugar Growers, Inc. 

Literature Cited 

Amsel, H. G. 1949. On the Microlepidoptera col- 
lected by E. P. Wiltshire in Irak and Iran in the 
years 1935 to 1938. Bull. Soc. Fouad I Entomol. 

Bleszynski, S. 1963. Studies on the Crambidae (Lep- 
idoptera). Review of the genera of the family 
Crambidae with data on their synonyms and types. 
Acta Zool. Cracov. 8: 91-132. 

. 1965. Crambinae. In Amsel, H. G., Reisser, 

H., and Gregor, F. eds., Microlepidoptera Pa- 
laearctica 1. Verlag Fromme, Wien. 553 pp. 

. 1967. Studies of the Crambinae (Lepidop- 

tera). Part 44. New neotropical genera and species. 
Preliminary checklist of neotropical Crambinae. 
Acta Zool. Cracov. 12: 39-1 10. 

Bleszynski, S. and R. J. Collins. 1962. A short cat- 
alogue of the world species of the family Cram- 
bidae (Lepidoptera). Acta Zool. Cracov. 7: 197- 

Butler, A. G. 1881. On a collection of nocturnal Lep- 
idoptera from the Hawaiian Islands. Ann. Mag. 
Nat. Hist. (5) 7: 392-408. 

Fletcher, D. S. and I. W. B. Nye. 1984. The Generic 
Names of Moths of the World. Volume 5 [Pyra- 
loidea]. Brit. Mus. (Nat. Hist.). London. 185 pp. 

Gaskin, D. E. 1973. A revision of New Zealand Chi- 
lonini (Lepidoptera: Pyralidae) and redescription 
of some Australian species. N. Z. J. Sci. 16: 435- 

Hampson, G. F. 1926. Descriptions of new genera 
and species of Lepidoptera Phalaenae of the 
subfamily Noctuinae (Noctuidae) in the British 
Museum (Natural History). British Mus. (Nat. 
Hist.), London. 641 pp. 

Hiibner, J. 1796 [1836]. Sammlung europaischer 
Schmetterlinge 8: (Tineae). 78 pp., 71 pis. Augs- 

. 1816 [1826]. Verzeichniss bekannter 

Schmettlinge [sic], pp. [1]-431. Augsburg. 

Klots, A. B. 1 970. North American Crambinae: Notes 
on the tribe Chiloini and a revision of the genera 
Eoreuma Ely and Xubida Schaus (Lepidoptera: 
Pyralidae). N.Y. Entomol. Soc. 78: 100-120. 

. 1983. Crambinae. In Hodges, R. W. et al, 

eds., Check List of the Lepidoptera of America 
North of Mexico. E. W. Classey Ltd. & Wedge 
Entomol. Res. Found. London. 284 pp. 

Nye, I. W. B. 1975. The Generic Names of Moths of 
the World. Volume 1 [Noctuoidea, part]. Brit. Mus. 
(Nat. Hist.). London. 568 pp. 


89(1), 1987, pp. 103-121 


Norman E. Woodley 

Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, c/o 
U.S. National Museum of Natural History NHB 168, Washington, D.C. 20560. 

Abstract.— Tht two Afrotropical pachygastrine genera Ashantina Kertesz and Meris- 
tomeringina James are reviewed. Two new generic synonymies are proposed: Neosolva 
Seguy is a new junior synonym o{ Ashantina (based on type species synonymy, N. dubia 
Seguy is a new junior synonym o{ Ashantina antennata Kertesz) and Agnathomyia Lindner 
is a new junior synonym of Meristomeringina. Lectotypes are designated for Ashantina 
antennata Kertesz and Neosolva dubia Seguy. A key to the six known species of Meris- 
tomeringina is provided as well as illustrations of male genitalia of all species, four of 
which are new (country of type locality in parentheses): M. aka (Zaire), M. cholo (Malawi), 
M. kontagora (Nigeria), and M. praestigiator (Uganda). 

The Pachygastrinae is a large subfamily 
of stratiomyid flies. Their extreme struc- 
tural diversity has made them a difficult 
group to work with and has resulted in the 
description of a large number of genera, each 
with few included species. Of the 62 Afro- 
tropical genera recognized by James ( 1980), 
38 are monotypic, only seven have five or 
more included species, and only eight occur 
outside of the Afrotropical Region. The only 
recent key to genera from the region is that 
of Lindner (1966), which excludes those 
found only in Madagascar ( 1 5 genera), and 
nine other genera subsequently described or 
discovered from mainland Africa. Conse- 
quently, even generic determinations within 
the subfamily can be difficult to make. 

The character states used to define the 
Pachygastrinae, loss of vein M, beyond the 
discal cell and the abdomen composed of 
five major segments (i.e. those beyond five 
much reduced), are found in other stratio- 
myids. Thus the monophyly of the subfam- 
ily has not been adequately demonstrated. 
Based on their studies of female terminalia 

of representative stratiomyid taxa, Naga- 
tomi and Iwata (1978) noted that the "Cli- 
tellariinae and Pachygasterinae may be het- 

This paper reviews two genera included 
by James (1952) in the tribe Meristomerin- 
gini, a tribe not maintained by him in sub- 
sequent publications. None of the seven 
character states mentioned by James (1952) 
is unique to the group. While the Afrotrop- 
ical genera included in the tribe by James 
are quite similar in general appearance, the 
possible monophyly of this suprageneric 
grouping requires much further documen- 
tation. The two genera treated in this paper 
may be identified using Lindner's ( 1 966) ge- 
neric key. The male genitalia for all known 
species in both genera are illustrated, which 
should allow for detection of additional 
species if they exist. 

Genus ASHANTINA Kertesz 

Ashantina Kertesz, 1914: 539. Type species, 
A. antennata Kertesz, by original desig- 



Fig. 1 . Ashantina antennata Kertesz, left lateral view of male head. 

Neosolva Seguy, 1953: 152. Type species, 
N. duhia Seguy, by original designation. 

T>\2ignos\s.— Ashantina is most easily rec- 
ognized by the structure of its antenna (Fig. 
1), in particular the eighth flagellomere, 
which is thick basally but quickly tapers to 
an arista-like apex. Only the basal portion 
of the eighth flagellomere is haired. Also, 
the rounded but protruding face, which ex- 
tends well below the lower eye margin (Fig. 
1), and which is squared when viewed an- 
teriorly, is characteristic. Both of these char- 
acter states are likely autapomorphic for the 
genus; I am aware of no other pachygas- 
trines with similar features. 

Other features exhibited by Ashantina, al- 
though found in other pachygastrines, are 
useful in characterizing the genus. They in- 
clude the elongate antennal flagellum, the 
spineless scutellum, vein R.+3 arising well 
beyond crossvein r-m, and the elongate ab- 

Remarks.— The synonymy of Neosolva 
with Ashantina has been confirmed by ex- 
amination of type material for each of the 
included species. Seguy 's (1953) original de- 
scription of Neosolva was brief, not partic- 
ularly useful, and has made subsequent rec- 

ognition of his taxon difficult. Neosolva was 
excluded from the Xylomyidae by Naga- 
tomi and Tanaka (1971), who did not ex- 
amine the types, but guessed that it might 
be a pachygastrine. James (1980: 274) treat- 
ed Neosolva as an unplaced genus of Stra- 

Ashantina runs rather ambiguously 
through paragraph 19 of Lindner's generic 
key. It should probably be traced through 
the alternate, paragraph 14, as the last flagel- 
lomere is "bristle-form," with only the base 
thickened and haired. 

Ashantina antennata Kertesz 

Ashantina antennata Kertesz, 1914: 539. 
Neosolva duhia Seguy, 1953: 152. NEW 

Type material.— One female syntype of 
A. antennata, hereby designated as the lec- 
totype, still exists in the British Museum 
(Natural History), London. The other syn- 
types, which were in the Hungarian Natural 
History Museum, were destroyed in 1956. 
The lectotype female is labeled: "Caught 
on leaf./Dunkwa, Ashanti, W. Africa. 
2. VIL 1 907. Dr. W. M. Graham. 1 908-245./ 
Ashantina 9! antennata Kert det. Kertesz/ 
LECTOTYPE Ashantina antennata Ker- 



tesz, 1914: 539 des. N. E. Woodley 1981." 
The specimen is in reasonably good con- 
dition but is missing the left antennal fla- 
gellum, the entire right antenna, the last two 
tarsomeres of the left foreleg, and the right 
hindleg beyond the trochanter. 

Four syntypes of A^. duhia Seguy are pres- 
ent in the Museum national d'Histoire na- 
turelle, Paris. I have examined one of these, 
a female, that is hereby designated as the 
lectotype. It is labeled: "Thio/MUSEUM 
PARIS Nimba (Guinee) M. Lamotte 
II.VI.42/Neosolva dubia Typ. Seguy 
9/LECTOTYPE Neosolva dubia Seguy, 
1953: 152 des. N. E. Woodley 1981/Ash- 
antina antennata Kertesz, 1914: 539 Det. 
N. E. Woodley 1981." The lectotype is in 
excellent condition, lacking only the last four 
tarsomeres of the left hindleg. 

Diagnosis.— As Ashantina is monotypic, 
A. antennata may be recognized by the ge- 
neric characters discussed above. Figs. 2, 4, 
and 5 illustrate the male terminalia, which 
should allow determination of this species 
with certainty and allow detection of further 
undescribed species if they exist. 

Distribution.— James (1980) states that 
A. antennata is found from Liberia to Zaire. 
Its distribution is shown in Fig. 6. 

Material examined. -CAMEROUN: 1 9, 
Dept. Nyong-Sanaga, Nkolbisson, Septem- 
ber 1963, L. G. Segers (MRAC); 2 3, 
Yaounde, 2600 ft., 29-30 May 1936, Van 
Zwaluwenberg& McGough (USNM). CEN- 
Dept. Lobaye, La Maboke, sur Whitfieldia 
longifolia, 31 August and 4 October 1970, 
L. Matile (MNHN). GHANA: 1 9 (lecto- 
type) Ashanti, Dunkwa, caught on leaf, 2 
July 1907, W. M. Graham (BMNH). 
GUINEA: (1 9 (lectotype ofN. dubia), Nim- 
ba [Mountains], Thio, February-April 1942, 
M. Lamotte (MNHN). LIBERIA: 4 <5, 1 9, 
Robertsport, Bendu, 26 February to 1 April 
1943, F. M. Snyder (AMNH). NIGERIA: 
1 5, Ile-Ife, 20 June 1970, J. T.Medler 
(CNC); 3 3, 4 9, Ile-Ife, 26 January 1 975 and 
16 March 1975, J. T. Medler (USNM). 

ZAIRE: 6 <5, Luebo, 27 April 1958, F. J. 
Frangoise (IRNSB); 1 <3, 1 9, Mbanza-Ngun- 
gu [as Thysville], 15°0'E, 5°30'S (AMNH); 
1 (5, Pare National Albert, W. Ruwenzori 
(1200-1500 m), March 1937, Hackars 
(MRAC); 1 a. Pare National Albert, Kivu, 
Rutshuru (riv. Rutshuru) 1000 m, 3 July 
1935, G. F. de Witte (MRAC). 

Remarks.— Ashantina antennata is a dis- 
tinctive Afrotropical species and is fairly 
common in collections. Lindner (1938, 
1955, 1966, 1970) gives numerous records. 
The sexes are quite similar in habitus. The 
frons and face in females are slightly wider 
than in males. Females are generally some- 
what darker in coloration, which is espe- 
cially noticeable on the thoracic pleura and 
dorsum of the abdomen. The species is quite 
variable in size; specimens examined range 
from 5.5 to 9.0 mm in length. 

The end of the abdomen in both sexes is 
unusual as the fifth tergite is shorter and 
narrower than the sternite, and it is appar- 
ently movable and capable of closing the 
end of the abdomen (Fig. 3). The aedeagal 
complex is complicated in structure and is 
somewhat simplified in my illustration (Fig. 
5). In profile it is nearly straight, and is not 
illustrated in this view because it was dif- 
ficult to delimit structures which were ob- 
scured by membrane. The homologies are 
difficult to determine, but the aedeagus is 
apparently strongly fused to the aedeagal 
valves (the lateral processes in the normal 
trifid aedeagal complex of stratiomyids). 


Meristomeringina James, 1952: 38. Type 

species, M. mimetes James, by original 

Agnathomyia Lindner, 1958: 124. Type 

species, A. combinata Lindner, by mono- 


Diagnosis.— Meristomeringina is the only 
genus of Afrotropical Pachygastrinae that 
has a combination of a simple antennal fla- 
gellum and a scutellum with two spines. The 



Figs. 2-5. Features of Ashantina antennata Kertesz. 2, Male genital capsule, dorsal view. 3, End of female 
abdomen, dorsal view. 4, Male epandrium and post-genital segments, dorsal view. 5, Male aedeagal complex, 
dorsal view. 

only other genus on mainland Africa with 
two scutellar spines is Goetghebueromyia 
Lindner, but this genus has the last antennal 
flagellomere aristate and has a much differ- 

ent habitus. Several pachygastrines from 

Madagascar also have two scutellar spines 

but are also very different in general form. 

Ktvadivks.— Meristomeringina is difficult 



Fig. 6. Distribution of Ashantina antennata Kertesz. 

to define with a conclusively autapomorph- 
ic character state. The shape of the antennal 
flagellum is characteristic but is of the more 
general, plesiomorphic form, i.e. without any 
fusion of flagellomeres or modification of 
the terminal ones to form a style or arista 
(Figs. 7-9). The antennae of the pachygas- 
trines are so variable in structure that char- 
acter polarities are difficult to determine in 
many cases. Presence of scutellar spines is 
also plesiomorphic within the subfamily, al- 
though the character state of having only 
two is rare in pachygastrines. The aedeagal 

complex is similar in all species included in 
the genus, and the presence of toothlike pro- 
cesses on the aedeagal valves may be aut- 
apomorphic for the genus. 

The six species now known of Meristo- 
meringina are extremely similar in mor- 
phology. Several species from central Africa 
can be identified with certainty only by ex- 
amining the male genitalia. Females are vir- 
tually identical, and the descriptions are 
based on specimens associated with males 
that have identical collecting data. The fe- 
male genitalia are not distinctly different. 




Figs. 7-9. Left lateral views of heads of Meristo- 
meringina. 7, Male of M. mimetes James. 8, Female 
of M. mimetes James. 9, Male holotype of M. com- 
binata (Lindner). 

The differences in structure of the female 
furca that I have observed between speci- 
mens is sHght, and none appear to be con- 
stant within and diagnostic between species. 
Further studies of females are needed based 
on additional material in series associated 
with males to assess variation and possible 
diagnostic characters. 

Virtually nothing is known of the natural 
history of Meristomeringina. They are per- 
haps crepuscular or nocturnal as both M. 
mimetes and M. kontagora have been col- 

lected at light. Two females that are prob- 
ably M. mimetes were collected on Zea 
mays. One unassociated female from Zaire 
was collected on large leaves of shrubs in 
gallery forest along a river. 

In the following portion of the paper, M. 
mimetes is redescribed completely, and 
treated first as it is the type species of the 
genus. For the five additional species, de- 
scriptions are based only on differences be- 
tween the respective species and M. mi- 
metes, because they are all so similar 
morphologically. It should be noted that all 
of the characters described for M. mimetes 
have been examined for each species. 

Key TO Species of Meristomeringina 

1 . White spot of anepistemum extending from 
dorsal, posterior comer at base of wing along 
notopleural suture, reaching anterior spiracle 

- White spot of anepistemum confined to dorsal, 
posterior comer 5 

2. Middle and hind femora with basal halves yel- 
lowish-white, apical halves brownish-black, the 
colors strongly contrasting; male gonostylus not 
strongly explanate 3 

- Middle and hind femora with basal thirds yel- 
lowish-white, apical two-thirds dark brownish- 
orange, the colors less strongly contrasting than 
in the three species keying out above; male 
gonostylus strongly explanate, more or less tri- 
angular in outline (Fig. 23) 

M. combinata (Lindner) 

3. Gonostylus of male genitalia with a long, slen- 
der, posteriorly directed digitate process; pos- 
terior margin of hypandrium notched medi- 
ally, thus bilobed (Fig. 11) ... A/, mimetes James 

- Gonostylus of male genitalia without long dig- 
itate process, but with shorter, more dorsally 
directed process; posterior margin of hypan- 
drium rounded, without medial notch (Figs. 
17,31) 4 

4. Male gonostylus with apical process simple, 
short and dorsally directed, sharp apically; pos- 
terodorsal comer of gonocoxites more truncate 

than in M. aka (Fig. 31) 

M. praestigiator, new species 

- Male gonostylus with apical process longer and 
recurved in a dorsal direction, the apex blunter, 
with several minute denticles; posterodorsal 
comer of gonocoxites narrower and more 
rounded than in M. praestigiator (Fig. 17) . . 
M. aka. new species 



Fig. 10. Distribution of species of A/m5/(?wm/?g/«a. 

Second segment of palpus dark; all femora 
strongly bicolored, basal halves yellowish-white, 
apical halves brownish-black; hind tibia whol- 
ly brownish black; second tarsomeres of mid- 
dle and hind legs blackish on at least apical 
halves; male gonostylus without posterior dig- 
itate process (Fig. 27) . . M. kont agora, new species 
Entire palpus pale; all femora weakly bicol- 
ored, basal halves yellowish-white, apical halves 
brownish-orange; hind tibia brownish-orange 
with moderately distinct paler ring medially; 
second tarsomeres of middle and hind legs 
wholly yellow; male gonostylus with posterior 
digitate process (Fig. 20) . . M. cholo, new species 

Meristomeringina mimetes James 

Meristomeringina mimetes J amts, 1952: 38. 

Type material.— The holotype male, in 
the collection of the California Academy of 
Sciences, San Francisco, is labeled: "Ber- 
tona Fr. Cameroons VII- 10-49 B. Malkin/ 
At light/Meristomer-ingina mimetes S James 
HOLOTYPE." The specimen is in excellent 
condition, missing only the last three tar- 
someres of the right front leg. 

Diagnosis. — This species is extremely 



similar to M. praestigiator and M. aka, dif- 
fering significantly only in structure of the 
male genitalia. The bilobed posterior mar- 
gin of the hypandrium and the gonostylus 
bearing a long, digitate posterior process 
(Fig. 11) are features that will separate M. 
mimetes from these other two species. The 
process of the gonostylus is longer in M. 
mimetes than in any other species in the 
genus. Unassociated females cannot be re- 
liably determined. 

Description.— Mfl/^: Head (Fig. 7) black- 
ish; eyes very large, hence frons very re- 
duced; ocellar tubercle moderately promi- 
nent, ocelli large; face very slightly convex, 
receding, finely punctate; occiput concave; 
face with very narrow, whitish tomentose 
margins along eyes; pilosity of head evenly 
set, hairs white, short, more or less ap- 
pressed, longer and more erect on lower fa- 
cial margin and genae; eyes with extremely 
short, sparse pilosity, visible only at high 
magnification; antennae simple, 0.82 to 0.90 
length of head, ratio of segments (for one 
paratype) 12:12:49[7:5:5:5:5:5:5:12], flagel- 
lum slightly tapering apically, first two seg- 
ments and base of flagellum brownish, be- 
coming nearly black distally, sometimes 
more yellowish-brown basally in some 
specimens; scape and pedicel with short, 
dark hairs, flagellum with velvety vestiture 
and short stubby hairs, most noticeable on 
last two flagellomeres; palpi small, with two 
subequal segments, pale whitish-yellow, 
with a few apical hairs; proboscis pale yel- 
lowish. Thorax black, pleura more brown- 
ish, post-pronotal lobes with outer halves 
yellowish, postalar calli brownish, scutellar 
spines whitish; anepistemum with creamy 
white spot on posterodorsal comer, becom- 
ing narrower and extending anteriorly along 
notopleural suture to anterior spiracle; pro- 
sternum brownish yellow; mesonotum 
moderately convex, densely but finely, gran- 
ulately punctate; scutellum simple, mod- 
erately convex, punctate as mesonotum, the 
two separated by about their length; meron 
+ katepimeron, laterotergite, and subscu- 

tellum with whitish tomentum; most of tho- 
rax set with whitish, semi-appressed pilos- 
ity, coppery colored on mesonotum and 
middle of scutellum, bare anterodorsally on 
anepistemum, posterior half of anepime- 
ron, entire meron + katepimeron, medio- 
tergite, and subscutellum; legs yellowish- 
white, but entire front tarsi, distal halves of 
mid and hind femora, hind tibiae except for 
indistinct annulus just basad of middle 
(sometimes extending more extensively to- 
ward base), and apical halves of second tar- 
someres and tarsomeres 3-5 of mid and hind 
legs brownish black; middle tibiae brown- 
ish, becoming paler apically; front femora 
gradually becoming orangish-yellow distal- 
ly; and front tibiae entirely orangish-brown; 
pilosity of legs very short, mostly pale but 
with some dark hairs on darkly pigmented 
regions, with longer, more erect hairs pres- 
ent on posterior surfaces of middle and to 
a lesser extent the hind femora; halter pale 
yellowish, knob partly brownish; wings 
evenly infuscated with grayish-brown, 
slightly darker in vicinity of radial sector 
veins, paler near base of wing; wings evenly 
set with microtrichia except at extreme base. 
Abdomen nearly twice as long as wide, 
brownish-black, proctiger and cerci more 
yellowish; finely, granularly punctate; with 
short, appressed pilosity, mostly pale but 
some is dark medially on tergites, longest 
on first segment. Male genitalia large, gon- 
ocoxites (Fig. 11) rounded, posterodorsal 
region truncate, the intemal corner with fine 
denticles near margin; posterior margin of 
hypandrium slightly produced with sharp, 
U-shaped medial notch rendering it distinct- 
ly bilobed; gonostyli strongly concave dor- 
sally, with sharp anteromedial comer and 
long, slender, posteriorly directed digitate 
process that is gently bent dorsally and long- 
er than the basal portion of the gonostylus; 
aedeagal complex (Figs. 12, 13) large, gently 
arcuate in profile, the three posterior lobes 
long and attenuated posteriorly, completely 
fused, with a few asymmetrically placed 
projecting teeth; attachment structure of ae- 



deagal complex fairly large, simple, broadly 
attached to gonocoxal apodemes; epan- 
drium (Fig. 14) large, explanate, gradually 
tapered posteriorly to a truncate apex. 
Length, 6.9 to 7.5 mm. 

Female: Differs from male as follows: 
Head (Fig. 8) with eyes smaller, dichoptic; 
frons 0.36 to 0.40 width of head, parallel 
sided above transverse sulcus, finely punc- 
tate except for a feebly developed medial 
carina extending from anterior ocellus to 
transverse sulcus, sometimes undeveloped 
anteriorly; transverse sulcus and surround- 
ing area slightly depressed; lower frons and 
face wider, but similar in structure to male; 
vertex slightly produced posteriorly, margin 
rounded; postocular orbits rounded, wid- 
ening ventrally; antennae 0.94 to 1.0 length 
of head, ratio of segments in one paratype 
10:10:36[5:4:4:4:3:3:4:9]. Thorax with me- 
sonotal pilosity usually wholly whitish, at 
most with slight golden cast on narrow, me- 
dial strip; halter entirely pale. Abdomen with 
pale pilosity more conspicuous along tergal 
sutures. Female genital furca extremely sim- 
ilar to the illustration for that of M. aka (see 
Fig. 16). Post-genital segments with cerci 
cylindrical, slender, second segment very 
slightly shorter than first. Length 7.5 to 7.9 

Distribution. — Known from Cameroun, 
Nigeria, and Zaire (Fig. 1 0). 

Material examined. -CAMEROUN: 8 3, 
2 2 (holotype, allotype, and 8 paratypes), 
Bertona, 10 July 1949, at light, B. Malkin 
(CAS, USNM, WSU); 1 <5, Env. Douala 
(MNHN). NIGERIA: 1 <5 (labeled as para- 
type), Ife Oyo Province, 19 March 1949, B. 
Malkin (WSU); 1 9, Zaria, Samaru, June 
1979, J. C. Deeming (NEW); 1 3, 2 9, Sa- 
maru, 21-29 July 1970, P. H. Ward, mer- 
cury vapor light trap (BMNH). ZAIRE: 1 3, 
1 9 Haut-Uele, Mauda, March 1925, H. 
Schouteden (MRAC). 

Two females with data NIGERIA: U. C. 
Ibadan, 1-2 August 1956, G. H. Caswell, 
Zea mays (BMNH) are almost certainly M. 
mimetes, but males are not known from this 

locality. Three additional unassociated fe- 
males seem most likely to be this species, 
but their identity is also uncertain: ZAIRE: 
Mayidi, 1945, P. Van Eyen (MRAC); Elis- 
abeth ville, January 1956-January 1957, Ch. 
Seydel, "a la lumiere" (MRAC); Elisabeth- 
ville, M. Bequaert, 29 November 1933, 
"dans galerie forestiere de la riv. Kimilolo 
courant sur large feuille d'arbuste" (WSU). 
The first two of these were reported by Lind- 
ner (1966) as M. mimetes. 

Remarks. — The type locality for this 
species was spelled "Bertona" on the labels 
and in James' paper, but it probably refers 
to Bertoua, Cameroun, as I have not been 
able to find any locality with the former 
spelling. The male from Ife Oyo Province, 
Nigeria was labeled as a paratype by James 
but not cited in the original publication. As 
the label is handwritten and looks very sim- 
ilar to the Bertona, Cameroons labels, I as- 
sume this was an oversight by James and 
the specimen was in the original type series. 

James (1952: 39) figured the male geni- 
talia, but his drawing is crude, as it was 
apparently drawn in situ without clearing. 
The male genitalia of M mimetes have gon- 
ostyli that are very similar to those found 
in M. cholo, differing mainly in the length 
of the digitate process which is shorter in 
the latter species. This structural similarity 
may be synapomorphic indicating a sister- 
species relationship between these two 
species. The gonostyli of M kont agora have 
the same general form, but lack a posterior 
process. It is not clear if this is a derived 
loss or a more plesiomorphic character state. 

A small amount of color variation on the 
legs has been observed, especially on the 
hind tibiae. Pale color is usually restricted 
to a vague, narrow annulus near the middle 
of the tibia, but sometimes is more exten- 
sive basally. Aside from slight size varia- 
tion, the species is quite uniform in ap- 

The females of M. mimetes and M aka, 
and presumably females of M praestigiator 
(presently unknown), are virtually identical. 








They cannot be identified reliably unless as- 
sociated with males. Although there is con- 
siderable overlap in size, M. mimetes has a 
general appearance slightly larger and more 
robust than M. aka. Also, the anterior ex- 
tension of the white spot on the anepister- 
num along the notopleural suture is usually 
slightly wider in M. mimetes than in M. 
aka. It is not known, however, if these subtle 
differences will remain distinct when more 
adequate series of each species are at hand. 

Meristomeringina aka Woodley, 
New Species 

Type material. — The holotype male, 
housed at the Institut Royal de Sciences Na- 
turelle de Belgique, Brussels, is labeled: 
"Congo Beige, P.N.G. Miss. H. De Saeger 
Aka/2, 22-v- 1 952 H. De Saeger. 35 1 4/Mer- 
istomerin-gina mimetes Lindner det Jam/ 
HOLOTYPE $ Meristomeringina aka N. E. 
Woodley 1986." The apical half of the left 
scutellar spine is missing, otherwise the 
specimen is in excellent condition. The ab- 
domen beyond the fifth segment is pre- 
served in glycerin in a microvial on the spec- 
imen pin. An allotype female, 10 male and 
1 female paratypes are designated, data being 
summarized in the material examined sec- 
tion below. 

Diagnosis. — Extremely similar to M. mi- 
metes and M. praestigiator, this species may 
be identified reliably only by examination 
of the male genitalia. It differs from M. mi- 
metes in not having a long digitate process 
on the gonostylus. Meristomeringina aka has 
the apical process of the gonostylus (Fig. 1 7) 
more strongly recurved than in M. praes- 
tigiator, and it possesses a few small den- 
ticles which are absent in the latter species. 
Also, the posterodorsal comer of the gon- 

ocoxites of M aka is narrower and less trun- 
cate than in other species in the genus. Fe- 
males unassociated with males cannot be 
reliably determined. 

Description.— Differs from M. mimetes 
as follows. Male: Head with antennae 
slightly shorter, 0.78 to 0.83 length of head, 
ratio of segments of holotype 8:8:23[4:3:2: 
2:2:2:2:6]. Thorax with creamy white spot 
of anepistemum with anterior extension 
along notopleural suture slightly narrower; 
coppery appressed pilosity of mesonotum 
less conspicuous; knob of halter with dark- 
ened areas more intense. Abdomen with 
male genitalia relatively larger than in M. 
mimetes, gonocoxites (Fig. 17) rounded, 
posterodorsal margin truncate but internal 
angle more produced and rounded, with 
denticles present over slightly larger area; 
posterior margin of hypandrium more 
strongly produced, sharply rounded but not 
emarginate medially; gonostyh (Figs. 15, 17) 
only slightly concave dorsally but narrower, 
with posterior region produced and re- 
curved dorsally, the apex fairly sharp with 
a few denticles; aedeagal complex (Figs. 18, 
1 9) quite similar to that of M. mimetes and 
M. praestigiator. Length 5.9 to 7.0 mm. 

Female: Extremely similar to female of 
M. mimetes. Differs from male as follows. 
Head: with frons 0.40 width of head, struc- 
ture and coloration virtually identical to fe- 
male of M mimetes; antennae 0.79 length 
of head, ratio of segments in allotype 8:8: 
29[4:3:3:3:3:3:3:7]. Thorax with mesonotal 
pilosity wholly whitish; haltere entirely pale. 
Abdomen with pale pilosity more conspic- 
uous along tergal sutures; genital furca (Fig. 
1 6) elongate; posterior bridge wide, rounded 
posteriorly; a long, very slender anterior ex- 
tension present on anterior bridge; median 

Figs. 1 1-14. Male genitalia oi Meristomeringina mimetes James. 1 1, Male genital capsule, dorsal view. 12, 
Male aedeagal complex, right lateral view. 13, Male aedeagal complex, dorsal view. 14, Male epandrium and 
post-genital segments, dorsal view. Abbreviations: as, attachment structure of aedeagal complex, c, cercus. ep. 
epandrium. gc, gonocoxites. gs, gonostylus. hyp, hypandrium. ic, internal comer of posterodorsal portion of 



Figs. 15-19. Male (drawn from holotype) and female (drawn from allotype) genitalia of Meristomehngina 
aka, new species. 15, Right gonostylus, posterolateral view. 16, Female genital furca, ventral view. 17, Male 
genital capsule, dorsal view. 18, Male aedeagal complex, dorsal view. 19, Male aedeagal complex, right lateral 
view. Abbreviations: ab, anterior bridge of furca, with anterior process, ma, median aperture, pb, posterior 
bridge, pip. posterolateral process. 

aperture moderately small, diamond shaped 
with rounded comers; posterolateral pro- 
cesses moderate in size, slender, apices 
slightly convergent; ninth tergite well scler- 
otized but incomplete medially. Cerci as in 
M. mimetes. Length 6.5 mm. 

Distribution. — Known only from Zaire 
(Fig. 10). 

Material examined.— ZAIRE (all as 
"Congo Beige"): 4 $ (holotype, 3 paratypes), 
P[arc] N[ational de la] G[aramba], Aka/2, 
22 May 1952, H. De Saeger 3514 (2 at 



IRNSB, 2 at MRAC); 1 9 (allotype), same 
data but 19 May 1952, H. De Saeger 3476 
(MRAC); 1 6, same data but Aka, 14 May 
1952, H. De Saeger 3450 (IRNSB); 1 6, 
P.N.G., Dedegwa, 2 1 May 1 952, H. De Sae- 
ger 3499 (IRNSB); 2 <5, P.N.G., Garamba/2 
(source), 6 June 1952, H. De Saeger 3583 
(MRAC); 1 <5, P.N.G., PFSK. 22/8, 10 June 
1952, H. De Saeger 3608 (IRNSB); 1 5, Uele, 
Buta, 450 m, 1 1 June 1935, G. F. de Witte 
1359 (MRAC); 1 <5, 1 9, Uele, Monga (riv. 
Bill), 450 m, 18 April to 8 May 1935, G. F. 
de Witte 1344 (MRAC). 

Etymology.— The specific name, a noun 
in apposition, refers to the type locality. 

Remarks. — Other than size, little varia- 
tion has been detected in this species. The 
size of the median aperture of the genital 
furca varies in the two females associated 
with males. This species is extremely sim- 
ilar to both M. mimetes and M. praestigia- 
tor, and can be reliably identified only by 
the structure of the male genitalia. All of the 
specimens from Pare National de la Garam- 
ba were identified previously as M. mimetes 
by Lindner and published as such (1965). 

The male genitalia of M. aka are quite 
similar in general form to those of M praes- 
tigiator. These two species are the only ones 
with the posterior margin of the hypan- 
drium not medially emarginate. The general 
form of the gonostyli in the two species is 
similar in being only slightly concave dor- 
sally, and in having the apical portion pro- 
duced and somewhat recurved dorsally, the 
apex of the process being acutely pointed in 
posterolateral view. 

Meristomeringina cholo Woodley, 
New Species 

Type material.— The holotype male is in 
the British Museum (Natural History). It is 
labeled: "Nyasaland. Cholo. R. C. Wood./ 
867 19.12.16/HOLOTYPE <5 Meristome- 
ringina cholo N. E. Woodley 1986." The 
type is in good condition, but is missing 
both antennae and the right hind leg beyond 
the trochanter. The abdomen beyond the 

fifth segment is preserved in a microvial 
with glycerin on the specimen pin. 

Diagnosis.— This species is one of two in 
which the white spot of the anepisternum 
does not extend anteriorly along the noto- 
pleural suture, the other being M. konta- 
gora. Meristomeringina cholo may be sep- 
arated from M. kontagora by its pale second 
palpal segment and presence of a digitate 
process on the male gonostylus (Fig. 20). It 
is the only species in the genus with an en- 
tirely pale second tarsomere on the middle 
and hind legs. 

Description. — Differs from M. mimetes 
as follows. Male: Head with creamy white 
coloration narrowly surrounding antennal 
bases, and a very narrow creamy band along 
lateral margin of occiput behind middle of 
eye; antennae missing. Thorax with creamy 
white spot of anepisternum confined to pos- 
terodorsal comer, not extending beyond 
posterior half of notopleural suture; anepi- 
meron and dorsal margin of katepisternum 
just below it brownish-orange; legs with 
overall lighter appearance: distal halves of 
all femora and entire front and middle tibiae 
brownish-orange; hind tibiae brownish with 
more distinct pale median annulus; and en- 
tire front tarsi and distal three tarsomeres 
of middle and hind legs dark brownish. Ab- 
domen slightly more brownish in color- 
ation; male genitalia with gonocoxites (Fig. 
20) more quadrate in general shape, pos- 
terodorsal region not as strongly truncate, 
the internal comer with finer denticles; pos- 
terior margin of hypandrium similar in 
shape but medial notch more nearly 
V-shaped and slightly smaller; gonostyli very 
similar, but anteromedial corner not as 
acute, and digitate process shorter, only 
about as long as basal part of gonostylus; 
aedeagal complex (Figs. 21, 22) shorter and 
broader, nearly straight in profile, with lat- 
eral margins developed outwardly and only 
thinly connected to the three medial lobes, 
appearing almost as a pair of accessory lobes, 
and bearing several irregular teeth. Length 
6.8 mm. 



Figs. 20-22. Male genitalia (drawn from holotype) of Meristomehngina cholo, new species. 20, Genital 
capsule, dorsal view. 2 1 , Aedeagal complex, dorsal view. 22, Aedeagal complex, right lateral view. 

Female: Unknown. 

Distribution. — Known only from the type 
locality in southern Malawi (Fig. 10). 

Material examined. — MALAWI (as Ny- 
asaland): 1 3 (holotype), Cholo, 1 9 Decem- 
ber 1916, R. C. Wood (BMNH). 

Etymology.— The species name, a noun 
in apposition, refers to the type locality. 

Remarks.— This species has a slightly 
paler visage than most others in the genus, 
the darker regions of the legs being brown- 
ish-orange rather than blackish. In this re- 
gard it resembles M. combinata. The male 
genitalia are most similar to those of M. 
mimetes, as discussed under that species. 
However, the aedeagal complex of M cholo 
is much shorter and proportionately broad- 
er than in any other known species in the 

Meristomeringina combinata (Lindner), 
New Combination 

Agnathomyia combinata Lindner, 1958: 

Type material.— The male holotype, in 
the Natal Museum, Pietermaritzburg, South 
Africa, is labeled: "Trelawney. S.R. 10:XII: 

53 N. J. Myers./Agnathomyia combinata 
Lind Lindner det./Typus Lindner 1957/ 
Meristomeringina combinata (Lindner) det. 
Woodley 1984." The specimen is in good 
condition, although the head is slightly de- 
tached. The terminalia are preserved in a 
microvial with glycerin attached to the pin 
of the specimen. 

Didignosis. — Meristomehngina combi- 
nata may be distinguished from the other 
three species that have the anepistemal spot 
extending to the anterior spiracle by its paler 
legs, the darkest color on femora being 
brownish-orange rather than blackish. It is 
unique in the genus in having the male gon- 
ostyli large, more or less triangular in out- 
line, and strongly explanate in dorsal view 
(Fig. 23). 

Description. — Differs from M. mimetes 
as follows. Male: Head (Fig. 9) with slight 
brownish tinge; antennae shorter, 0.60 
length of head, ratio of segments in holotype 
10:7:26[5:3:3:2:3:3:2:5], flagellum some- 
what more compact; palpi with second seg- 
ment distinctly darker yellow on apical half 
Thorax slightly more brownish in general 
coloration, creamy white spot of anepister- 

Figs. 23-26. Male genitalia (drawn from holotype) of Meristomeringina combinata (Lindner). 23, Genital 
capsule, dorsal view. 24, Aedeagal complex, right lateral view. 25, Aedeagal complex, dorsal view. 26, Epandrium 
and post-genital segments, dorsal view. 


num not extending quite as far ventrally 
along anterior border of anepimeron; scu- 
tellar spines slightly shorter and thicker; 
mesonotal pilosity a little more bronzy and 
less conspicuous; legs in general paler in col- 
or, only front tarsi, distal halves of second 
tarsomeres and tarsomeres three to five of 
middle and hind legs brownish, otherwise 
coxae and bases of femora pale yellowish, 
remainder of legs brownish-orange, with all 
tibiae grading to brownish distally; wings 
with very slightly paler infuscation. Abdo- 
men slightly more brownish; male genitalia 
with gonocoxites (Fig. 23) with posterodor- 
sal region narrower, the internal comer 
without denticles; posterior margin of hy- 
pandrium more strongly produced, medial 
notch deeper and v-shaped, thus the process 
more sharply bilobed; gonostyli much dif- 
ferent, large and explanate, shallowly con- 
cave dorsally, more or less triangular in out- 
line in dorsal view, without distinct 
processes; aedeagal complex (Figs. 24, 25) 
more robust posteriorly, with a pair of large, 
lateral tooth-like projections that are nearly 
symmetrical. Length 8.0 mm. 

Female: Unknown. 

Distribution.— Known only from Zim- 
babwe (Fig. 10). 

Material examined. — ZIMBABWE (as 
Southern Rhodesia): 1 $ (holotype), Trelaw- 
ney, 10 December 1953, N. J. Myers (NM). 

Remarks. — Lindner (1958) considered 
Agnathomyia to be characterized by anten- 
nal structure with "einem Endgriffel und 
einer feinen apikalen Borste," this appar- 
ently being the main diflference between his 
genus and Meristomeringina. Examination 
of the type failed to confirm his description 
of this feature. The last flagellomere is sim- 
ple with a few short apical hairs. Lindner's 
(1958: 123, Fig. 3) figure is crude and does 
not adequately show details of the anten- 
nae. Thus in Lindner's (1966) later key, this 
species should not trace through paragraph 
14, but through its alternate, paragraph 19. 

The slightly paler coloration of this species 
differs from most members of the genus but 

is similar to that found in M. cholo. The 
large, explanate male gonostyli are unique 
in the genus, while other features of the male 
genitalia are quite similar to those of con- 

Meristomeringina kontagora Woodley, 

New Species 

Type material.— The male holotype is in 
the British Museum (Natural History) and 
bears the labels: "NIGERIA: N.W. State. 
Kontagora River. 3 mis. from Niger. 
13.viii.l970. P.H. Ward. B.M. 1970-604./ 
Meristomeringina mimetes James det. S. J. 
Chambers 1973/HOLOTYPE <5 Meristo- 
meringina kontagora N. E. Woodley 1986." 
The specimen is in excellent condition. The 
abdomen beyond the fifth segment is pre- 
served in glycerin in a micro vial on the spec- 
imen pin. An allotype and one paratype are 
designated, with data presented in the ma- 
terial examined section. 

Didignosxs. — Meristomeringina konta- 
gora is unique in the genus in having a dark 
second palpal segment. This species and M. 
cholo both have the pale anepisternal spot 
not extending forward to the anterior spi- 
racle, but M. kontagora is darker with con- 
spicuously bicolored legs. It is the smallest 
species in the genus. 

Description. — Differs from M. mimetes 
as follows. Male: Head with antennae 0.87 
length of head, ratio of segments of holotype 
10:8:30[5:4:3:3:3:3:3:6], flagellum slightly 
more compact, first segment brownish, sec- 
ond yellowish-brown, ffagellum with first 
flagellomere brownish, others brownish- 
black; palpi with second segment brownish, 
conspicuously darker than first; proboscis 
yellowish. Thorax with postalar calli dark 
brownish; anepistemum with creamy white 
spot confined to posterodorsal comer, not 
extending anteriorly along notopleural su- 
ture; prostemum brownish; mesonotum 
with coppery hairs less noticeable; legs yel- 
lowish-white with bases of hind coxae 
brownish, apical halves of all femora, all 
tibiae, front tarsi, and apical halves of sec- 



Figs. 27-29. Male genitalia (drawn from holotype) oi Meristomeringina kontagora. new species. 27, Genital 
capsule, dorsal view. 28, Aedeagal complex, dorsal view. 29, Aedeagal complex, right lateral view. 

ond tarsomeres and tarsomeres three to five 
of middle and hind legs brownish black; hal- 
ter with knob brownish black; wing with 
infuscation grayish, a little more extensively 
hyaline basally, extreme apex whitish hya- 
line. Abdomen black; dorsal pilosity mostly 
brownish-black; male genitalia with gono- 
coxites (Fig. 27) rounded, internal comer of 
posterodorsal region without denticulate 
margins; process of posteromedial margin 
of hypandrium more quadrate, the medial 
notch slightly more v-shaped; gonostyli 
strongly concave dorsally but without dig- 
itate process, only vaguely bilobed along 
posterointernal margin; aedeagal complex 
(Figs. 28, 29) with posterior portion broad- 
er, more sharply bilobed apically, and with 
the tooth-like projections larger; attach- 
ment structure reduced in size and more 
narrowly connected to gonocoxal apo- 
demes. Length 5.3 mm. 

Female: Similar to male except as fol- 
lows. Head with frons 0.37-0.38 width of 
head, parallel-sided above transverse sul- 

cus, very similar to M. mimetes in structure, 
except that anterior portion of medial carina 
is not developed and this region is more 
impressed; antennae 0.77-0.85 length of 
head, ratio of segments in allotype 7:7:26[4: 
3:3:3:3:3:2:5]. Thorax with mesonotal pi- 
losity entirely pale; halter entirely pale; ter- 
gal sutures mostly surrounded by pale pi- 
losity; female genitalia not examined. Length 
6.0-6.3 mm. 

Distribution. — Known only from Nigeria 
(Fig. 10). 

Material examined.— NIGERIA: 1 <5 (ho- 
lotype), N. W. State, Kontagora River 3 
miles from Niger, 13 August 1970, P. H. 
Ward (BMNH); 1 9 (allotype), Igbo Ora, 18 
August 1964, B.R.L., at light (BMNH); 1 9, 
Ile-Ife, 2 August 1969, J. T. Medler (CNC). 

Etymology.— The species name, a noun 
in apposition, refers to the type locality. 

Remarks.— This species, the smallest in 
the genus, is uniform in appearance as far 
as the few known specimens indicate. The 
general structure of the male gonostyli is the 



Figs. 30-33. Male genitalia (drawn from holotype) of Meristomeringina praestigiator, new species. 20, Right 
gonostylus, posterolateral view. 3 1 , Genital capsule, dorsal view. 32, Aedeagal complex, dorsal view. 33, Aedeagal 
complex, right lateral view. 

same as in M. mi metes and M. cholo, but 
in M. kontagora there is no digitate process. 

Meristomeringina praestigiator Woodley, 
New Species 

Type material. — The holotype male is in 
the British Museum (Natural History) 
and is labeled: "Para-type/van Someren 
BWAMBA 7-8, 1946/Pres by Com Inst Ent 
BM 1953-357/Meristomer-ingina mimetes 
ristomeringina praestigiator N. E. Woodley 
1986." The specimen is in excellent con- 
dition although missing the right antennal 
flagellum. One paratype is designated with 
data presented in the material examined 

Diagnosis.— Extremely similar to M. aha 
and M. mimetes, Meristomeringina praes- 
tigiator may be identified reliably only by 

the male genitalia. It lacks the digitate pro- 
cess of the gonostylus and median notch of 
the posterior margin of the hypandrium 
found in M. mimetes (Fig. 30). The apical 
portion of the gonostylus is not as sharply 
produced and recurved as in M. aha, and 
lacks apical denticles. 

Description. — Differs from M. mimetes 
as follows. Male: Head with antennae 
slightly shorter, 0.80 length of head, 11:8: 
30[4:3:3:3:2:3:3:9], second segment and base 
of flagellum yellowish-brown. Thorax with 
legs essentially as in M. mimetes, but with- 
out trace of pale annulus on hind tibiae; 
knob of halter more extensively brownish. 
Abdomen with male genitalia with gono- 
coxites (Fig. 3 1 ) rounded, posterior margin 
of hypandrium more strongly produced, but 
evenly rounded, without medial notch; gon- 
ostyli (Figs. 30, 31) rather small, simple, 


with a sharp process directed dorsally near 
apex, without denticles; aedeagal complex 
(Figs. 32, 33) with slightly larger tooth-like 
projections. Length 7.4 mm. 

Female: Unknown. 

Distribution. — Known only from south- 
western Uganda and eastern Zaire (Fig. 10). 

Material examined. — UGANDA: 1 <5(ho- 
lotype), Bwamba, July-August 1946, van 
Someren (BMNH). ZAIRE: 1 <5, Pare Na- 
tional Albert, Secteur Nord, riv. Ngokoi, 
affl. Talya, 1 100 m, P. Vanschuytbroeck VS 
37 (MRAC). 

Etymology.— The species name, a noun 
in apposition meaning "imposter" or "de- 
ceiver," refers to the extreme similarity of 
M. praestigiator to M. mimetes, causing its 
inclusion in the type series of the latter. 

Remarks. — This species is most similar 
to M. aka, and the features in common of 
the male genitalia are discussed under that 
species. Females are unknown, but are 
probably nearly indistinguishable from those 
of M. mimetes and M. aka. 


I thank the following curators who have 
been generous in loaning material used in 
this study: Paul H. Amaud, Jr., California 
Academy of Sciences, San Francisco, Cali- 
fornia (CAS); John Chainey, British Mu- 
seum (Natural History), London, England; 
Eliane de Coninck, Musee Royal de I'Af- 
rique Centrale, Tervuren, Belgium (MRAC); 
Patrick Grootaert, Institut Royal des Sci- 
ences Naturelle de Belgique, Brussels, Bel- 
gium (IRSNB); J. G. H. Londt, Natal Mu- 
seum, Pietermaritzburg, South Africa (NM); 
Loi'c Matile, Museum national d'Histoire 
naturelle, Paris, France (MNHN); H. J. Tes- 
key, Canadian National Collection, Biosys- 
tematics Research Institute, Ottawa, Can- 
ada (CNC); William J. Turner, James 
Entomological Collection, Washington State 

University, Pullman, Washington (WSU); 
and P. Wygodzinsky, American Museum of 
Natural History, New York (AMNH). J. C. 
Deeming kindly gave me one specimen of 
M. mimetes (NEW). I also thank Wayne N. 
Mathis of the Smithsonian Institution, 
Washington, D.C. and D. A. Nickle and R. 
V. Peterson of the Systematic Entomology 
Laboratory for critically reviewing the 

Literature Cited 

James, M. T. 1952. The pachygastrine tribe Meris- 

tomeringini, with descriptions of a new genus and 

species (Diptera. Stratiomyidae). Ann. Entomol. 

Soc. Am. 45: 38-43. 
. 1980. 20. Family Stratiomyidae, pp. 253- 

274. In R. W. Crosskey (ed.). Catalogue of the 

Diptera of the Afrotropical Region. British Mu- 
seum (Natural History), London, 1437 pp. 
Kertesz, K. 1914. Vorarbeiten zu einer Monographic 

der Notacanthen. XXIII-XXXV. Ann. Hist.-Nat. 

Mus. Natl. Hung. 12: 449-557. 
Lindner, E. 1938. Stratiomyiiden aus dem Kongo- 

Gebiet. (Diptera). Bull. Mus. Roy. Hist. Nat. Belg. 

14: 1-35. 
. 1955. Congo-Stratiomyiidae (Dipt.). Rev. 

Zool. Bot. Afr. 52: 241-245. 
. 1958. Athiopische stratiomyiiden (Diptera). 

IV. J. Entomol. Soc. South. Afr. 21: 121-128. 
. 1965. Stratiomyiidae (Diptera Brachycera). 

Explor. Pare. Natl. Garamba Miss. H. de Saeger 

46: 45-65. 
. 1966. Stratiomyiiden aus dem Kongo im 

Musee Royal de TAfrique centrale in Tervuren, 

mit einer Bestimmungstabelle der Unterfamilie der 

afrikanischen Pachygasterinae. Rev. Zool. Bot. Afr. 

73: 351-384. 
. 1970. Westafrikanische Stratiomyiden aus 

dem Museum national d'Histoire naturelle de Paris. 

Bull. Inst. Fr. Afr. Noire 32: 817-828. 
Nagatomi, A. and K. Iwata. 1978. Female terminalia 

of lower Brachycera-II. Beitr. Entomol. 28: 263- 

Nagatomi, A. and A. Tanaka. 1971. The Solvidae of 

Japan. Mushi 45: 101-146. 
Seguy, E. 1953. La reserve naturelle integrale du Mt. 

Nimba. Fasc. 1. X. — Dipteres. Mem. Inst. Fr. Afr. 

Noire 19: 151-164. (1952). 


89(1), 1987, pp. 122-125 






(GD) Alan Retcher Research Station, Queensland Department of Lands, 27 Magazine 
Street, Sherwood, Queensland, Australia 4075; (WAP) North American Field Station, 
Queensland Department of Lands, 2714 Pecan Drive, Temple, Texas 76502 USA. 

Abstract.— Tht gall forming cecidomyiid fly Neolasioptem lathami Gagne, collected on 
Baccharis halimifolia and B. neglecta, was studied to determine its host specificity and, 
thereby, its suitability as a biological control agent for B. halimifolia in Australia. Emerging 
flies were confined to small containers in which cut stems of a range of plant species were 
available for oviposition. No eggs were laid on a plant other than a Baccharis species. 
These results and the absence of any field host records to the contrary demonstrated that 
this fly was specific to Baccharis. Permission to release this fly in Australia was granted 
by the appropriate authorities. 

The North American shrub Baccharis 
halimifolia L. (Asteraceae: Astereae) is a se- 
rious weed in southeastern Queensland and 
northeastern New South Wales, Australia. 
For over 25 years a concerted eflbrt has been 
made to find suitable biological control 
agents. During that time extensive surveys 
of both North and South America were con- 
ducted to locate host specific insects suitable 
for release into Australia. The most suc- 
cessful introduction to date has been a ceci- 
domyiid fly, Rhopalomyia californica Felt 
that galls the terminal stems of B. pilularis 
DC. in California (McFadyen, in press). 

Five species (two found in the United 
States and three in Argentina) of the genus 
Neolasioptera (Diptera: Cecidomyidae) also 
form galls on Baccharis spp. (Gagne 1971). 
The North American species, both reported 
from B. halimifolia, are A^. lathami Gagne, 
which forms a soft globular gall, and the 

much rarer A^. baccharicola Gagne, which 
forms a hard cylindrical stem gall. There 
are a further 1 2 Nearctic species of Neola- 
sioptera that have been reared from stems 
of Asteraceae (Gagne, 1971). 

This paper reports studies that were con- 
ducted to determine the host specificity of 
A^. lathami to obtain permission for the re- 
lease of the organism in Australia. 


N. lathami is a small, rather fragile look- 
ing fly with a greatly reduced gut and ap- 
parently non-functional mouthparts. In the 
laboratory the flies were never observed to 
ingest either food or water or to produce 
feces. Mating occurred soon after emer- 
gence of the flies. The females were readily 
distinguished by the reddish color of their 
abdomens that were distended by vermil- 



lion colored eggs. Oviposition occurred soon 
after mating with each female producing 
100-150 eggs. The adults were short lived 
with most surviving 3-5 days. 

Eggs were laid mostly on the surface of 
stem terminals but some were laid in leaf 
axils some distance down the stems. Neo- 
nate larvae entered the stems by pushing 
between the unopened leaf bud scales. Mor- 
tality was high at this stage and only a few 
gained entry and became established. 

Communal galls formed as a result of sev- 
eral larvae entering the one bud. Fully de- 
veloped galls vary from 1 to 3 cm in di- 
ameter; the size of the gall depending on the 
number of larvae present and also the rate 
of plant growth. A large gall might have as 
many as 15 larvae each in an individual 

Before pupating, the mature larva re- 
moved all but a layer of epidermal cells from 
the distal end of the chamber giving it a 
"windowed" appearance. The larva then 
pupated in the chamber. Before eclosion the 
pupa moved to the outer end of the chamber 
and pushed through the remaining epider- 
mal cells with the aid of the antennal horns. 
When all but a few abdominal segments 
protruded from the chamber the pupal skin 
split irregularly and the fly emerged. The 
pupal case remained attached to the gall. 

Host Range, Distribution and 

Galls of this fly have been reported pre- 
viously on B. halimifoUa along the eastern 
seaboard of the United States from New 
York to Mississippi (Gagne, 1971). In the 
present study galls of this fly were also col- 
lected west of this distribution as far as Del 
Rio in western Texas. Furthermore, galls 
were often collected from B. neglecta Brit- 
ton, a new host record for this insect. 

The galls were most abundant in spring. 
In Texas, mature galls were found as early 
as the end of March. By end of spring, par- 
asitism appeared to be the most important 

Table 1 . Mean egg counts on bouquets of plants 
exposed to N. lathami emerged from galls collected 
from either B. neglecta or B. halimifoUa. 








Baccharis halimifoUa L. 



Baccharis neglecta Britt. 



Solidago altissima L. 

Chrvsothamnus nauseosus 

(Pall.) Britt. 

Aster novae-angliae L. 

Helianthus annuus L. 

Dahlia pinnata Cav. 

mortality factor. It was then quite common 
to fail to recover a single fly from a collec- 
tion of galls. Only occasional galls were seen 
in Texas during the remainder of the year. 
In Texas, galls were never very abundant 
and only rarely were more than 10-20 galls 
found on the one plant. At these population 
levels the insect had little effect on the growth 
of mature plants. However, in Florida seed- 
ling plants only a few inches high were seen 
with as many as five large galls and these 
caused severe stunting. 

Host Specificity Testing 

Oviposition. — Because the aduh fly is the 
only mobile stage the species could be con- 
sidered monophagous if specificity of ovi- 
position could be satisfactorily demonstrat- 
ed. Two series of tests were therefore 
conducted to determine the specificity of 

A smaller test using plants closely related 
to Baccharis was conducted in Texas. In 
each of two 35 x 27 x 17 cm clear plastic 
cages two bouquets of foliage and growing 
tips of each of seven plants were randomly 
placed. Into one cage were placed galls col- 
lected from B. halimifolia near Conroe, 
Texas and into the other were placed galls 
collected from B. neglecta near Temple, 
Texas. After sufficient exposure to the 
emerging flies, the bouquets were examined 



Table 2. List of plant species against which N. la- 
thami was tested in order to obtain permission to re- 
lease it in Australia. 

APIACEAE: Daucus carota L.; Pastinaca saliva L. 
ANACARDIACEAE: Mangifera indica L. 
ASTERACEAE: Bacchahs halimifolia L.; Carthamus 

tinctorius L.; Chrysanthemum sp.; Dahlia sp.; He- 

lianthus annuus L.; Lactuca sativa L. 
BRASSICACEAE: Brassica oleraceae (L.) Alef.; Bras- 

sica rapa L. 
BROMELIACEAE: Ananas comosus (L.) Merr. 
CARICACEAE: Carica papaya L. 
CHENOPODIACEAE: Beta vulgaris L. 
CONVOLVULACEAE: Ipomoea batatas (L.) Lam. 
CUCURBITACEAE: Cucumis melo L.; Cucumis sa- 

tivus L.; Curcubita maxima Duch. 
FABIACEAE: Arachis hypogaea L.; Centrosema pu- 

bescens Benth.; Desmodium canum (Gmel.); Glycine 

wightii (R. Grab, ex Wight & Am.) Verde; Glycine 

max L. Merr.; Medicago sativa L.; Phaseolus atro- 

purpureus DC; Phaseolis vulgaris L.; Pisum sativum 

L.; Stizolobium 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. 
^OhCEAE: Avena sativa L.; Digitaria decumbens Stent.; 

Pannicum maximum Jacq.; Paspalum dilatatum 

Poir.; Pennisetum clandestinum Chiov.; Saccharum 

ojficinarum L.; Sorghum vulgare L.; Triticum aes- 

tevum L.; Zea mays L. 
PROTEACEAE: Macadamia integrifolia Maid & 

ROSACEAE: Fragaria vesca L.; Malus sylvestris Mill.; 

Prunus domestica L.; Prunus persica (L.) Batch.; Fy- 

ri« communis L.; i?05a sp. 
RUTACEAE: OYrM5 //mo« (L.) Burm. F.; Citrus par- 

adisi Macfady.; Citrus reticidata Blanco; Citrus sin- 

sensis (L.) 
SAPINDACEAE: Litchi chinensis Sonn. 
SOLANACEAE: Capsicum annuum L.; Lycopersicum 

esculentum Miller; Nicotiana tabacum L.; Solanum 

tuberosum L. 
VITACEAE: Vitis vinifera L. 
ZINGIBERACEAE: Zingiber officinale Roscoe. 

under a binocular microscope and any eggs 
deposited were counted. The results (Table 
1) indicated that both populations of flies 
oviposited on both species of Baccharis but 
not on any other plant. 

A more comprehensive testing program, 
designed to satisfy the Australian Depart- 
ment of Health's requirements for intro- 
duction into Australia, was conducted using 
65 species of plants (Table 2). This hst of 
plants included the most important agri- 
cultural species. Bouquets of the growing 
tips of 8 plant species from the list, together 
with a bouquet of 5. halimifolia were placed 
into glass and stainless steel aquaria (60 x 
36 X 40 cm). Twenty-five pairs of flies from 
galls collected in Florida were placed in each 
cage. Each test was duplicated. Eggs were 
counted after a three day duration. Many 
eggs were deposited on all the B. halimifolia 
controls but none were found on any other 

Gall development tests.— A series of tests 
was carried out in an insectary to determine 
on which plants galls would develop. Each 
group of well-developed potted plants was 
infested with 25 pairs of flies collected in 
Florida. Each test was duplicated. After three 
days the plants were placed in a glasshouse. 
Hatching, movement of larvae and gall for- 
mation were observed and recorded. Eggs 
were found only on the B. halimifolia plants. 
After three to five days, larvae emerged and 
entered the young tips. There was no move- 
ment of these larvae onto other plants. Galls 
were formed in two weeks and the life cycle 
was completed in six to eight weeks. 


The experimental work supported the 
previous host records from the field by in- 
dicating that N. lathami is host specific to 
those species of Baccharis on which it is 
found in the field (i.e. B. halimifolia and B. 
neglecta). Permission was therefore sought 
and obtained to import and release this in- 
sect for the biological control of 5. halimifo- 
lia in Australia. 

Although this fly does not cause dramatic 
control in its native habitat, this does not 
preclude it from being successful in a new 
environment. The fly has a high reproduc- 
tion rate and a short life cycle. The fact that 



it is usually at low population densities is 
due primarily to the high rate of attack by 
parasites. Before being released in Australia 
parasites will be very carefully eliminated 
from the population in a quarantine facility. 
Without these parasites a much higher pop- 
ulation of the fly may develop in Australia, 
perhaps to a level where significant effect on 
the population of the weed may occur. 


We thank Raymond J. Gagne of the Sys- 
tematic Entomology Laboratory, Agricul- 

tural Research Service, USDA, Washing- 
ton, D.C. for authoritative identifications of 
our material and other information and help. 

Literature Cited 

Gagne, R. J. 1971. Two new species of North Amer- 
ican Neolasioptera from Baccharis (Diptera: Ce- 
cidomyiidae— Compositae). Proc. Entomol. Soc. 
Wash. 73: 153-157. 

McFadyen, P. J. In press. Introduction of the gall fly 
Rhopalomyia californica from the USA into Aus- 
tralia for the control of the weed Baccharis hali- 
mifolia. Proc. VI Int. Symp. Biol. Contr. Weeds, 
Vancouver, 1984. 


89(1), 1987, pp. 126-131 





Jose A. Mari Mutt 

Department of Biology, University of Puerto Rico, Mayagiiez, Puerto Rico 00708. 

Abstract.— Se\x2L cryptica n. sp. is described from specimens collected in Florida and 
Arizona. This species was previously confused with S. bipunctata (Packard), which is 
redescribed from type material and specimens from Nebraska. Seira distincta Mari Mutt 
is recorded for the first time from the United States and Mexico. 

During a recent review of the Puerto Ri- 
can species of Seira (Mari Mutt, 1987) I 
considered one local species as possibly S. 
bipunctata (Packard), described in 1873 
from Texas and redescribed by Christiansen 
and Bellinger (1980) from specimens col- 
lected in various localities of the United 
States. These authors reported a degree of 
variation in coloration and chaetotaxy that 
suggested their samples contained more than 
one species. Christiansen and Bellinger had 
considered this possibility but preferred to 
list all their records under one species. 

Thanks to Kenneth Christiansen, who has 
placed at my disposal his collection of S. 
bipunctata, and to the Museum of Com- 
parative Zoology of Harvard University, 
which lent the necessary type material, I 
have been able to reinterpret the North 
American material of S. bipunctata. Some 
of the specimens in the Christiansen collec- 
tion belong to Packard's species while oth- 
ers belong to a new species that is described 
below. I received also from Dr. Christiansen 
some specimens from Massachusetts and 
Mexico that represent new records for Seira 
distincta Mari Mutt (1986), known previ- 
ously only from Puerto Rico. 

The type series of Seira bipunctata and 
the holotype of Seira cryptica n. sp. are de- 

posited in the Museum of Comparative Zo- 
ology, Cambridge, Massachusetts. The re- 
maining material is deposited in the 
collection of Dr. Christiansen, Department 
of Biology, Grinnell College, Iowa. In the 
species descriptions and discussions. Ant. 
2, Th. 2, Abd. 2, etc. means second antennal 
segment, second thoracic segment, etc. 

Seira cryptica Mari Mutt, New Species 

Seira bipunctata (Packard): Christiansen and 
Bellinger 1980: 925-926, Fig. 756 B, E, 
F (Florida), misidentification. 

Description. — Length from front of head 
to end of Abd. 6 up to 1.8 mm. Body with 
a lateral band of pigment along its length, 
pigment extending dorsally but with dimin- 
ishing intensity, leaving dorsum of body un- 
pigmented (Fig. 12, see also Christiansen 
and Bellinger, 1980: 926, Fig. B). Some 
lighter specimens almost white, darker 
specimens with pigment extending almost 
to midline of body. Antennae and legs light- 
ly pigmented throughout length. Manu- 
brium with some pigment basally. Anterior 
margin of mesonotum rounded, not pro- 
jecting over head. Apex of Ant. 4 without 
proturberance or pin seta but with bilobed 
papilla (Fig. 13). Head rounded, distribu- 

Figs. 1-7. Seira bipunctata. 1, Distribution of head macrochaetae. 2, Distribution of body macrochaetae, 
bothriotricha (wavy lines) and pseudopores (x) (setae signalled by arrows are absent in some specimens— see 
text. On Th. 2, a broken line surrounds the external posterior group of macrochaetae). 3, Mucro. 4, Metathoracic 
claws. 5, Apex of Ant. 4. 6, External labial papilla. 7, Labral papillae. 





tion of macrochaetae as in Fig. 14. Eyes G 
and H reduced, not visible in cleared spec- 
imens. Outer labral papillae rounded, inner 
papillae conical and not bifurcated (Fig. 1 3). 
Body macrochaetotaxy as in Fig. 1 1 ; 2 + 2 
setae on Abd. 1 . Other characters described 
under S. bipunctata identical in this species. 

Comments. — The specimen from locality 
2425 has three outer macrochaetae on both 
sides of Abd. 3; all other specimens possess 
two setae. I have examined two specimens 
from a cave in Puebla, Mexico (coll. no. 
4067) which possess the typical coloration 
of this species but their guts are filled with 
so much food that the abdominal chaeto- 
taxy cannot be studied. 

Diagnosis. — 5'^zra crypt ica is very similar 
to S. distincta Mari Mutt (1986), described 
from Puerto Rico and now reported in this 
paper from the United States (Massachu- 
setts) and Mexico. These species differ in 
coloration, the pigment in S. distincta is re- 
stricted to the sides of Th. 3 to Abd. 2 (Mari 
Mutt, 1986: Fig. 21). In addition, S. dis- 
tincta lacks the macrochaeta inserted above 
the pseudopore of Abd. 4 and consistently 
possesses three outer macrochaetae on Abd. 

3 (two in S. crypt ica). 

Seira cryptica can be distinguished from 
S. bipunctata by the presence of 2 + 2 mac- 
rochaetae on Abd. 1 (4 + 4 in 5". bipunc- 
tata) and also by the coloration, although 
well pigmented specimens oiS. cryptica may 
closely resemble specimens of S. bipunc- 
tata. Specimens of S. bipunctata possess a 
somewhat conical mesonotum which pro- 
jects over an elongated head. In S. cryptica 
the mesonotum does not project over the 
head and the head is rounded. Seira bi- 
punctata has an apical protuberance on Ant. 

4 (absent in S. cryptica) and always has 

three outer macrochaetae on Abd. 3 (usually 
two in S. cryptica). Finally, the posterior 
outer group of setae on Th. 2 is usually com- 
posed of six setae in S. bipunctata and of 
five in S. cryptica. 

Material examined. — Florida, Monroe 
County, on several small mangrove islands, 
4. VII. 1 969 to 1 6. VII. 1970, collection num- 
bers: 2425, 2432, 2433, 2506, 2827, 2831, 
2834; D. Simberloff, holotype and 17 para- 
types on slides. Arizona, nr. Phoenix, 
McDowell Mt. Park, on dry stream bed, 
1 9. III. 1986, coll. no. 6620, K. Christiansen, 
1 specimen on slide. 

Seira bipunctata (Packard) 

Lepidocyrtus bipunctatus Fackard 1873: 37 

Seira bipunctata— ChvisXiansen & Bellinger 

1980: 925-926, Fig. 756 A, C (Nebraska 

and New Mexico). 

Description. — Length from front of head 
to end of Abd. 6 up to 2.4 mm. Coloration 
variable (see comments). Anterior margin 
of mesonotum conical, projecting over head. 
Distribution of scales: head and body, dor- 
sum of Ant. 1 and Ant. 2, dorsal proximal 
half of Ant. 3, all leg segments, and furcula. 
Ant. 4 slightly annulated; its apex (Fig. 5) 
with distinct protuberance and bilobed pa- 
pilla but without pin seta. Head elongate, 
macrochaetae distributed as in Fig. 1 . Eyes 
G and H not greatly reduced. Interocular 
chaetotaxy consists of a small ciliated seta 
external to eye D, a similar seta between 
eyes E and F, and 5 longer setae in area 
between eyes C to H (arrangement identical 
in S. ca/2^«/ Jacquemart— Mari Mutt, 1986: 
Fig. 10). Prelabral setae ciliated, labral 
setae smooth. Labral papillae well 
developed, sometimes bifurcated apically 

Figs. 8, 9. Seira bipunctata. 8, Distribution of violet pigment, lectotype. 9, Ventral manubrial chaetotaxy, 
basal area of manubrium at top of page. Figs. 10-14. Seira cryptica. 10, Labral papillae. 11, Distribution of 
body macrochaetae, bothriotricha (wavy lines) and pseudopores (x). 1 2, Distribution of violet pigment. 1 3, Apex 
of Ant. 4. 14, Distribution of head macrochaetae. 



(Fig. 7). Labial chaetotaxy: al-a5,MlM2r 
(rcduced)ELlL2. One long ciliated seta on 
each side of ventral groove near posterior 
margin of head. One to 3 pairs of scales 
inserted between this seta and setae of post- 
labial quadrangle. Setae of maxillary palpus 
smooth, subequal in length and shape. Dif- 
ferentiated seta of outer labial papilla clearly 
surpassing apex of papilla (Fig. 6). Body 
macrochaetotaxy as in Fig. 2; 4 + 4 setae 
on Abd. 1. Detailed chaetotaxy of Abd. 2 
and Abd. 3 and distribution of setae asso- 
ciated with anterior bothriotricha of Abd. 
4 as in 5". cahcni {Man Mutt, 1986: Figs. 16- 
18). Trochanteral organ with up to 30 slen- 
der smooth setae. Structure of claws as in 
Fig. 4. Ventral manubrial chaetotaxy as in 
Fig. 9. Mucro as in Fig. 3. Male genital plate 
circinate, with 1 1 smooth setae in a circle 
around genital pore and at east 1 + 1 setae 
near pore. 

Comments.— The types of this species 
possess a small patch of pigment along the 
anterior margin of the mesothorax and a 
larger patch on the lateral-posterior margin 
of Abd. 4 (Fig. 8). On the other hand, the 
specimens from Nebraska are almost com- 
pletely pigmented as illustrated by Chris- 
tiansen and Bellinger ( 1 980: 926, Fig. 756C). 
The first two antennal segments of the latter 
specimens are violet but Ant. 3 and Ant. 4 
are pale. Legs are violet to the apical '/» of 
femur, remainder of leg is pale yellow. The 
specimens from New Mexico have all the 
antennal segments, head, and thorax pig- 
mented but the abdomen lacks color. No 
anatomical differences were detected among 
these populations. 

Setae signalled by arrows on Th. 2 and 
Abd. 4 in Fig. 2 are absent on one side of 
the body of one specimen from Nebraska. 
The signalled seta on Abd. 3 is absent on 
both sides of the body of one specimen from 
the same locality. 

Diagnosis.— This species is similar to S. 
cahcni Jacquemart (1976). described from 
the Galapagos Islands and reported from 
Cuba (Gruia, 1983) and Puerto Rico (Mari 
Mutt, 1986). Both species possess identical 

head and body macrochaetotaxy except that 
5'. bipunctata has six setae on the outer pos- 
terior group of Th. 2 (Fig. 2) while most 
specimens of Jacquemart's species have five 
setae (some specimens have six setae). Seim 
cahcni lacks an apical protuberance on Ant. 
4 and has a pair of setae between the apical 
pair on the manubrium of S. bipunctata (cf. 
Fig. 9 and Mari Mutt, 1986: Fig. 1 1). These 
species also differ in typical coloration. 

Material examined. — Texas, McLennan 
County, Waco, Lectotype (here designated), 
4 paralectotypes on slides and 35 paralec- 
totypes in alcohol; Nebraska, Thomas 
County, Halsey, 13.n.l957, coll. no. 46 Ix, 
Henzlik, col., 4 specimens on slides; New 
Mexico, Eddy County, Carlsbad Caverns 
National Park, bat cave site no. 4, 
25. IV. 1975, coll. no. 3668, 3 on a slide with 
specimens of an Entomobrya. 

Seim distincta Mari Mutt 

Seira distincta Mari Mutt, in press. 

Described originally from Puerto Rico, 
this species is reported for the first time from 
the United States and Mexico. Seira dis- 
tincta is very similar to S. cryptica\ the dif- 
ferences between both are listed in the lat- 
ter's diagnosis. 

Material examined. — USA, Massachu- 
setts, Cambridge, Biological Laboratories, 
26.11.1948, K. Christiansen, coll. no. 5985, 
4 specimens on slides; Mexico, Veracruz, 
Cueva del Rio, 3 km E of Atoyac, 6. II. 1 957, 
coll. no. 4093, J. Reddell, 1 specimen on 

Literature Cited 

Christiansen, K. and P. Bellinger. 1980. The Collem- 
bola of North America north of the Rio Grande, 
a taxonomic analysis. Part 3. Family Entomobryi- 
dae. Grinnell College, Iowa. pp. 785-1042. 

Gruia, M. 1983. Collemboles arthropleones de Cuba 
recoltes par les expeditions cubano-roumaines en 
1969-1973, II. Result. Exped. Biospeleol. Cu- 
bano-Roumaines Cuba 4: 191-205. 

Jacquemart, S. 1976. Collemboles nouveaux des lies 
Galapagos. Mission zoologique beige aux lies Ga- 
lapagos et en Ecuador (N. et J. Leleup, 1 964- 1 965) 
3: 137-157. 



Mari Mutt, J. A. In press. Puerto Rican species of 

Seira. Carib. J. Sci. 22(3-4). 
Packard. A. S. 1873. Synopsis of the Thysanura of 

Essex County, Mass., with descriptions of a few 
extralimital forms. Rep. Peabody Acad. Sci. 5: 23- 

89(1), 1987, p. 131 


Brachymeria discretoidea, a new junior synonym of Brachymeria discreta 
(Hymenoptera: Chalcididae) 

Brachymeria (Gahanula) discreta Gahan 
and Brachymeria {Gahamda) discretoidea 
Gahan were described in 1942 (Proc. U.S. 
Natl. Mus. 92: 43-44) and considered to be 
similar to one another. Brachymeria dis- 
creta was based on specimens from Mexico, 
discretoidea on specimens from Panama. 
Brachymeria discreta was later recorded 
from Texas, Arizona, Southern California, 
and Hawaii, and discretoidea from Arizona, 
Texas, and Mexico (Burks, 1979. Chalcid- 
idae, pp. 860-874. In Krombein, K. V. et 
al., eds., Catalog of Hymenoptera in Amer- 
ica North of Mexico. Vol. I. Smithson. Inst. 
Press., Wash., D.C.). Brachymeria discreta 
has been reared from the nest of Polistes 
instabilis Sauss. (Hymenoptera: Vespidae), 
probably from a pyraustid moth that in- 
fested the wasp nest, and from a species of 
Tinea (Lepidoptera: Tineidae) breeding in 
chicken droppings. Brachymeria discretoi- 
dea has been reared from the nest of Trigona 
amalthea (Oliv.) (Hymenoptera: Apidae) 
infested by an unidentified moth (Gahan, 
ibid.; Burks, 1960, Trans. Am. Entomol. 
Soc. 86: 225-273). 

Burks (1960) indicated difficulty in de- 
termining material as either discreta or dis- 
cretoidea and that the differences noted by 
Gahan tended to intergrade. He chose not 
to synonomize the two species and indicat- 
ed that, except for the characters in his key 
("frontal carina always well developed ver- 
sus weak-virtually absent; ocellocular line 
-/s versus y^ as long as diameter of lateral 
ocellus; marginal vein 4 versus 3V2 times as 
long as postmarginal; propodeum with an 

elongate-median areola versus lacking"), the 
two species were alike in color and struc- 

Upon examining the types of both species 
(in the U.S. Museum of Natural History, 
Washington, D.C., Nos. 55149 and 55150) 
and additional material from various lo- 
calities, I have concluded that the material 
represents one species, that the color differ- 
ences and intergrading charcters of Gahan 
and Burks merely represent intraspecific 
variation. I therefore consider discretoidea 
a junior synonym of discreta. 

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 Office, Vi- 
salia, California for editorial comments on 
this paper. I thank also the following insti- 
tutions and their personnel for the oppor- 
tunity to have examined their material of 
B. discreta: University of California, Davis; 
California Department of Food and Agri- 
culture, Sacramento; Florida Department of 
Agriculture and Consumer Affairs, Gaines- 
ville; Bishop Museum, Honolulu, Hawaii; 
National Museum of Natural History, 
Washington, D.C.; Systematic Entomology 
Laboratory, USDA, Washington, D.C.; and 
the Fresno County Agricultural Commis- 
sioner's Office, Fresno, California. 

Jeffrey A. Halstead, 21 JON. Hayes, Fres- 
no, California 93722. 


89(1), 1987, pp. 132-136 




Charles E. Turner, Robert W. Pemberton, and Sara S. Rosenthal 

Biological Control of Weeds, U.S. Department of Agriculture, Agricultural Research 
Service, Western Regional Research Center, Albany, California 94710. 

Abstract. —Seventeen new host records are reported for the artichoke plume moth Pla- 
typtilia carduidactyla (Lepidoptera, Pterophoridae) from native and naturalized thistles 
(Asteraceae, Cardueae) in California. All are adult rearing records from field populations 
of the host thistles. The number of moths reared and the amount of host material collected 
for rearing are reported. The new records include the naturalized species Arctium minus 
and Carduus pycnocephalus, and 1 5 native Cirsium species: Cirsium andersonii, C. an- 
drewsii, C. brevistylum, C. californicum, C. campylon, C. ciliolatum, C. crassicaule, C. 
cymosum, C. douglasii, C. fontinale obispoense, C. hydrophilum vaseyi, C. loncholepis, C. 
pastoris, C. rhothophilum, and C. tioganum. Seven of the newly reported Cirsium hosts 
are considered rare. Previous host records are also summarized. With one exception, all 
known hosts are in the thistle subtribe Carduinae. 

The so-called artichoke plume moth, Pla- 
typtilia carduidactyla (Riley) (Lepidoptera, 
Pterophoridae) is the most serious pest of 
introduced cultivated globe artichoke {Cyn- 
ara scolymus L., Asteraceae) in California 
(Haynes et al, 1981; Lange, 1 94 1 ; Lange et 
al., 1954). This moth, which is native and 
widespread in North America, is muitivol- 
tine and feeds on leaf, stem, and flowerhead 
tissue of its host plants (Lange, 1950). The 
thistle tribe (Asteraceae, Cardueae) is an im- 
portant group of native species, naturalized 
weeds, and the cultivated crops safflower 
(Carthamus tinctorius L.) and globe arti- 
choke (Moore and Frankton, 1 974; Ownbey 
et al., 1975; Reed, 1970; Robbins et al., 
1970; USDA, SCS, 1982). Our study of the 
host pattern of the plume moth is part of a 
broader entomofaunal study of the thistle 
tribe in CaUfomia (Pemberton et al., 1985; 
Turner et al., in press) as it relates to bio- 
logical control of weedy naturalized thistles. 

The previously reported host records for 
P. carduidactyla are as follows: Centaurea 
melitensis L. (Lange, 1941, 1950); Cirsium 
arvense (L.) Scop. (Lange, 1950); C calli- 
lepis (Greene) Jeps. (reported as C. ameri- 
canum Daniels var. callilepis Jeps.) (Lange, 
1950); C discolor {M\M. ex Willd.) Spreng. 
(Marcovitch, 1916); C. edule Nutt. (possi- 
bly confused with C brevistylum Cronq.) 
(Lange, 1941, 1950); C. occidentale {^\xXi) 
Jeps. (Lange, 1941, 1950); C proteanumh 
T. Howell (reported as C. occidentale var. 
venustum Jeps.) (Lange, 1941, 1950); C. 
quercetorum (Gray) Jeps. (Lange, 1941, 
1950); C undulatum (Nutt.) Spreng. (Lange, 
1941, 1950); C. vulgare (Savi) Tenore (re- 
ported as C. lanceolatum (L.) Scop.) (Riley, 
1869; Lange, 1941, 1950); Cynara cardun- 
culus L. (Lange, 1941, 1950); C scolymus 
L. (Essig, 1922; Lange, 1941, 1950); Sily- 
bum marianum (L.) Gaertn. (Essig, 1922). 
Where we use a different host name from 



that originally reported, this was done in 
accordance with more recent treatments of 
the genus Cirsium (Jepson, 1925; Munz, 
1973). There has been confusion between 
C brevistylum and C edule, which are 
closely related and similar (Moore and 
Frankton, 1962). It is doubtful that true C. 
edule occurs in California (Moore and 
Frankton, 1962; Munz, 1973). Of these pre- 
viously reported hosts, explicit mention of 
emerged adults from host material is only 
made for C discolor, C. vulgare, and C. 
scolymus. Of the previously reported hosts, 
C. callilepis, C. discolor, C. edule, C. occi- 
dentale, C. proteanum, C. quercetorum, and 

C. undulatum are native to North America 
(USDA, SCS, 1982). 

All the new host records reported here are 
from material collected and reared by the 
authors except for the records prior to 1 982, 
which are from material collected and reared 
by K. E. Frick and R. B. Hawkes, formerly 
affiliated with our Laboratory. Host mate- 
rial collected in 1982 consisted of periodic 
samples of ten whole plants from the same 
field populations of native and naturalized 
Carduus2Ln(^ Cirsium species. In 1983, 1984, 
and 1985 we sampled the flowerheads only 
of field populations of native and intro- 
duced species o{ Arctium, Carduus, Cartha- 
mus, Centaurea, Cirsium, Cynara, Onopor- 
dum, Saussurea, and Silybum. We collected 
lateral and terminal flowerheads at a stage 
between flowering and seed dissemination. 
All of the introduced species sampled were 
from naturalized populations except for 
samples from planted fields of artichoke and 
safflower. The host material was brought 
back to the lab for the rearing of adult plume 
moths and other thistle insects (Pemberton 
et al., 1985; Turner et al., in press). 

D. C. Ferguson and W. H. Lange, Jr. de- 
termined the plume moths. The authors, G. 

D. Barbe, T. C. Fuller, and G. B. Ownbey 
determined the thistles. Voucher specimens 
of most plume moths and host thistles are 
retained in our lab. 

We report here 17 new host records for 

P. carduidactyla from native and natural- 
ized thistle species, including information 
on host location and sample date, number 
of flowerheads sampled, and the number of 
emerged adults. The new host records are 
from the relatively common native thistles 
Cirsium andersonii (Gray) Petrak, C an- 
drewsii (Gray) Jeps., C brevistylum Cronq., 
C. californicum Gray, C. cymosum (Greene) 
J. T. Howell, C. douglasii DC, C pastoris 
J. T. Howefl, and C tioganum Congd.; and 
from the relatively rare (Federal Register, 
1985) native thistles C. campylon H. 
Sharsm., C ciliolatum (L. Henders.) J. T. 
Howell, C crassicaule (Greene) Jeps., C 
fontinale (Greene) Jeps. var. obispoense J. 
T. Howell, C hydrophilum (Greene) Jeps. 
var. vaseyi (Gray) J. T. Howell, C loncho- 
lepis Petrak, and C rhothophilum Blake. 
New host records also are from the natu- 
ralized thistles Arctium minus Bemh. and 
Carduus pycnocephalus L. 

These newly reported host species occur 
in a wide variety of habitats located 
throughout cismontane California (Munz, 
1973). The number of emerged adult plume 
moths was generally smaller than the num- 
ber of larvae and pupae associated with the 
host plants. This diflerence in number be- 
tween adult versus immature plume moths 
could have been due to insect parasitoids 
(Lange, 1950) or to our rearing conditions. 
We also reared adult plume moths from C. 
occidentale, C proteanum, C quercetorum, 
C. undulatum, and C. vulgare among the 
hosts previously reported. 

Native Cirsium species are the only known 
native North American hosts for P. car- 
duidactyla. In North America Cirsium con- 
tains ca. 130 native species (Moore and 
Frankton, 1974; Ownbey et al., 1975), with 
ca. 30 species native to California (Munz, 
1973). The moth is now known from 21 
Cirsium species (excluding C. edule) native 
to North America, and 1 9 species native to 
California. Ironically, the plume moth was 
first described from specimens reared from 
naturalized Cirsium vulgare in Missouri 



(Riley, 1 869). Riley ( 1 869) used the perhaps 
more appropriate colloquial name, thistle 
plume moth, for P. carduidactyla. 

Globe artichoke has been grown com- 
mercially in California at least since 1900 
(Stokdyk, 1932). P. carduidactyla has been 
a pest of globe artichoke in California at 
least since 1922 (Essig, 1922; Lange, 1941). 
Both Cirsium and Cynara are in the sub- 
tribe Carduinae (Dittrich, 1977), which in- 
dicates that they are relatively closely re- 
lated. The plume moth is a native 
stenophagous insect that is capable of host 
utilization of an introduced crop plant that 
is relatively closely related to its native hosts. 

All known host taxa are in the thistle tribe 
Cardueae; and all known hosts are in the 
subtribe Carduinae with the exception of 
Centaurea melitensis, which is in the sub- 
tribe Centaureinae (Dittrich, 1977). Lange 
(1950) reported a larval feeding record of 
P. carduidactyla from a C. melitensis pop- 
ulation adjacent to a planting of globe ar- 
tichoke that had the moth. We did not rear 
P. carduidactyla from five samples of C 
melitensis, or from any of the 14 samples 
from 3 other species of Centaurea that we 
studied (unpublished data). 

New Host Records 

Arctium minus. Dimmick State Park, Men- 
docino Co., CaHf.: host plants coll. 6-IX- 

62 by K. E. Frick, 9 adults emerged by 

Carduus pycnocephalus. 2.5 km N. of Napa, 
Napa Co., Calif.: host plants coll. 15-V- 

63 by R. B. Hawkes, 1 adult emerged on 

Cirsium andersonii. Gumboot Rd., 1 9.0 km 
from Castle Lake Rd., Siskiyou Co., Cal- 
if.: 108 host heads coll. 22-VIII-84, 1 
adult emerged by 25-IX-84. 

Cirsium andrewsii. Abbott's Lagoon, Pt. 
Reyes, Marin Co., Calif: host plants coll. 
ll-V-82, 2 adults emerged by 5-VI-82; 
host plants coll. l-VI-82, 1 adult emerged 
from head by 22-VI-82; host plants coll. 

13-VII-82, 5 adults emerged from heads 
by 17-XI-82; host plants coll. 3-VIII-82, 
3 adults emerged from heads by 1 1 -VI- 
83; 100 host heads coll. 22-VI-85, 1 adult 
emerged by 5-XI-85. 

Cirsium brevistylum. ca. 1 4 km W. of Wil- 
lits, Mendocino Co., Calif.: host plants 
coll. 25-VII-61 by K. E. Frick, 2 aduhs 
emerged by 4-VIII-61; host plants coll. 
2-VII-62 by K. E. Frick, 20 adults 
emerged by 23-VII-62. Crescent City, Del 
Norte Co., Calif: host plants coll. 16- 
VIII-6 1 by K. E. Frick, 5 adults emerged 
by 5-IX-61. Mt. Tamalpais, Marin Co., 
Calif.: host plants coll. 5-VII-62 by K. E. 
Frick, 3 adults emerged. (Additional later 
host records from Del Norte and Men- 
docino counties coll. by K. E. Frick avail- 
able from the authors.) Abbott's Lagoon, 
Pt. Reyes, Marin Co., CaHf.: 100 host 
heads coll. 1 7-V-84, 3 adults emerged by 
l-VIII-84; 100 host heads coll. 5-VII-84, 
3 adults emerged by 14-VIII-84. 
MacKerricher Beach, Mendocino Co., 
CaHf: 89 host heads coH. lO-VII-84, 17 
adults emerged by 14-VIII-84. 

Cirsium californicum. Hwy. 49 between 
Downieville and Sierra City, Sierra Co., 
Calif.: 100 host heads coH. 21-VI-84, 1 
aduh emerged by 31-VII-84. 

Cirsium campylon. Blackbird Valley off 
Mines Rd., Santa Clara Co., Calif.: host 
plants coll. 8-VI-82, 1 adult emerged from 
head by 30- VI-82; host plants coH. 1 -VII- 
82, 1 aduh emerged from head by 21- 
VII-82; host plants coH. 31-VIII-82, 2 
adults emerged from head by 17-XI-82. 
Mines Rd., 11.6 km S. of Alameda— San- 
ta Clara county line, Santa Clara Co., 
Calif.: 100 host heads coU. 30-VI-85, 1 
adult emerged by 5-XI-85. 

Cirsium ciliolatum. York Rd. at Mulloy Rd., 
Siskiyou Co., Calif.: 100 host heads coU. 
27-VI-84, 1 aduh emerged by 30-VII-84. 

Cirsium crassicaule. Kern Nat. Wildlife 
Refuge, Kern Co., CaHf.: 60 host heads 
coH. 23-VI-85, 1 aduh emerged by 5-VII- 



Cirsium cymosum. Callahan — Cecil ville 
Rd., 8.0 km from Callahan, Siskiyou Co., 
CaUf.: 67 host heads coll. 5-VIII-83, 1 
adult emerged by 22-XII-83. 

Cirsium douglasii. Callahan— Cecil ville Rd., 
8 km from Callahan, Siskiyou Co., Calif.: 
host plants coll. 27-VII-82, 1 adult 
emerged from heads by 18-VIII-82; host 
plants coll. 17-VIII-82, 1 adult emerged 
from heads by 1 7-XI-82. Hwy. 3, 3.9 km 
S. of Scott Mt. Pass, Trinity Co., Calif.: 
100 host heads coll. 4-VIII-84, 2 aduhs 
emerged by 3-IX-84. 

Cirsium fontinale obispoense. San Simeon 
Cr. Rd., 8.4 km from Hwy. 1, San Luis 
Obispo Co., Calif: 100 host heads coll. 
26-V-84, 1 aduh emerged by 31-VII-84. 

Cirsium hydrophilum vaseyi. West Point Inn 
trail, Mt. Tamalpais, Marin Co., Calif.: 
100 host heads coll. 22-VI-85, 1 adult 
emerged by 5-XI-85. 

Cirsium loncholepis. Guadalupe Dunes, San 
Luis Obispo Co., Calif.: 60 host heads 
coll. 14-VI-83, 1 aduh emerged by 22- 

Cirsium pastoris. Callahan— Cecil ville Rd., 
2-8 km from Callahan, Siskiyou Co., 
CaUf : host plants coll. 7-VII-82, 1 aduh 
emerged from stem by 21-VII-82; 100 
host heads coll. 9-VIII-84, 1 adult 
emerged by 4-X-84. Slough Rd., S. of 
Louie Rd. exit from 1-5, Siskiyou Co., 
Calif: 100 host heads coll. 29-VI-83, 3 
adults emerged by 1 -X-83. Edgewood exit 
from 1-5, 5.6 km N. of Weed, Siskiyou 
Co., CaUf: 100 heads coll. 9-VIII-84, 3 
adults emerged by 4-X-84. 

Cirsium rhothophilum. Guadalupe Dunes 
nr. Oso Flaco Lake, San Luis Obispo Co., 
Calif.: 42 host heads coll. 15-VI-83, 2 
adults emerged by 5-VII-84. 

Cirsium tioganum. Willow Cr. Rd., Siski- 
you Co., Calif.: 82 host heads coll. 12- 
VII-84, 1 aduh emerged by 30-VII-84. 
Dorris Brownell Rd., nr. jet. with Willow 
Cr. Rd., Siskiyou Co., Calif: 100 host 
heads coll. 18-VII-85, 9 adults emerged 
by 6-XI-85. 


K. F. Haynes and D. MacNeill reviewed 
the manuscript. D. C. Ferguson (Systematic 
Entomology Laboratory, Agricultural Re- 
search Service, U.S. Department of Agri- 
culture) and W. H. Lange, Jr., helped de- 
termine the P. carduidactyla. G. D. Barbe, 
T. C. Fuller, and G. B. Ownbey helped de- 
termine the thistle species. R. Colville and 
M. T. Johnson, Jr., helped with the rearing 
and processing of insects, and the latter also 
helped with the sampling of thistleheads. 

Literature Cited 

Dittrich, M. 1977. Cynaraea-systematic review, pp. 
999-1038. In Heywood, V. H., J. B. Harbome, 
and B. L. Turner, eds., The biology and chemistry 
of the Compositae, 2 vols. Academic, New York. 

Essig, E. O. 1922. The artichoke plume moth. Calif. 
Dept. of Agric. Monthly Bull. 11: 454-456. 

Federal Register. 1985. Endangered and threatened 
wildlife and plants; review of plant taxa for listing 
as endangered or threatened species; notice of re- 
view. Federal Register 50: 39526-39584. 

Haynes, K. F., M. C. Birch, and J. A. Klun. 1981. 
Sex pheromone offers promise for control of ar- 
tichoke plume moth. Calif Agric. 35: 13-14. 

Jepson, W. L. 1925. A manual of the flowering plants 
of California. Univ. of California, Berkeley. 

Lange, W. H., Jr. 1941. The artichoke plume moth 
and other pests injurious to the globe artichoke. 
Calif Agr. Exp. Sta. Bull. 653. 

. 1950. Biology and systematics of plume moths 

of the genus Platyptilia in California. Hilgardia 
19: 561-668. 

Lange, W. H., R. H. Sciaroni, and A. S. Greathead. 
1 954. Artichoke plume moth damage. Calif Agric. 
8(7): 7-8, 12. 

Marcovitch, S. 1916. Insects attacking weeds in Min- 
nesota. Minn. State Entomol. 16th Report: 135- 

Moore, R. J. and C. Frankton. 1962. Cytotaxonomy 
and Canadian distribution of Cirsium ediile and 
Cirsium brevistylum. Can. J. Bot. 40: 1 187-1 196. 

. 1974. The thistles of Canada. Research 

Branch, Canada Dept. Agriculture, Monograph No. 

Munz, P. A. 1973. A California flora and supplement. 
Univ. of California, Berkeley. 

Ownbey, G. B., P. H. Raven, and D. W. Kyhos. 1975. 
Chromosome numbers of some North American 
species of the genus Cirsium. III. Western United 
States, Mexico, and Guatemala. Brittonia 27: 297- 



Pemberton, R. W., C. E. Turner, and S. S. Rosenthal. 
1985. New host records for tephritid flies from 
Cirsium and Saussurea thistles in California. Proc. 
Entomol. Soc. Wash. 87: 790-794. 

Reed, C. F. 1970. Selected weeds of the United States. 
U.S. Dept. of Agric, Agric. Hdbk. No. 366. 

Riley, C. V. 1 869. First annual report on the noxious, 
beneficial, and other insects of the state of Mis- 
souri, pp. 180-181. State of Missouri, Jefferson 

Robbins, W. W., M. K. Bellue, and W. S. Ball. 1970. 
Weeds of California. Calif Dept. Agric, Sacra- 

Stodyk, E. A. 1932. Marketing globe artichokes. Cal- 
if Agric. Exp. Sta. Bull. 524. 

Turner, C. E., R. W. Pemberton, and S. S. Rosenthal. 
Host utilization of native Cirsium thistles (Aster- 
aceae) by the introduced weevil Rhinocyllus coni- 
cus (Coleoptera: Curculionidae) in California. En- 
viron. Entomol. (In press.) 

U.S. Dept. Agriculture, Soil Conservation Service. 
1982. National list of scientific plant names. Vol. 
1 . List of plant names. 


89(1), 1987, pp. 137-140 


T. Michael Peters 

Professor and Curator, Department of Entomology, University of Massachusetts, Am- 
herst, Massachusetts 01003. 

Abstract.— Dixa adleri, new species, from Pennsylvania is described with illustrations 
of the mature larva, male genitalia, and female bursa copulatrix. Adults are compared 
with other known Nearctic dixids. Larval habitats of dixid genera are compared. 

Larval Dixidae live in or just above the 
meniscus in a wide variety of freshwater 
habitats, from stagnant bogs to rushing 
brooks. The three Nearctic genera have been 
associated in the literature with distinct types 
of freshwater environments, differences that 
have been correlated with unique anatom- 
ical characteristics of each of the genera. 
However, comparison of environmental 
data available for a new species of Dixa 
found in eastern United States with those 
of my previous collections, revealed some 
anomalies. Following the description of the 
new species is a discussion of overlap in 
larval habitats of the two most widely dis- 
tributed genera; Dixa Meigen and Dixella 
Dyar and Shannon. 

Dixa adleri Peters, New Species 

Figs. 1-4 

Xd\i\\.—Head: Medium brown in speci- 
mens preserved in alcohol; without micro- 
trichia; a line of 14 long setae along periph- 
ery of compound eye from dorsum of vertex 
to venter of eye; frontoclypeus with 2 or 3 
setae near ventral edge; antennae concol- 
orous with head, first flagellomere subcylin- 
drical, width:length 1:10. Thorax: Medium 
brown in mature specimens, but with dis- 
tinctly darker vittae on scutum, vittae in- 
distinguishable in teneral individuals; an- 
terior pronotum of male with 6-8 setae (9- 

11 in 9) as long as width of sclerite; 1 seta 
near dorsal suture on posterior pronotum; 
scutellum of males with a transverse row of 
1 1 setae, the central one with another pos- 
terior to it, scutellum of females with an 
anterior transverse row of 1 3 setae, followed 
by a transverse row of 3 setae and followed 
by a single mesal seta. Wing: Clear, without 
pigmented areas, length 3.25-3.69 mm 
(<5)(3. 69-4.08 in 9); in 3 M3 + 4:M1 + 2 
as 1:1.4-1.6, M3 + 4:Mst as 1:1.5-2.5; 
R2 + 3:R3 as 1:2. 1-3.1; in 9, vein ratios are 
within same range as those of 5. Crossvein 
m-cu broken. Halter: Hyaline. Legs: Distal 
spiniform seta on 3rd tarsomere of foreleg 
(on tarsomeres 3-4 of 1 9), on 1-3 of midleg 
and on 1-4 of hindleg; basal recurved spi- 
niform seta on tarsomere 5 of fore and mid- 
leg of (3; claws simple in 9, in $ hind claw 
simple, fore and midclaws with 5-6 ventral 
teeth (3-4 long, 1 short, 1 long); femuntibia: 
tarsus length ratios of forelegs as 1:0.96- 
1.00:1.50-1.90 in <5, in 9 1:0.96-1.00:1.40- 
1.48; midleg 1:0.92-1.05:1.13-1.74 in 3, in 
9 1:1.00-1.05:1.32-1.41; hindleg 1:1.00- 
1.05:1.54-1.74 in <5, in 9 1:1.14-1.21:1.67- 
1.95. Abdomen: Mottled brown and grey, 
mixed lighter and darker areas; in 3, 9th 
stemite widest laterally, wider than the more 
lightly sclerotized tergite 9; tergite 10 with 
non-segmented cerci (Fig. 1); gonocoxite and 
gonostylus as in Figs. 1 and 2; ejaculatory 



Figs. 1-4. Dixa adleri. 1, Lateral view of male terminalia. 2, Dorsal view of same. 3, Bursa copuiatrix. 4, 
Posterior abdominal structures of fourth instar larva; left half dorsal, right half ventral. 

duct very short and very lightly sclerotized; 
claspette and penis valves not discemable 
in cleared glycerin mount, even under in- 
terference microscopy; 5 with three tufts of 
setae in bursa copuiatrix (Fig. 3). 

Fourth instar larva. — Length 6.6-9.0 mm 
(n = 8); coloration of specimens available is 
bleached out, probably due to preservation 
techniques. Terminology follows Peters and 
Adamski (1982). Head: Seta 10 about Vi 



length of 9, length of 7 intermediate between 
them; head seta 1 long, wide; seta 2 slender 
but longer than 1, 3 very short and wide, 
widest at middle. Antenna: Tip with several 
sharp spines surrounding a larger, pear- 
shaped, thin-walled sensillum. Thorax: An- 
terior ventral setae of prothorax not ex- 
tending beyond mouth brushes, arranged as 
1 - 1 -4- 1 - 1 -4- 1 - 1 , each group of 4 without 
common sclerotized base. Abdomen: Pro- 
legs of segments 1 and 2 with uniordinal, 
biserial crochets of subequal length, anterior 
proleg preceded by row of spicules subequal 
in length to crochets, 25 crochets in first 
row, second row thickest, with 24 crochets; 
posterior proleg with 21 crochets per row, 
posterior row thickest, followed by an un- 
even line of thin spicules which are subequal 
in length to crochets; corona of branched 
spicules on dorsum of segments 2-7; am- 
bulatory combs on venter of segments 5-7 
each with 9-10 spines on either side of a 
medial subrectangular plate, spines in 2 dis- 
tinct rows of alternating heavy and more 
slender spines, heavier ones projecting at 
less acute angle from surface of segment; 
segment 8 with only 6 slender setae on ven- 
ter, shorter than length of segment; paraspi- 
racular setae, postspiracular process, and 
metaspiracular plate as in Fig. 4; segment 9 
with fringe of setae continuous from pos- 
terolateral process anterior to postspiracu- 
lar process, median plate as in Fig. 4, pecten 
of anterolateral plate simple, as in Fig. 4; 
ventrolateral plate of segment 1 with a thick 
seta directed posteriorly, another arising near 
its base, is much more slender and directed 
laterally, anterior to the plate is another more 
dorsal hair; caudal hairs of postanal process 
over 2 x length of process. 

Specimens examined. — Holotype <5, Slab 
Cabin Run, Pine Grove Mills, Centre Co., 
Pennsylvania, 18-IX-81, collected by Peter 
H. Adler with Malaise trap over stream; 
deposited in the Peters dixid collection. De- 
partment of Entomology, University of 
Massachusetts, Amherst, Massachusetts. 
Eighteen paratype 7 5 and 1 1 2, same data 
as holotype, with associated immature skins. 

Additional sites, all in Pennsylvania: 29 
specimens from Scott Road (Slab Cabin 
Run), State College, and Briesly. 


If keyed using Peters and Cook (1966), 
mature D. adleri males come out as Dixa 
fratema Garrett. They may be separated on 
the basis of the ejaculatory duct: it is large 
and heavily sclerotized in D. fratema, but 
scarcely visible in D. adleri. Teneral males 
without distinctly darker scutal vittae key 
to D. inextricata but have more than 5 setae 
on the anterior pronotum. Females key to 
D. fluvica but may be separated by details 
of the bursa copulatrix. The three tufts of 
setae each possesses are subequal in length 
in D. fluvica, with the median group con- 
sisting of 3-4 setae. In D. adleri the median 
group is distinctly shorter than the other two 
and consists of 5-6 setae. All setae in the 
bursa copulatrix of D. adleri are very stout. 
They are slender in D. fluvica. 

Collection sites for Dixa adleri are char- 
acterized in Adler et al. (1983) and Adler 
and Kim (1984). Adults were collected with 
a Malaise trap erected over the stream. Lar- 
vae were collected with drift nets. Several 
adults with associated larval and/or pupal 
skins were reared by Dr. Adler after whom 
the species is named. 

In his lengthly discussion of larval habi- 
tat, Nowell (1952) observed that Dixa is a 
fast water form "always found out in the 
center of streams where the water is swiftest 
. . .," Meringodixa inhabits "quieter, 
smoother streams . . . congregating in quiet 
niches, or quiet pools ..." while Dixella is 
"in two habitats [it] may be in swiftly run- 
ning streams or in very still waters." He 
continued, "This is the only one of the 
groups which is represented in the quiet 
pools." These observations were based on 
California dixids and fit well the anatomical 
differences among larvae of each of the gen- 
era. Dixa has coronae of branched hydro- 
fuge hairs on abdominal segments 2-7, while 
Dixella has no flotation structures other than 
those on the posterior that surround the spi- 



racks in all known dixid larvae. Meringo- 
dixa has only the posterior flotation struc- 
ture, but it has long lateral tufts of hairs on 
abdominal segments 2-6 that are reported 
to aid in flotation (Nowell, 1951). 

While my own observations of Dixa and 
Dixella from other parts of the U.S. gen- 
erally agree with Nowell's, the actual situ- 
ation is more complex. Average current for 
my field collections is measured by timing 
a half-filled plastic vial over a measured dis- 
tance. Dixa was collected from moving 
waters in 8 of 12 instances, with average 
current of 1.83 ft/s (range = 0.67-3.5 ft/s). 
Dixella was collected from predominantly 
still water— ponds, bogs, pools— in 14 of 23 
instances, with average current in the other 
collections of 0.94 ft/s (range of 0.1-3.0 ft/ 
s). An additional difference is that Dixa 
sometimes (4 of 12 instances) occurs where 
"emergent" vegetation is lacking, but Dix- 
ella, with one exception noted below, al- 
ways (23 of 23 instances) is found where 
vegetation emerges from the water or vege- 
tation on the bank (grasses, roots) hangs into 
the water. In six collections, where both Dixa 
and Dixella were present, four of six sites 
were still water, but current in the other two 
ranged from 1.0-2.6 ft/s. In one site, both 
genera were taken from rocky banks; all oth- 
ers had "emergent" vegetation. 

Larval habitats of Dixa and Dixella ap- 
parently overlap to a great extent, with Dixa 
predominant in rapidly moving water and 
Dixella more common in still water. Dixa 
with its 6 coronae of branched, hydrofuge 

spicules should more effectively stay affoat 
in rapid water if caught in the current or 
undergoing a diel drift (Waters, 1962). Thus, 
Dixa has an anatomical advantage in rap- 
idly moving water, but is not restricted to 
this environment. Dixella occurs in both 
moving and still water, but usually is found 
in areas with less rapid current than Dixa. 
They do occur together in both habitats, 
suggesting that larval habitat is controlled 
by two factors, maternal selection of ovi- 
position site and sites available to the ovi- 
positing female. 


This research was partially supported by 
Hatch project #541 of the Massachusetts 
Agricultural Station. 

Literature Cited 

Adler, P. H. and K. C. Kim. 1984. Ecological char- 
acterization of two sibling species, IIIL-1 and IS- 
7, in the Simulium vittatum complex (Diptera: 
Simuliidae). Can J. Zool. 62: 1308-1315. 

Adler, P. H., R. W. Light, and K. C. Kim. 1983. The 
aquatic drift patterns of black flies (Diptera: Sim- 
uliidae). Hydrobiologica 107: 183-191. 

Nowell. W. R. 1951. The dipterous family Dixidae 
in Western North America (Insecta:Diptera). Mi- 
croentomol. 16: 187-270. 

Peters, T. M. and D. Adamski. 1982. A description 
of the larva of Dixella nova (Walker)(Diptera:Dix- 
idae). Proc. Entomol. Soc. Wash. 84: 521-528. 

Peters, T. M. and E. F. Cook. 1966. The Nearctic 
Dixidae (Diptera). Misc. Publ. Entomol. Soc. Am. 
5: 231-278. 

Waters, T. F. 1962. Diurnal periodicity in the drift 
of stream invertebrates. Ecology 43: 316-320. 


89(1), 1987, pp. 141-146 


B. P. Stark and B. C. Kondratieff 

(BPS) Department of Biology, Mississippi College, Clinton, Mississippi 39058; (BCK) 
Department of Entomology, Colorado State University, Fort Collins, Colorado 80523. 

Abstract.— k new species, Peltoperla tarteri Stark and Kondratieff, from Virginia and 
West Virginia is described. This new species was previously confused with P. arcuata 
Needham, but males lack the peculiar, basally curved cerci of that species. Both species 
inhabit springs and springbrooks of the central Appalachians but there are no known co- 

The Nearctic genus Peltoperla presently 
includes a single eastern species, P. arcuata 
Needham (Stark and Stewart, 1981). This 
species was described from a female adult 
collected in Ithaca, New York (Needham, 
1905). Twenty years later Needham and 
Claassen (1925) described the male from 
two specimens taken in Ithaca and at Pres- 
ident (Venango County), Pennsylvania. The 
nymph of this species was reared and de- 
scribed by Claassen (1931), also from Ith- 
aca. Prison (1942) redescribed the male in 
order to note the unusual cereal shape, pre- 
viously not emphasized. 

During a study of Nearctic peltoperlid 
genera by Stark and Stewart (1981), speci- 
mens lacking this peculiar cereal character 
were examined from Virginia and West Vir- 
ginia. These specimens were used to illus- 
trate the male genitalia for the genus Pel- 
toperla in the above paper. A study of these 
and additional material indicates that these 
specimens actually represent an unde- 
scribed species. Terminology used in this 
paper follows Stark and Stewart (1981). 

The following museums, institutions and 
individuals provided material for study: R. 
W. Baumann, Brigham Young University; 
O. S. Flint, United States National Museum 

of Natural History; R. F. Kirchner, Hun- 
tington, West Virginia; J. K. Liebherr, Cor- 
nell University; E. C. Masteller, Behrend 
College; R. F. Surdick, Front Royal, Vir- 
ginia; and J. R. Voshell, Virginia Polytech- 
nic Institute and State University. 

Peltoperla arcuata Needham 

Needham and Claassen (1925), Prison 
(1942) and Hitchcock (1974) provided ad- 
equate descriptions of this species. We are 
providing additional comparative figures of 
the genitalia and eggs to facilitate identifi- 
cation of both species (Figs. 3, 4, 1 1-1 5, 1 7). 

Specimens were examined from the fol- 
lowing locations: KENTUCKY: Boyd Co., 
Ashland; Powell Co., Mill Crk, Natural 
Bridge State Park. NEW YORK: Tompkins 
Co., Ithaca (holotype); Ringwood. PENN- 
SYLVANIA: Centre Co., Penn-Roosevelt 
Dam; Elk Co., Ridgeway Spring; Watercress 
Spring; Erie Co., 6-Mile Crk; Fulton Co., 
Crystal Spring; Westmoreland Co., Laugh- 
intown, Furnace Run. VIRGINIA: Dick- 
erson Co., Laurel Branch, Breaks Int. Park; 
Shenandoah Co., Springs, Little Sluice 
Mountain; Wythe Co., East Fork; Stoney 
Fork Reed Crk. WEST VIRGINIA: Brax- 
ton Co., Laurel Run, Falls Mill; Greenbrier 



Figs. 1-4. Feltopcrki eggs. 1, /'. tartcri. 300 x. 2, F. tartcri. detail of chorion and micropyli^s (M), 1000 x. 3, 
P. arcuata, 340 x. 4. /'. airmta, detail of chorion and micropyles, lOOOx. 

Co., Coats Run, North Fork Cherry Riv; 
Logan Co., Frogtown Hollow Copperas 
Mine Fork; Mingo Co., Laurel Fork Pigeon 
Crk; Pocahontas Co., Fork of Tea Crk. 

Peltoperla tarteri Stark and KondratiefT, 
Nk.w Spkciks 

Pchopciia arcuata: Stark and Stewart ( 1 98 1). 
in part. 

Male. — Forewing length 11-12.5 mm. 
General color brown patterned with pale 
brown. Tergum 9 unmodified, without dor- 
sal lobe. Paraprocts and tergum 10 typical 
ofgenus. Cerci essentially straight; first seg- 
ment elongate but not swollen basally (Figs. 
5, 8, and 9). Vesicle on sternum 9 slightly 
wider than long and rounded apically (Fig. 

6). Aedeagus membranous and multilobed; 
apex terminates in two sparsely setose lat- 
eral lobes and 3 mesal finger-like lobes cov- 
ered with fine spicules; a pair of spiculate 
mesolateral lobes are occasionally not 
everted (Fig. 10). 

Female. — Forewing length 12-13.5 mm. 
Subgenital plate truncate apically; vaginal 
sclerite sides parallel (Fig. 16). 

Egg. — Spherical to slightly ovoid; collar 
absent. Chorionic surface finely punctate 
giving surface a rough appearance. Micro- 
pyles typical ofgenus (Figs. 1, 2). 

Nymph. — Presently indistinguishable 
from /'. arcuata. 

Etymology. — This species is named in 
honor of Donald C. Tarter, Marshall Uni- 
versity. Huntington, West Virginia. 

Material examined.- Holotype <5, alio- 



/ /. 


Figs. 5-10. P. tarteri, male genitalia. 5, Terminalia dorsal. 6, Sternum 9. 7, Epiproct, lateral. 8, Paraprocts 
and epiproct, ventral (Giles Co., Virginia). 9, Paraprocts and epiproct, ventral (Wyoming Co., West Virginia). 
10, Aedeagus, ventral. 

type 9, 23 paratype $ and 43 paratype 9, 
West Virgina, Fayette Co., Big Hollow of 
Paint Creek, 19 May 1979, R. F. Kirchner. 
Additional paratypes: VIRGINIA: Craig 
Co., Hollow Hill Farms, 26 June 1977, B. 
Kondratieff, 1 <5, 1 9 (VPI); Floyd Co., trib- 
utary of Little Riv., 8 June 1978, B. Kon- 
dratieff, 5 S (VPI); Giles Co., Little Stoney 
Crk, 14 July 1971, M. Kosztarab, 2 6 
(USNM); same location, 26 June 1977, B. 

Kondratieff, 3 5, 1 9 (VPI); same location 
24 June 1978, B. Kondratieff, 7 5, 1 9 (VPI); 
Mountain Lake, small spring on Co. Rt. 700, 

B. Kondratieff, 1 5, 1 9 (BPS); Mud Branch, 
Mountain Lake, 26 June 1977, B. Kondra- 
tieff, 6 3, 2 9 (VPI); Virgin Timber Area, 
Mountain Lake, 15/18 July 1 978, K. A. and 

C. R. Parker, 5 3, 1 9 (VPI); Spring near Co. 
Rt. 613, 24 June 1978, B. Kondratieff, 9 5, 
5 9 (VPI); Greene Co., Pocosin Cabin, Shen- 



Figs. 1 1-15. P. arcuata, male genitalia. 1 1, Terminalia dorsal. 12, Sternum 9. 13, Paraprocts and epiproct. 
ventral. 14, Aedeagus, ventral. 15, Epiproct, lateral. 

andoah National Park, 20 July 1973, O. S. 
Flint, 1 5, 1 9 (USNM); Nelson Co., Stony 
Crk, Co. Rt. 751, 28 June 1983, 1 $ (BPS); 
Patrick Co., Spring into Rock Castle Crk, 
10 May 1983, B. Kondratieff, 1 5, 1 9 (BCK); 
Rockingham Co., Shenandoah National 
Park, 30 May 1976, O. S. Flint, 1 <5, 1 9 
(USNM); Rappahannock Co., Shenandoah 
National Park, Skyline Drive MP 71, 24 

June 1961, O. S. Flint, 4 <5, 1 9 (USNM). 
WEST VIRGINIA: Wyoming Co., Clear 
Fork Guyandotte Riv, 19 March 1978 
(reared), L. Evans, 3 <5 (RFK). 

The holotype and allotype are deposited 
in the United States National Museum of 
Natural History (USNM), the paratypes in 
the authors' collections (BPS, BCK), and in 
those of R. F. Kirchner (RFK), Virginia 





Figs. 16, 
teri. 11, P. 

1 7. Peltoperla female genitalia. 1 6, P. tar- 

aedeagal apex is also quite different, being 
apically multilobed in P. tarteri (Fig. 10). 
The epiprocts of both species are variable. 
Figures 8, 9 indicate the range of variation 
for this structure in P. tarteri. 

Females of both species are similar; how- 
ever, the subgenital plate of P. tarteri is 
longer and typically more truncate apically, 
whereas that of P. arcuata is broadly round- 
ed apically. Internally the vaginal sclerite of 
P. arcuata is broader across the anterior 
margin than posteriorly, whereas in P. tar- 
teri the sides of the sclerites are parallel. The 
eggs are similar for both species, although 
subtle differences are evident in Figs. 2 and 
4. Nymphs of both species are very similar 
and examination of reared material provid- 
ed no useful characters for separation. 

The distribution of these species is some- 
what enigmatic, since their ranges overlap 
broadly in the Blue Ridge, Ridge and Valley 
and Appalachian Plateau physiographic 
provinces of Virginia and West Virginia. 
Both species of Peltoperla typically occur 
only in crenon habitats (spring sources and 
springbrooks) of the higher, central Appa- 
lachians. The two species have not been tak- 
en together, however, they are often col- 
lected with species of Tallaperla such as T. 
anna (Needham and Claassen) and T. ma- 
ria (Needham and Claassen). 


We thank the individuals listed above for 
arranging the loans of specimens. Sarah Fai- 
son, Univ. of Mississippi School of Den- 
tistry, assisted in preparing SEM micro- 
graphs. This study was supported by NSF 
grant #BSR-8407455. 

Polytechnic Institute and State University 
(VPI), and the Monte L. Bean Museum, 
Brigham Young University (MLBM). 

Diagnosis and discussion. — Males of P. 
tarteri are easily separated from P. arcuata 
by the absence of the curved cerci and by 
the lack of a dorsal lobe on tergum 9. The 

Literature Cited 

Claassen, P. W. 131. Plecoptera nymphs of North 
America north of Mexico. Thomas Say Found. 
Entomol. Soc. Am. 3: 1-199. 

Frison, T. H. 1942. Studies of North American Ple- 
coptera with special reference to the fauna of Il- 
linois. Bull. 111. 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. State Geol. Nat. Hist. Surv. Conn. 

107: 1-262. 
Needham, J. G. 1905. New genera and species of 

Perlidae. Proc. Biol. Soc. Wash. 18: 107-110. 
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. Am. 2: 1-397. 
Stark, B. P. and K. W. Stewart. 1981. The nearctic 
genera of Peltoperlidae (Plecoptera). J. Kans. 
Entomol. Soc. 54: 285-311. 


89(1), 1987, pp. 147-156 


David R. Smith and Edward M. Barrows 

(DRS) Systematic Entomology Laboratory, BBII, Agricultural Research Service, U.S. 
Department of Agriculture, c/o National Museum of Natural History NHB 168, Wash- 
ington, D.C. 20560; (EMB) Department of Biology, Georgetown University, Washington, 
D.C. 20057. 

Abstract. — Malaise traps were used to determine the species, abundances, and seasonal 
occurrences of adult sawflies in the Washington, D.C, metropolitan area. The traps, placed 
in two urban environments for six years and three more natural habitats for two years 
from March to November, obtained 948 sawflies representing 117 species. Peaks in num- 
bers of species and specimens were in May. The more abundant species were Acordulecera 
pellucida (Konow), Schizocerella pilicornis (Holmgren), AUantus nigritibialis (Rohwer), 
and Ametastegia pallipes (Spinola). Known hosts for all captured species include repre- 
sentatives of 53 plant genera, with Rosaceae being the most utilized plant family in urban 
environments. Notes on selected species are given. 

Urban environments are becoming more 
common and are of increasing biological in- 
terest (Frankie and Koehler, 1983), but no 
quantitative study has yet been done on the 
urban sawfly fauna. As larvae, all sawfly 
species are plant feeders, and many are, or 
have the potential of becoming, important 
plant pests in urban environments. There 
are about 1100 species in North America 
(Smith, 1979). The purpose of this study is 
to determine what sawfly species are present 
in urban environments, their abundances, 
seasonal occurrences, and host-plant utili- 
zations. The urban environment in our study 
consisted of relatively well manicured and 
ornamentally landscaped yards and semi- 
natural areas within a metropolitan region. 

Materials and Methods 

Five sites were studied in the Washing- 
ton, D.C, metropolitan area, two urban en- 
vironments for six years and three more 
natural environments for two years. The 

natural environments were less disturbed, 
non-landscaped areas. The urban environ- 
ments were at residences in Glen Echo, 
Montgomery Co., Maryland (yard-garden 
trap), and near Annandale, Fairfax Co., Vir- 
ginia (yard trap). The natural environments 
were woodland (wooded-stream-habitat 
trap), woods edge (ecotone trap), and field 
(field trap), at the David W. Taylor Naval 
Ship Research Center (TNSRC) in Mont- 
gomery Co., Maryland, 5 km NNW of Glen 
Echo. These sites are described in more de- 
tail by Barrows (1986). Because only three 
specimens were collected in the field trap, 
they are combined with the ecotone trap in 
Table 1. 

Bioequip® (Santa Monica, Calif) Mal- 
aise traps were used at the Glen Echo and 
TNSRC sites. They are 2 m tall, pyramidal, 
with four 0.8 m- rectangular openings. In 
Fairfax Co., a Townes-style Malaise trap 
was used. It is rectangular with a 1.8 m- 
opening on each side and a killing jar at one 



Table 1. Families, species, number captured, flight times (earliest and latest dates), and recorded hosts of 
sawflies. YT = yard trap, Fairfax Co., Va.; YGT = yard-garden trap. Glen Echo, Md.; WSHT = woodland- 
stream habitat trap, TNSRC; ET = ecotone trap, TNSRC. Recorded hosts are from Smith (1979) and Gibson 

Families, Species 







Recorded Hosts 


Xyela bakeri Konow 



III-20; IV-7 


Xyela obscura (Strobl) 







Pamphilius middlekauffi Shi- 





nohara and Smith 

Pamphilius ochreipes (Cres- 







Pamphilius pullatus (Cresson) 






Onycholyda luteicornis (Nor- 







Onycholyda rufofasciatus 



VII- 12 



Acordulecera dorsalis Say 





30; VIII-4 

Quercus. Corylus, 
Juglans. Casta- 

Acordulecera maculata Mac- 






Acordulecera mellina Mac- 






Acordulecera pellucida (Ko- 







Sphacophilus cellularis (Say) 





Ipomaea, Convol- 

Arge humeralis (Beauvois) 






Arge clavicornis complex 




Schizocerella pilicornis 








Sterictiphora serotina Smith 






Atomacera decepta Rohwer 





Atomacera debilis Say 







Zaraea lorucerae (L.) 









Monoctenus sp. 





Diprion similis (Hartig) 






Xiphydria maculata Say 





Xiphydria tibialis Say 



VI- 12; VI- 17 

Ulmus, Betula, 

Quercus, Rhus, 
Tilia, Prunus, 



Table 1. Continued. 

^T \GT WSHT ET Total 

Recorded Hosts 


Tremex columba (L.) 

Janus integer (Norton) 


Hemitaxonus albidopictus 

Hemitaxonus dubitatus (Nor- 

A neugmenus flavipes (Norton) 

Heptamelus ochwleucus (Ste- 

Dolerus nitens Zaddach 
Dolerus unicolor (Beauvois) 
Loderus vestigialis apricus 

Cladius difformis (Panzer) 
Priophorus pallipes (Lepele- 

Hoplocampa marlatti Rohwer 
Cratewcercus fraternalis (Nor- 
Euura sp. 

Nematus lipovskyi Smith 
Nematus erythrogaster (Nor- 
Nematus hudsoniimagnus 

10 1 VIII-4 

3 3 V-26-VI-2 

1 V-16 

3 VI-23-VII-16 

1 VII-15 

3 V-26;IX-16 

37 1 





2 IV-7; V-8 

3 V-22-V-29 

2 16 









deciduous trees 





Prunus, Cratae- 
gus, Alnus 
Prunus ? 





Nematus oligospilus Foerster 





Nematus abbotii Kirby 




V-8; V-9 


Nematus (ribesii gp.) 




Nematus tibialis Newman 





Neopareophora litura (KJug) 






Pachynematus corniger (Nor- 








Pristiphora rufipes Lepeletier 




V-20; VII-14 


Pristiphora bivittata (Norton) 






Pristiphora abbreviata (Hartig) 






Pristiphora banksi Marlatt 





Pristiphora acidovalva Wong 





and Ross 

Pristiphora cincta Newman 




Betula, Salix, 

Pristiphora zella Rohwer 




Rubus, Geum, 

Pristiphora chlorea (Norton) 




IV-27; V-14 




Table 1. Continued. 

Families. Species 







Recorded Hosts 

Pristiphora sp. (bivittata gp.) 



IV- 17 

Pristiphora micronematica 






Amaiironematus sp. 





Caliroa liinata MacGillivray 




Caliroa quercuscoccineae 




V-29; VIII-3 



Caliroa fasciata (Norton) 






Caliroa obsoleta (Norton) 





Endelomyia aethiops (F.) 





Met alius rohweri Mac- 






Nefusa ambigua (Norton) 





Fenusa pusilla (Lepeletier) 






Eutomostethus ephippium 





Poa. grass 


Eutomostethus luteiventris 






Eupareophora parca (Cresson) 






Phymatocera fumipennis 







Phymatocera racemosae 



VII- 15 

Smilacina, Poly- 



Paracharactus rudis (Norton) 






Monophadnoides geniculatus 






Periclista marginicollis (Nor- 







Periclista albicollis (Norton) 







Periclista inaequidens (Nor- 



IV- 10 



Halidamia affinis (Fallen) 








Pseudosiobla excavata (Nor- 






Allantus nigritibialis (Rohwer) 







Monostegia abdominalis (F.) 





Ametastegia aperta (Norton) 






Ametastegia articulata (Klug) 





Rumex, Polygo- 

Ametastegia equiseti (Fallen) 





Ametastegia becra Smith 




Ametastegia pallipes (Spinola) 






Ametastegia pulchella (Roh- 
Ametastegia tener (Fallen) 









Empria maculata (Norton) 






Fragaria, Poten- 

Empria muhicolor (Norton) 




Alnus, Betula 



Table 1. Continued. 

Families, Species 







Recorded Hosts 

Macremphytus testaceus (Nor- 









Taxonus epicera (Say) 





Taxonus pallidicornis (Nor- 



VI- 12 



Taxonus pallipes (Say) 





Taxonus terminalis (Say) 







Lagium atroviolaceum (Nor- 





Sambucus, Vi- 



Aglaostigma semiluteum 






Tenthredo rufopecta (Norton) 






Tenthredo sp. 




Macrophya alba MacGillivray 




Macrophya albomaculata 






Macrophya cinctula (Norton) 




Macrophya flavicoxae (Nor- 






Macrophya flavolineata (Nor- 




V-8; V-29 


Macrophya formosa (KJug) 






Macrophya goniphora (Say) 





Macrophya lineatana Rohwer 




Vl-9; VI-29 

Macrophya macgillivrayi Gib- 




Macrophya mensa Gibson 






Macrophya mixta Mac- 






Macrophya pannosa (Say) 




V-7; V-14 


Macrophya pulchella (Klug) 




Macrophya senecca Gibson 




Macrophya simillima Rohwer 





Macrophya succincta Cresson 



IV-21; V-1 

Macrophya tibiator Norton 



V-29; Vl-12 

Macrophya trisyllaba (Norton) 





Macrophya varia (Norton) 







Macrophya zoe Kirby 





V-21; V-25 







end about 2 m above the ground. The traps 
were left up continually from the last week 
in March to the first week in November 
from 1980-1985 for the yard traps and 
1983-1984 for the TNSRC traps. Collec- 
tions were made about once a week. Spec- 
imens were identified by DRS, except for 
most Pristiphora which were identified by 
H. R. Wong. Voucher specimens are de- 

posited in the National Museum of Natural 
History, Washington, D.C. 

Results and Discussion 

Species and abundance. — Species and 
numbers of specimens collected from each 
site are listed in Table 1. At all sites, 948 
specimens representing 10 families, 53 gen- 
era, and 1 1 7 species were caught. This rep- 







30 . 


20 . 


10 . 




■ ■__«_■• « _ _ _ _ 


1 2 3 4 5 6 7 9 1011 141516171819 20 38 40 48 51 63106138 

Number of Individuals 

Frequencies of specimens per species in the total sample. 

resents more than half of the approximately 
220 species of Symphyta that arc expected 
to occur in the Washington, D.C., metro- 
politan area (DRS, personal notes). In the 
two urban environments, 820 specimens of 
104 species were collected. For all sites, 35 
species were found only in Fairfax Co., 16 
in Glen Echo, and 13 at TNSRC. Fairfax 
Co. and Glen Echo had 20 species in com- 
mon; Fairfax Co. and TNSRC, 14 species; 
and Glen Echo and TNSRC, 9 species. Ten 
species were found in all three study sites. 
Differences in species composition at each 
site (Table 1) are likely due to the different 
hosts present near the traps, length of sam- 
pling time (two years at TNSRC), and/or 
the type of trap used. For example in Glen 
Echo, 402 specimens were caught in two 
traps, whereas in Fairfax Co., 418 speci- 
mens were caught in one trap. 

Frequencies of specimens per species (Fig. 
1) indicated that only a few species are pres- 
ent in large numbers. For 83 species (81%) 
only four, or fewer, individuals were caught. 
The collection of only one to several spec- 
imens of a species may indicate species rar- 
ity, accidentally wind-blown individuals 
from other habitats, or representatives of 

groups, e.g. Macrophya, that are strong fliers 
and may fly far in search of hosts in other 

Several species, not listed in Table 1 , were 
hand collected in the Fairfax Co. yard but 
never collected in the traps. These are Ato- 
macera decepta Rohwer defoliating Hibis- 
cus sp., Allantiis viennensis (Schrank) on 
Rosa sp., Uroccrus cressoni Norton collect- 
ed at large, and a Neodiprion sp. on Finns 
sp. However, most species collected in the 
traps were never observed, or hand collect- 
ed, in the yard. This indicates a limitation 
of the use of a single sampling method. A 
combination of trapping and observation 
gives a more complete picture of the fauna. 

Seasonal distribution.— The earliest saw- 
flies collected were Xyela spp. on March 20 
and the latest was Pachynematus corniger 
on November 4. Sawflies were present 
throughout the intervening period, but the 
peak in numbers of species and numbers of 
specimens was from the second half of April 
through the second half of May (Figs. 2, 3). 
The flight time for each species is given in 
Table 1, and examples of different types of 
flight periods are shown in Fig. 4. Adults 
appear for a short time once a year for most 



species, indicating that most are univoltine, 
and most of these appear in the spring: 64 
species appeared during April to June, sim- 
ilar to the flight period of Eutomostethus 
ephippium (Fig. 4); 16 species appeared 
mostly during June to July; 1 1 species ap- 
peared during July to August; and 4 species 
were late fliers, appearing only during Au- 
gust and September, similar to the flight pe- 
riod of Metallus rohweri (Fig. 4). One 
species, Allantus nigritibialis, had two flight 
periods, one in spring and one in late sum- 
mer (Fig. 4). Seventeen species were col- 
lected through a large part of the season, 
and peaks of occurrence were apparent dur- 
ing several periods, similar to the flight pe- 
riods of Acordulecera pellucida and Schizo- 
cerella pilicornis in Fig. 4; these apparently 
have several generations a year. 

Host-plants. — Sawflies collected in all 
sites feed on 53 plant genera in 30 families 
based on recorded hosts (Tables 1, 2). Host- 
plant records are from Smith (1979) and 
Gibson (1980). If more than one host is 
recorded for a species (e.g. Acordulecera 
dorsalis), all are listed, and the total speci- 
mens for the species is given for each host 
in Table 2 since it is not known which host(s) 
is utilized in the study sites. For sawflies 
collected in the urban environments, the 
most common hosts are those used as or- 
namentals or those present as weeds, and 
Rosaceae was the most utilized plant family 
based on numbers of sawfly species and 
specimens. At TNSRC, however, only two 
species for which the host is known are as- 
sociated with members of Rosaceae. Since 
sawfly species and abundance are depen- 
dent on available hosts, the trend in urban 
environments should be toward dominance 
of those species associated with cultivated 

Urban species. — Most species collected in 
the urban environments also occur in nat- 
ural environments, though some (Table 1) 
were much more abundant in the yards 
sampled. Only two species, Allantus nigri- 
tibialis and A. viennensis (both associated 


1 1^^ I 1 

1 1 1 





^^^^F^^^^V^^:^ . ■ 

^ • 


M A 

O N 

Figs. 2, 3. Numbers of species and individuals 
caught from March (M) to November (N), based on 
half month intervals. 1 = first half of the month; 2 = 
second half of the month. 

with cultivated Rosa spp.) have yet to be 
found outside urban environments (DRS, 
personal notes). 

Notes on Species 

Xyela spp. —The two species were the ear- 
liest flying sawflies, caught from March 20 
to April 7. Larvae feed in staminate pine 
cones, and adults fly to flowers of other 
plants to feed on pollen; their flight time 
usually coincides with the availability of 
adult food. When numerous, larvae are 
sometimes a nuisance when dropping from 
the pine trees to the ground where they enter 
the soil and form pupal cells. 

Onvcholvda luteicornis.— This was the 



Table 2. Host plants and number of species and 
number of individuals collected associated with those 

Table 2. Continued. 

Number Number 

of of 

Sawfly Speci- 

Planl Hosts 

Species mens 











































Pop u I us 



Corn us 




Number Number 

of of 

Sawfly Speci- 

Plant Hosts 

Species mens 















Polygonatum 1 1 

Smilacina 2 8 

Juncus 1 1 

"grasses" 5 105 

most common pamphiliid and was cap- 
tured only in the yard traps. Larvae live and 
feed in rolled leaves and webs of their own 
making on Rubus spp. 

Acordulecem dorsalis.— The peak flight 
was in May and the species was most com- 
mon in Fairfax Co. and TNSRC. In Fairfax 
Co., larvae may feed on Quercus sp. or Car- 
ya sp. nearby. This group has not been stud- 
ied, and there may be more than one species 
under this name. 

Acordulecera pellucida. —The host of this 
species is not known. It was collected in 
both yard traps from April to October (Fig. 
4) with peak flight periods twice during the 

Schizocerella pilicornis. —This species 
represented 14% of the specimens collected 
(138 specimens), most of which were from 
the yard-garden trap. It is common in urban 
environments where its larvae feed on purs- 
lane (Portulaca spp.), a commonly culti- 
vated, or weed, plant. Flight time is shown 
in Fig. 4. 

Atomacera decepta. —This species is a de- 
foliator of ornamental Hibiscus on which it 
was found in Fairfax Co. It was not collected 
in the trap. It may be present from May to 



September, passing through several gener- 

Zaraea lonicerae. —This adventive Eu- 
ropean species has become common in the 
D.C. area during the past 20 years. Larvae 
feed on Lonkera spp., and adults fly during 
April and May. 

Neodiprion sp. — No adults were collected 
in traps, but larvae were found feeding on 
Pinus spp. in Fairfax Co. Several species are 
known to be destructive to ornamental pines. 

Diprion similis.— One specimen of this 
adventive European species was collected 
in Fairfax Co., and it is the first record of 
this species for northern Virginia. This pine- 
feeding species was known from northeast- 
em United States south to Pennsylvania and 
from an isolated area in extreme south- 
western Virginia into North Carolina. 

Xiphydria maculata. — The larvae of this 
species bore in wood in branches of Acer 
spp. Adults were found only at the Fairfax 
Co. site where maple is a dominant tree. 

Heptamelus ochroleucus.— This adven- 
tive European species feeds on ferns and 
was known only from British Columbia and 
New York. More recently it was found in 
Maryland, and now in Fairfax Co. Adults 
were collected in spring and fall. 

Dolerus nitens.— This adventive Euro- 
pean species feeds on grasses and has spread 
rapidly in North America. It is sometimes 
abundant during its very short flight period. 

Cladius diffor mis.— Commonly called the 
bristly rose slug, it is one of the common 
defoliators of cultivated roses and was col- 
lected at both yard trap sites and found feed- 
ing on roses in Fairfax Co. This species has 
several generations a year. 

Amauronematus sp.— This species can- 
not be identified until taxonomic difficulties 
are resolved. Known hosts for members of 
the genus are Alnus spp. and Salix spp. 

Metallus rohweri.— This species was col- 
lected only in late summer and early fall 
(Fig. 4). The larva is a leafminer of Rubus 

212'121 21212 

M A M J J A 

Fig. 4. Adult flight periods for selected species. Eu- 
tomostethiis ephippium (Ee), early-spring; Allantus ni- 
gritibialis (An), spring and fall; Acordulecera pellucida 
(Ap), entire season; Schizocerella pilicornis (Sp), pri- 
marily mid-summer; and Metallus rohweri (Mr), late 
summer and early fall. 

Fenusa pusilla.—Knov^n as the birch 
leafminer, this species is a common pest 
farther north, though it is sometimes abun- 
dant on ornamental birch in the D.C. area. 
A single specimen was collected in the field 
trap at TNSRC. 

Eutomostethus ephippium.— This is ad- 
ventive from Europe and has spread rapidly 
in North America. It can be abundant dur- 
ing its short flight period in early spring (Fig. 

Allantus nigritibialis. —This was a rarely 
collected species before we initiated trap- 
ping. It was found on and reared from larvae 
feeding on cultivated roses in Fairfax Co., 
and adults were collected in both yard trap 



sites. Collection records indicate a spring 
and a late summer to fall generation (Fig. 

Allantus viennensis. —This species was not 
collected in the traps but was reared from 
larvae feeding on cultivated roses in Fairfax 
Co. It is an adventive European species, first 
recorded in North America from near Ith- 
aca, New York. The Virginia specimens 
represent the second North American rec- 

Monostegia abdominalis. —Specimens 
collected in the Fairfax Co. trap represent 
a new record for Virginia and the southern- 
most record for the species. 

Ametastegia pallipes.— This species was 
collected in large numbers and only in both 
yard trap sites. Its host, Viola, is a common 
weed species. Adults were collected 
throughout the season from April to Oc- 
tober with peak numbers during the end of 
May and first of April, the end of July and 
first of August, and the end of September. 

Macrophya spp. — Of the approximately 
40 species in eastern North America (Gib- 
son, 1 880), we collected 20 during this study. 
Most collections are represented by very few 
specimens. Hosts are not known for many 
of the species. They are strong fliers and 
many may have been accidental catches and 
may not be associated with plants in the trap 


H. R. Wong, Canadian Forestry Service, 
Northern Forestry Centre, Edmonton, Al- 
berta, identified most of the Pristiphora. J. 
R. Gauthey permitted us to trap sawflies at 
the Taylor Naval Ship Research Center. The 
Washington Field Biologists' Club and 
Georgetown University provided part of the 
financial support for this work. We appre- 
ciate the comments of the following review- 
ers: H. R. Wong; H. Goulet, Biosystematics 
Research Institute, Agriculture Canada, Ot- 
tawa; Robert D. Gordon and E. E. Grissell, 
Systematic Entomology Laboratory, USDA, 
Washington, D.C. 

Literature Cited 

Barrows, E. M. 1986. A hornet, paper wasps, and 
yellowjackets (Hymenoptera: Vespidae) in sub- 
urban habitats of the Washington, D.C, area. Proc. 
Entomol. Soc. Wash. 88: 237-243. 

Frankie, G. W. and C. S. Koehler, eds. 1983. Urban 
Entomology: An Interdisciplinary Approach. 
Praeger Press, New York. 493 pp. 

Gibson, G. A. P. 1 980. A revision of the genus Mac- 
rophya Dahlbom (Hymenoptera: Symphyta, Ten- 
thredinidae) of North America. Mem. Entomol. 
Soc. Can. No. 114, 167 pp. 

Smith, D. R. 1979. Symphyta, pp. 3-137. In Krom- 
bein, K. V. et al., eds.. Catalog of Hymenoptera 
of America north of Mexico. Vol. 1. Smithsonian 
Institution Press, Washington, D.C. 


89(1), 1987, pp. 157-166 





V. Baimai, R. G. Andre, B. A. Harrison 


(VB, UK) Department of Biology, Faculty of Science, Mahidol University, Rama VI 
Road, Bangkok 10400, Thailand; (RGA, BAH) Department of Entomology, Walter Reed 
Army Institute of Research, Washington, D.C. 20307-5 100, USA; (LP) U.S. Army Medical 
Component, AFRIMS, Rajvithi Road, Bangkok 10400, Thailand. 

Abstract.— Cvo^sm% and chromosomal evidence is presented for two additional sibling 
species, dims C and dims D, within the taxon Anopheles dims Peyton and Harrison, in 
Thailand. The affinities of the four currently recognized species in this complex in Thailand 
and the limitations of certain techniques used to identify the species are discussed. 

Cytogenetics is one of the most useful tools 
for elucidating cryptic species of insects 
(Dobzhansky, 1970; White, 1973). The use 
of this method, together with biochemical, 
behavioral, ecological and morphological 
techniques has led to the recognition of a 
significant number of sibling species com- 
plexes of anopheline mosquitoes in different 
parts of the world (Bryan and Coluzzi, 1971; 
Kitzmiller et al., 1973; White et al., 1975; 
Kitzmiller, 1976; Coluzzi et al., 1979; Steg- 
nii and Kabanova, 1978; Peyton and Har- 
rison, 1979, 1980; Green and Miles, 1980; 
Subbarao et al., 1983; Green and Baimai, 
1984; Green et al., 1985). The discovery of 
these cryptic species is a highly significant 
step in the development of rational and ef- 
ficient control programs against the vectors 
of various mosquito-borne diseases. 

One of the most renowned vectors of hu- 
man malaria parasites in Southeast Asia is 
Anopheles balabacensis Baisas, a member 
of the widely distributed Leucosphyrus 
Group. Recently, it was demonstrated that 
An. balabacensis is a species complex (Pey- 

ton and Harrison, 1979, 1980; Baimai et 
al., 1981; Hii, 1982, 1984, \9^5). Anopheles 
dims Peyton and Harrison, was described 
as a species distinct from An. balabacensis 
in 1979, and is considered widespread in 
peninsular Malaysia and Thailand, while /I «. 
balabacensis sensu stricto is confined to the 
type-locality on Balabac Island and to 
neighboring areas of Palawan Island, Sabah 
and northeast Kalimantan (Peyton, unpub- 
lished data; Peyton and Harrison, 1979; Hii, 

Baimai et al. (1981) recently demonstrat- 
ed that the laboratory colony strain o{ An. 
balabacensis Perils form from The Institute 
of Medical Research (Kuala Lumpur) 
showed different sex chromosome charac- 
ters as seen in mitotic karyotype as well as 
on salivary gland polytene chromosomes. 
Genetic incompatibility between dims and 
the balabacensis Pedis form also was ob- 
served (Baimai and Harrison, 1980). The 
recognition of the balabacensis Perils form 
as a distinct genetic species from dims was 
confirmed later by the detailed studies of 



Table 1 . Laboratory family stocks of Anopheles dims complex from different localities in Thailand (otherwise 
indicated) used in this study. 

d,rus Species 

Locality (Date) 

No. of 

No. of Families Maintained (Code) 


Chonburi (1964) and Nakhon Rat- 



Bangkok colony (BK)* 
2 (TL)** 


Perlis State, Malaysia (1965) 
Trengganu State, Malaysia (1982) 


IMR colony (PR)* 
1 (MH) 


Kanchanaburi(1980, 1982) 




Ranong(1983, 1984) 



1 (PG) 

* The mixed colony. 
** The isofemale line now used as the reference stock for dims A. 

Hii (1982) who designated the Bangkok col- 
ony strain as dims A and the Perlis form as 
dims B. 

In 1979-80, E. L. Peyton, at the Smith- 
sonian Institution (personal communica- 
tion) advised the authors that two addition- 
al members of the Dirus Complex existed 
in Thailand which could be separated from 
dims A and dims B by morphological char- 
acters. Subsequently, these members were 
collected and colonies of both were estab- 
lished in 1980-81 at the Armed Forces Re- 
search Institute of Medical Sciences (AF- 
RIMS), Bangkok. 

Recently, Wibowo et al. (1984) discov- 
ered marked differences in the amount and 
distribution of constitutive heterochroma- 
tin in the sex chromosomes, as revealed by 
the Hoechst 33258 staining technique, of 
specimens from a Kanchanaburi colony de- 
rived from a single isofemale of one of the 
two new members. Based on comparative 
cytological data, these investigators con- 
firmed Peyton's morphological findings that 
the Kanchanaburi isoline represents a dis- 
tinct genetic species designated as dims C, 
which occurred sympatrically with dims A. 
Additional cytogenetic studies also have re- 
vealed that the second undescribed member 

noted by Peyton is distinct and represents 
a 4th member, dims D, of the complex in 

This paper presents crossing and chro- 
mosomal evidence supporting the existence 
of two additional sibling species, dims C 
and dims D, within the Dirus Complex in 

Materials and Methods 

Individual wild caught Anopheles females 
were identified to species by morphology, 
isolated, and allowed to oviposit. Subse- 
quent Fi larvae were reared in the labora- 
tory and used for cytogenetic confirmation 
of the species. Mitotic and salivary gland 
polytene chromosomes were prepared from 
4th instar larvae using the method of Bai- 
mai et al. (1981). Isofemale lines of each 
cytotype were set up with respect to the X 
and Y chromosome configurations and 
maintained in the laboratory for further 
crossing experiments (Table 1). 

The mosquitoes used in this study came 
from isofemale lines that were determined 
cytogenetically and maintained at the De- 
partment of Medical Entomology, AF- 
RIMS, Bangkok, and at the Department of 
Biology, Mahidol University, Bangkok. 
Anopheles dims A (Bangkok colony strain = 

VOLUME 89, NUMBER 1 159 

Table 2. Crossing combinations among the isolines of ^'Anopheles dims" from different geographic origins. 

No. of 




No. of 

Mean No. of 
tggs per 









% Eggs Hatching 























A (I) 




















































































Sterile male F, hybrids. 

BK) was used as the standard stock, whereas 
the IMR colony strain (Perhs form = PR) 
and the Trengganu colony strain (= MH) 
represented An. dims B in this study. The 
isoline strain from Kanchanaburi (= KN) 
represents dims C. Two other isoline strains 
of '^dims" were collected from Ranong (= 
RN) and Phangnga (= PG) in southern 
Thailand, and represent dims D (Table 1). 
Combinations of reciprocal pair-matings 
(Table 2) among the different cytotype 
strains were performed by the artificial mat- 
ing technique of Ow Yang et al. (1963). In 
each cross pair-mating 10-20 individual fe- 

males were mated with mature males. After 
successful copulation, each female was iso- 
lated in an oviposition vial. The number of 
females ovipositing, number of eggs ovi- 
posited and hatching and number of 
emerged F, adults were scored daily in the 
same manner as described by Klein et al. 
(1985). Fertility of F, hybrids was deter- 
mined later by self-crossing among them- 
selves as well as backcrossing to the respec- 
tive parental strains (Table 3). Genetic 
incompatibility could be inferred on the de- 
gree of synapsis in salivary gland polytene 
chromosomes of F, larval hybrids of anoph- 



Table 3. Backcrossing and selfcrossing experiments of F, hybrids from the crosses between isolines of Anoph- 
eles dims A and An. dims C. 


— No. of Females 

No. of Ovipositions 

Mean No. of Eggs 



Ovipositing (Total) 


per Oviposition 

% Eggs Hatching 

F, (D* 
































F, (11)** 






F, (11) 







F, (11) 





F, (II) 




F, (11) 

F, (II) 




* 1 = cross between female dims C x male dims A. 
** 11 = cross between female dims A x male dims C. 

elines, and the degree of viability was de- 
termined by egg hatch rates. 


Hybridization tests.— The resuUs of cross- 
mating experiments among the different 
sibling members of the complex are sum- 
marized in Table 2. Crosses between the 2 
dirus B isoline strains (MH and PR), which 
showed the same mitotic karyotype, yielded 
a large number of eggs per female (85.9 and 
142.2) and a high percentage of eggs hatch- 
ing (54.0% and 57.5%). Adult F, hybrids of 
both sexes from both directions were fully 
fertile and continued to produce progeny for 
several generations. There was no evidence 
of genetic incompatibility between these two 
strains. The crossing evidence clearly indi- 
cates that the MH and PR strains belong to 
the same genetic species. An. dirus B. 

Crosses between the dirus A and dirus C 
isoline strains also were successful in both 

directions, producing a large number of eggs 
(averaging 59.5 and 96.0 per female, re- 
spectively) and a high percentage of eggs 
hatching (47.3% and 50.0%, respectively). 
Many Fi adults of both sexes were obtained 
in each direction. Crosses between female 
dirus C X male dirus A (designated as cross 
I) yielded fully fertile male and female F, 
hybrids. In contrast, the reciprocal cross be- 
tween female dirus A and male dirus C (des- 
ignated as cross II) produced fertile Fi fe- 
males, but completely sterile F, males (Table 
2). These results indicate that genetic in- 
compatibility exists between these two iso- 
lines, at least in one direction of hybridiza- 
tion. These crossing data are supported by 
cytological evidence described below from 
the F, hybrids. 

All combinations of crosses involving the 
dirus B isoline strains (PR and MH) with 
the other dirus isolines were less successful 
than those matings mentioned earlier. 



Crosses between dims B (MH) and dims A 
in both directions produced a large number 
of eggs (average of 103.2 and 108.3 per fe- 
male). However, very low percentages of the 
eggs hatched in these crosses. Furthermore, 
very few adult F, offspring emerged, and of 
these, the F, males were sterile. Crosses be- 
tween female dims B (PR) x male dirus C 
were less successful because only 10.2% of 
the eggs hatched, and no adults emerged. 
The reciprocal cross between female dims 
C X male dims B (PR) gave more eggs (av- 
erage of 88.3 per female), but none hatched. 
Furthermore, a remarkable example of ge- 
netic incompatibility was obtained from the 
crosses between dims D (PG) and dims B 
(MH). A very small percentage of the eggs 
hatched (0.3%), and only in one direction. 
In the reciprocal mating of this cross, none 
of the 10 artificially inseminated females 
produced eggs. 

The dims D isoline strains (RN and PG) 
exhibit similar karyotype and polytene band 
sequences. Unfortunately, the PG strain was 
lost before the RN strain was obtained, thus 
cross mating tests between them were not 
made. However, cytological evidence in- 
dicates that they are conspecific strains. All 
combinations of cross matings among the 
dims D isoline strains (RN or PG) with oth- 
er isolines yielded either very small num- 
bers of F, female hybrids or no Fi hybrids 
at all (see groups 5, 6 and 7 in Table 1). 
These results clearly indicate that the RN 
and PG isoline strains of dims D were ge- 
netically distinct from the other dims iso- 
lines employed in this study. 

Cytological evidence. — The examination 
of F| hybrid larval salivary gland polytene 
chromosomes revealed some differences in 
banding sequences. Based on the standard 
salivary gland polytene chromosomes of ^z- 
ms A (Baimai et al., 1980), the F, female 
larvae from the cross between female dims 
A X male dims C exhibited approximately 
5-10% asynapsis of the chromosome com- 
plement (Fig. 1). In addition, marked dif- 
ferences in polytene banding sequences were 

observed at zone 6 (Fig. 1 , arrow) and at the 
tip of the X chromosome (Fig. 2, arrow), as 
well as at the tips of chromosome arm 2L 
and arm 2R (Figs. 3, 4, respectively). These 
differences are good chromosome markers 
for the dims C karyotype. 

Larval salivary gland polytene chromo- 
somes of F, hybrid females from the cross 
between female <^/>W5 A x male <iz>wj B (PR) 
showed approximately 80% asynapsis of the 
chromosome elements (Fig. 5). This sug- 
gests that genetic differentiation at a sub- 
microscopic level between these two species 
is more extensive than in the case of dims 
A and dims C. 

The chromosome complement of the di- 
ms D (RN and PG) strains is remarkably 
different from the standard dims A colony 
strain. The F, female larval chromosomes 
from the cross between female dims A x 
male dims D (RN) showed over 90% asyn- 
apsis along the 5 chromosome arms (Fig. 
6). The X chromosome of the dims D (RN) 
strain exhibited a fixed inversion covering 
zones 1 and 3 of the X chromosome com- 
pared with the standard sequence of the di- 
ms A strain (Fig. 7). Zone 6 of the X chro- 
mosome of the F, female hybrids was 
asynapsed completely. Moreover, asynapsis 
in chromosome arm 2R of F, hybrids in 
this case was more pronounced (Fig. 8) than 
in the case of dims C x dims A Fi hybrids. 
Overall, asynapsis was a persistent feature 
of the hybrid polytene chromosomes of the 
Dirus Complex. In addition, an analysis of 
the mitotic karyotype showed that the X 
and Y chromosomes of the dims D (RN) 
strain are shorter than those of dims A as 
can be observed in Fi hybrid larval chro- 
mosomes (Figs. 9, 10). Thus, the cytological 
observations clearly support the sterility and 
viability evidence from the hybridization 
experiments described above. 


Recent morphological, genetic and cyto- 
genetic studies of the taxon dims have re- 
vealed that it consists of at least 3 genetic 






Figs. 1-5. Figs. 1^. Larval salivary gland polytene chromosomes of F, hybrid females from the cross matings 
between female dims A x male dirus C. 1 , Condition of synapsis along the 5 chromosome elements, zone 6 of 
the X chromosome (small arrow) is almost totally asynapsed. 2, Complete synapsis of zones 1-5 of the X 
chromosome with a distinct banding difference at the tip (arrow). 3, 4, Tips of chromosome arm 2L and arm 



species namely dims A, B and C (Peyton, 
unpublished data; Baimai et al., 1981; Hii, 
1982; Wibowo et al., 1984). The present 
investigation confirms those findings. Fur- 
ther, the present cytogenetic evidence has 
confirmed the fourth species recognized 
morphologically by Peyton (unpublished) 
within this taxon. Provisionally this species 
is designated dims D, and it is represented 
by the RN and PG isoline strains from 
southern Thailand populations. Our results, 
however, seem to be in disagreement with 
the interpretation of Kanda et al. (1981). 
Based on their hybridization data, Kanda 
and co-workers are of the opinion that their 
colony strains from Chantaburi, Kanchan- 
aburi and IMR only represent geographical 
populations o{ An. balabacensis. 

The results of our cross mating experi- 
ments clearly indicate that An. dims is ac- 
tually a cluster of closely related species. 
The divergence of these siblings could have 
occurred comparatively recently. The pres- 
ent data suggest that dims A and dims C 
are very closely related and they occur sym- 
patrically, at least in Kanchanaburi prov- 
ince. Artificial mating between them is pos- 
sible under laboratory conditions. Whether 
gene flow between these two genetic species 
occurs in nature is not known. The species 
isolating mechanism for these two siblings 
probably involves premating isolation, as 
well as the unidirectional genetic incom- 
patibility detected in this study. 

Both dims A and dims C exhibit similar 
banding sequences of salivary gland poly- 
tene chromosomes and general mitotic 
karyotypes, although the former shows het- 
erochromatin variation in the sex chro- 
mosomes (Baimai et al., 1984). However, 
striking differences in the polytene chro- 
mosomes of Fi hybrids were observed at the 
tips of chromosome X and chromosome 

arms 2L and 2R. Furthermore, differences 
in the amount of heterochromatin are no- 
ticeable in the sex chromosomes (Wibowo 
et al., 1984). In general, the mechanism in 
the process of species differentiation for sib- 
lings in the Dirus Complex resembles that 
for some species groups of the picture- 
winged Hawaiian Drosophila (Ohta, 1980; 
Carson and Yoon, 1982). 

Anopheles dims B (= Perils form) is ge- 
netically distinct from dims A and dims C. 
Hybridizations between dims B and dims 
C were cross-sterile, producing no adult F, 
hybrids in either direction. Cross mating be- 
tween dims B and dims A, however, yielded 
very few adult F, hybrids, of which the males 
were completely sterile. Anopheles dims B 
showed cytological differences from dims A 
both in salivary gland polytene chromo- 
some (Fig. 5) and mitotic karyotype (Baimai 
et al., 1981). So far, distribution records in- 
dicate that dims B is confined to southern 
Thailand. Thus, dims B may be isolated 
geographically from dims A. On the other 
hand, dims A seems to be widespread in 
central and northern Thailand. We now are 
investigating the boundary limits of dims A 
and dims B in southern peninsular Thai- 
land. The distribution of dims C is limited 
to collection areas in Kanchanaburi Prov- 
ince, in western Thailand where it coexists 
with dims A and dims D, and an isolated 
questionable area in southern Thailand 
where it may be sympatric with dims B. 

The highest degree of genetic incompat- 
ibihty was found in all combinations of hy- 
bridization tests involving dims D. Most 
cross matings involving the RN or PG 
strains of dims D completely failed to pro- 
duce Fi hybrid adults. However, cross mat- 
ings between female dims D (RN) x male 
dims A produced a few Fi hybrid females 
which were very weak. Asynapsis in F, lar- 

2R, respectively (chromosomes of dirus C are indicated by arrows). Fig. 5. Larval salivary gland chromosome 
elements of a F, hybrid female from a cross between female dims A x male dirus B (PR) showing extensive 
regions of asynapsis. 

6 \| X •• 

1 4 



Figs. 6-10. Salivary gland polytene chromosomes and mitotic karyotypes of F, hybrid larvae from crosses 
between female dims A x male dims D (RN). 6, Asynaptic condition of the whole polytene chromosome 
complement. 7, A fixed inversion difference on the distal half and the complete asynaptic region of zone 6 of 
the X chromosome (arrows). 8, Asynapsis in chromosome arm 2R. 9, Comparison of heterochromatin differences 
of the X chromosomes in a F, female larval neuroblast cell. 10, A short Y chromosome of dims D compared 
with the X chromosome of dims A in a F, hybrid male. 



val polytene chromosomes was extensive, 
covering more than 90% of the chromo- 
some elements. Our data indicate that dims 
D is genetically remote from the other sib- 
lings in the Dirus Complex. Anopheles dirus 
D apparently is distributed widely in central 
and southern Thailand, and northern Ma- 
laysia, and has been found in sympatry with 
dirus A, dirus B and dirus C. 

The use of heterochromatic variation in 
sex chromosomes as a means for routine 
identification of our material from the field 
has distinct limitations. On the other hand, 
analysis of salivary gland polytene chro- 
mosomes now provides a better means of 
species identification of this sibling species 
complex than mitotic karyotypes, and this 
method is used routinely in our laboratory. 
First, X chromosome heterochromatin 
variation cannot be used routinely because 
it is difficult to score and is seen only in rare, 
superb preparations. The Y chromosome 
variation, on the other hand, is scored much 
more easily and is available for routine 
identification of families from wild-caught 
material. There is a quantitative difference 
in data from these two sources of variation. 
The X chromosome data can provide direct 
evidence for gene flow characteristics in na- 
ture and thus, evidence for mixtures of cryp- 
tic species in samples. The Y chromosome 
data cannot provide such evidence because 
of the combination of the obvious hemi- 
zygous condition of the Y in males and the 
knowledge that most female anophelines are 
mated successfully only once. Consequent- 
ly, different Y chromosomes are not ex- 
pected to occur together in single broods, 
and we cannot tell from their distribution 
in broods whether these Y chromosomes 
represent intra- or interspecific variation. 
The best we can do is to correlate Y chro- 
mosome variation with primary evidence 
for the different species within the Dirus 
Complex. The most widely used criterion is 
interspecific sterility as seen in laboratory 
crossing experiments. 

Our procedure has been to score Y chro- 

mosome variation in samples from nature 
and where a sample or sub-sample of fam- 
ilies shows the same Y chromosome, cross 
one of these families to laboratory reference 
stocks. At first, we were forced to use the 
non-isoline colonies of the Bangkok strain 
(species A) and the Perils form (species B). 
As identified isofemale lines became avail- 
able, we replaced the non-isoline colonies 
as reference stocks and also established iso- 
female lines as reference stocks for species 
C and D. 

The recognition of the existence of cryptic 
species within the taxon An. dirus has led 
to a better understanding of the process of 
species differentiation of the Leucosphyrus 
Group of Anopheles. Further information 
on species distributions, behavior and pop- 
ulation dynamics of these siblings undoubt- 
edly will lead to a better understanding of 
malaria transmission and strategies for ef- 
fective vector control in this region. Differ- 
ences in biological properties and behavior 
with respect to the vectorial capacity and 
the epidemiological significance of the four 
member species of the Dirus Complex are 
under investigation. Presently, the identi- 
fication of these genetic species from natural 
samples is a problem. A practical taxonom- 
ic key is now being developed at the Walter 
Reed Biosystematics Unit (Peyton, personal 
communication). Another technique which 
may be valuable in identifying these species 
is recombinant DNA for species specific 
DNA probes. 


This investigation was supported partial- 
ly by the UNDP/Worid Bank/WHO Special 
Program for Research and Training in 
Tropical Diseases and Mahidol University 
Fund. The reviews of the finished manu- 
script by R. A. Ward and E. L. Peyton are 
greatly appreciated. We thank C. A. Green 
for comments on the manuscript and the 
entomological field staff" and the Medical 
Audiovisual Section of the AFRIMS for 
technical assistance. 



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sibling speciation in the balabacensis complex of 
Southeast Asia (Diptera: Culicidae). Abst. 10th 
Intern. Cong. Trop. Med. Malaria, 9-15 Nov., 
Manila, pp. 83-84. 

Baimai, V., R. G. Andre, and B. A. Harrison. 1984. 
Heterochromatin variation in the sex chromo- 
somes in Thailand populations oi Anopheles dims 
A (Diptera: Culicidae). Can. J. Genet. Cytol. 26: 

Baimai, V., B. A. Harrison, and V. Nakavachara. 1 980. 
The salivary gland chromosomes of Anopheles 
(Cellia) dims (Diptera: Culicidae) of the Southeast 
Asian Leucosphyrus Group. Proc. Entomol. Soc. 
Wash. 82: 319-328. 

Baimai, V., B. A. Harrison, and L. Somchit. 1981. 
Karyotype differentiation of 3 anopheline taxa in 
the Balabacensis complex of Southeast Asia (Dip- 
tera: Culicidae). Genetica 57: 81-86. 

Bryan, J. H. and M. Coluzzi. 1971. Cytogenetic ob- 
servations on Anopheles farauti Laveran. Bull. 
W.H.O. 45: 266-267. 

Carson, H. L. and J. S. Yoon. 1982. Genetics and 
evolution of Hawaiian Drosophila, pp. 297-344. 
In Ashbumer, M., H. L. Carson and J. N. Thomp- 
son, eds.. The genetics and biology of Drosophila, 
Vol. 3b. Academic Press. New York, NY. 

Coluzzi, M., A. Sabatini, V. Petrarca, and M. A. Di 
Deco. 1979. Chromosomal differentiation and 
adaptation to human environments in the Anoph- 
eles gambiae complex. Trans R. Soc. Trop. Med. 
Hyg. 73: 483-497. 

Dobzhansky, T. 1970. Genetics of the Evolutionary 
Process. Columbia Univ. Press. New York, 505 

Green, C. A. and V. Baimai. 1984. Polytene chro- 
mosomes and their use in species studies of ma- 
laria vectors as exemplified by the Anopheles mac- 
ulatus complex. In B. C. Joshi, R. P. Sharma, H. 
C. Bansal, and V. L. Chopra, eds.. Genetics: new 
frontiers. Proc. XVth Int. Cong. Genet. Vol. 3: 
89-97. Oxford and IBH Publ. Co., New Delhi. 

Green, C. A. and S. J. Miles. 1980. Chromosomal 
evidence for sibling species of the malaria vector 
Anopheles (Cellia) culicifacies Giles. J. Trop. Med. 
Hyg. 83: 75-78. 

Green, C. A., V. Baimai, B. A. Harrison, and R. G. 
Andre. 1985. Cytogenetic evidence for a com- 
plex of species within the taxon Anopheles mac- 
ulatus (Diptera: Culicidae). Biol. J. Linn. Soc. 24: 

Hii, J. L. K. 1 982. Laboratory Studies of Three Mem- 
ber Species of the Anopheles balabacensis Com- 
plex (Diptera: Culicidae). Ph.D. Thesis. Univer- 
sity of London, 253 pp. 

. 1984. Involvement of the X-chromosome in 

hybrid male sterility from crosses between species 

A and species B of the taxon Anopheles dims. Mosq. 
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Kanda, T., K. Takai, G. L. Chiang, W. H. Cheong, and 
S. Sucharit. 1981. Hybridization and some bi- 
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89(1), 1987, pp. 167-174 




Paul J. Spangler and Richard C. Froeschner 

Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. 

Abstract.— Tht micro veHine water-bug Microvelia ayacuchana Drake and Maldonado 
Capriles [sic] (1952) is illustrated for the first time. A habitus view, the antennae, and the 
male and female genitalia are illustrated by pen and ink drawings and the surface sculpture 
of selected parts of the body are illustrated by scanning electron micrographs. Known 
distribution records for this species from Brazil, Guyana, Surinam, and Venezuela are 
reported. Habitats are discussed and a photograph of one of the biotopes is included. 

During a fieldtrip to collect aquatic Co- 
leoptera, Hemiptera, and other insects in 
the Takutu Mountains of Guyana in No- 
vember and December, 1983, numerous 
aquatic Hemiptera were collected by mem- 
bers of an Earthwatch Expedition. Among 
the water bugs were 10 distinctive winged 
specimens of a large species of Microvelia. 
An examination of those specimens and 
comparison with the type specimens of Mz- 
crovelia in the Drake collection and the gen- 
eral collection of aquatic Hemiptera in the 
U.S. National Museum of Natural History, 
Smithsonian Institution confirmed that the 
specimens from Guyana are conspecific with 
Microvelia ayacuchana Drake and Maldo- 
nado Capriles [sic] (1952). More recently, 
19 and 24 February 1986, Spangler collect- 
ed 22 more specimens of this species near 
the type locality, Puerto Ayacucho, T.F.A., 

Because M. ayacuchana is rare in collec- 
tions; belongs to a large genus with 8 1 de- 
scribed species in the Western Hemisphere 
that are similar in external morphology, 
sculpture, and color; has not been previ- 
ously illustrated; and has not been included 

in any keys; we have prepared this article 
with the hope that it will allow the reader 
to identify this species more easily. 

The type specimens of M. ayacuchana 
were collected by J. Maldonado at Puerto 
Ayacucho, Venezuela, in May and June 
1950; and Drake and Hussey ( 1955), in their 
checklist of the species of Microvelia, re- 
ported M. ayacuchana from British Guiana 
[now Guyana]. There are no specimens of 
the species from "British Guiana" in the 
NMNH but the record may have been based 
on borrowed specimens; unfortunately, the 
source of the specimens was not given. There 
has been nothing reported about M. aya- 
cuchana since Drake and Roze (1958) Hsted 
it as one of the seven species of Microvelia 
reported from Venezuela. 

The description of Microvelia ayacu- 
chana by Drake and Maldonado Capriles 
[sic] ( 1 952) is adequate for the external mor- 
phological characters as seen under a ste- 
reoscopic microscope. Additional external 
characters, some illustrated by scanning 
electron micrographs, and genitalic char- 
acters are discussed below. A habitus view 
(Fig. 1 ) and the dissected and cleared male 



Fig. 1 . Microvelia ayacuchana, male, habitus view. 

genitalia (Figs. 13, 14) and female genitalia 
(Figs. 15, 16) of M ayacuchana are illus- 
trated for the first time. 

Microvelia ayacuchana Drake and 
Maldonado Capriles [sic] 

Additional descriptive characters. — Mac- 
ropterous adult males longer (2.75 to 3.16 
mm) than most members of genus. Head 

(Fig. 2) with longitudinal groove on midline 
distinct, 3 pairs of trichobothria, ocelli ab- 
sent. Antennae (Fig. 3) long; 4 segmented; 
intemodial piece between segments 2 and 3 
and another between segments 3 and 4; an- 
tennal segment ratios 17:13:17:23. Rostrum 
3 segmented, extending to mesocoxae; man- 
dibular stylets toothed (Fig. 4). 
Mesoscutellum covered by posterior ex- 


Figs. 2-6. Microvelia ayacuchana. 2, Head. 3, Antenna. 4, Mandibular stylets. 5, Head, pronotum, and 
evaporatorium tuft. 6, Methathoracic scent gland channels. 







Figs. 13, 14. Microvelia ayacuchana. 13, Male genitalia, extended in alcohol, dorsal view. 14, Male genitalia, 
cleared in KOH, lateral view. 

Figs. 7-12. Figs. 7-10. Microvelia ayacuchana. 7, Campaniform sensillum, base of protarsal segment. 8, 
Campaniform sensillum enlarged. 9, Protibial grooming comb (arrow) and grasping combs, male. 10, Protibial 
grasping comb, male. Figs. 11, 12. Microvelia ayacuchana. 11, Protibial grasping comb, male, enlarged. 12, 
Protarsal claws. 




Figs. 15, 16. Micwvelia ayacuchana. 15, Female genitalia, extended in alcohol, dorsal view. 16, Female 
genitalia, lateral view. 



Fig. 1 7. Microvelia ayacuchana, pothole biotope at Tobogan area, 40 km south of Puerto Ayacucho, T.F.A., 

tension of pronotal lobe (Fig. 5). Metatho- 
racic scent gland channels (Fig. 6) terminate 
in an evaporatorium bearing a prominent 
tuft of long setae (Fig. 5, arrow) on pleuron 
in front of metacoxae. 

Protarsus with 1 campaniform sensillum 
dorsally at base of ultimate segment (Figs. 
7, 8). Front tibia of male with a short, trans- 
verse, apical, grooming comb of stout setae 
on dorsal margin (Fig. 9, arrow) and a long 
grasping comb (Figs. 9-11) of short stout 
spines on inner (ventral) surface of distal 
third. Claws (Fig. 1 2) distinctly inserted be- 
fore apex of last tarsal segment. 

Male genitalia (Figs. 13, 14) with aedea- 
gus with membranous vesica; parameres 
minute. Female genitalia (Figs. 15, 16) with 
proctiger broad, decurved, covering gono- 
coxal and genital opening. 

Specimens examined. — Specimens from 
the John T. Polhemus collection are indi- 
cated by (JTP) and those from the National 
Museum of Natural History, Smithsonian 
Institution are indicated by (NMNH). Rep- 
resentative specimens will be deposited in 
the Instituto de Zoologia Agricola, Univer- 

sidad Central de Venezuela, Maracay, Ven- 

BRAZIL: AMAZONAS: Lago Salgado, 
Jg. duebra d., 29.4.48, H. Sioh, 2 males 
UNI DISTRICT: Takutu Mountains, 
6°1 5'N 59°5"W, 3-10 Dec. 1 983, P. J. Span- 
gler, R. A. Faitoute, P. D. Perkins, 3 males, 
7 females ( 1 on SEM stub) (NMNH). SURI- 
NAM: Sipaliwini River, 13-VI-63, 6 males, 
6 females (NMNH). VENEZUELA: TER- 
to Ayacucho, 15 June 1950, J. Maldonado 
Capriles, holotype male, allotype (NMNH); 
same data, 1 male, 2 females (paratypes) 
(JTP); Puerto Ayacucho (40 Km S) at To- 
bogan, 19 Feb. 1986, P. J. Spangler, 1 male, 
1 female (NMNH); same data except 24 Feb. 
1986, 16 males, 4 females (NMNH). One 
additional paratype with the same data as 
the holotype except 15 May 1950 has not 
been found. 

Habitat.— The specimens from Guyana 
were collected from the shaded margin of a 
slowly flowing brook in the rainforest. The 
series from the Tobogan area south of Puer- 



to Ayacucho, Venezuela, were found on the 
water of a small pothole (Fig. 1 7) in bedrock 
beside a small stream; the pothole was ex- 
posed to full sunlight for a relatively short 
time during the day. 


We thank the following for their assis- 
tance: The Center for Field Research, for 
sponsoring the fieldwork in Guyana; the ad- 
ministrators of the Smithsonian Institu- 
tion's Research Opportunities Fund for 
supporting the fieldwork in Venezuela; 
Young T. Sohn, biological illustrator, for 
the pen and ink drawings; John T. Polhe- 
mus, for locality data from his specimens; 

and Phyllis Spangler for typing the manu- 
script into the word processor. 

Literature Cited 

Drake, C. J. and J. Maldonado Capriles. 1952. Water- 
striders from Territorio Amazonas of Venezuela 
(Hemiptera: Hydrometridae, Veliidae). The Great 
Basin Naturalist 12: 47-54. 

Drake, C. J. and R. F. Hussey. 1955. Concerning the 
Genus Microvelia Westwood, with Descriptions 
of Two New Species and a Check-list of the Amer- 
ican Forms (Hemiptera: Veliidae). The Florida 
Entomologist 38: 95-115. 

Drake, C. J. and J. A. Roze. 1958. A new Microvelia 
from Venezuela (Hemiptera: Veliidae). Bulletin of 
the Southern California Academy of Sciences 57: 


89(1), 1987, pp. 175-176 


J. F. Gates Clarke 

Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. 

Abstract.— The true identity of Acleris inana (Robinson) is revealed and the genitalia 
are figured. 

While curating the species of Acleris in 
the collection of the United States National 
Museum of Natural History it soon became 
abundantly clear that the identity of several 
species was confused, and that many series 
were mixed. Contributing to this confusion 
is the fact that the phenotypes are frequently 
repeated in several taxa and the study of the 
genitalia is the only means of correctly iden- 
tifying the species. 

Among the similarly marked species is 
Acleris inana (Robinson) (1869: 281) de- 
scribed as Teras inana. The number of spec- 
imens in his series was not indicated al- 
though he had at least two because he stated 
"Habitat. — Mass., N.Y." This species be- 
longs to a group of closely similar species 
consisting of A. inana (Robinson), A. fla- 
vivittana (Clemens), A. robinsoniana 
(Forbes), and perhaps others. 

Of inana, McDunnough (1934) stated that 
the ". . . determination of this species is 
based on a specimen compared by the writer 
in 1925 with a series in the United States 
National Museum under this name, and by 
a recent examination of a specimen in the 
American Museum Collection labelled 
"Homotype" by Kearfott." Unfortunately, 
McDunnough did not see or examine the 
Lectotype, which is in the Academy of Nat- 
ural Sciences, Philadelphia, but based his 
identification on specimens that appear to 
have been incorrectly identified. Mc- 

Dunnough's figure of the female genitalia 
(1934, p. 331, fig. 1) which he supposed to 
be inana. does not agree with the type, and 
I have not yet been able to associate defi- 
nitely any species with his figure. Subse- 
quently, Razowski (1966) figured as inana 
(fig. 526) what appears to be exactly the same 
as that figured by McDunnough (see above). 
In neither case have I been able to ascertain 
what species they illustrated. 

So far, I have been able to associate def- 
initely with inana, two females from Mis- 
souri, Jasper County, Sarcoxie, 14 June 
1975, in the J. R. Heitzman collection and 
four specimens in the United States Na- 
tional Museum as follows: One male and 
two females are labeled "NY." The male 
bears a further label inscribed "agrees per- 
fectly with type" and one female bears a 
label "Mass. Agrees with type." The lecto- 
type, designated by Klots (1942) and now 
in the Academy of Natural Sciences, Phila- 
delphia, bears the following label data: 
Small, square white label "36." Small white 
label "NY." Red label: Teras inana C. T. 
Robinson, 7439." Green label, genitalia slide 
by JFGC, female, USNM 25553. The gen- 
italia of the lectotype are illustrated in Fig. 1 . 

I am uncertain whether Acleris flavivit- 
tana (Clemens) is a distinct species. The type 
of Jlavivittana, apparently, is lost. The fe- 
male genitalia of a series of specimens in 
the United States National Museum from 




Fig. 1 . Acleris inana. a 
part of ductus bursae. 

Ventral view of female genitalia, b, Enlarged view of sterigma, ostium and posterior 

Pennsylvania, Virginia, and Washington 
state identified asflavivittana are practically 
identical to those of inatia. Until such time 
as the status of flavivittana can be estab- 
lished beyond a doubt I am retaining the 
two names. 

I wish to thank the authorities of the 
Philadelphia Academy of Sciences for the 
loan of the type of Acleris inana (Robinson). 
Also I wish to thank Victor Krantz for the 
photographs and Silver West for typing the 
manuscript, both of the Smithsonian staff. 

Literature Cited 

Klots, A. B. 1942. The Material of North American 
Microlepidoptera Other Than Aegeriidae in the 

American Museum of Natural History. Bulletin of 
the American Museum of Natural History 79: 391- 

McDunnough, J. 1934. The Canadian Species of the 
Tortricid Genus Peronea. Canadian Journal of Re- 
search 11: 290-332. 

Powell, J. A. 1964. Biological and Taxonomic Studies 
on Tortricine Moths, with Reference to the Species 
in California. I-IV. 1-317, 15 maps, 108 figures, 
8 plates. 

Razowski, J. 1966. World Fauna of the Tortricini 
(Lepidoptera, Tortricidae). Zaklad Zoologii Sys- 
tematycznej W. Krakowie. Polskiej Akademii 
Nauk. 1-575, 832 figures, 41 plates. 

Robinson, C. T. 1869. Notes on American Tortri- 
cidae. Transactions of the American Entomolog- 
ical Society 2: 261-288, pis. 1, 4-8. 


89(1), 1987, pp. 177-184 


R. D. Waltz and W. P. McCafferty 

Department of Entomology, Purdue University, West Lafayette, Indiana 47907. 

Abstract. —The genus Cloeodes is redefined to include species with hindwings and species 
without hindwings. Notobaetis Morihara and Edmunds (1980) and certain species pre- 
viously assigned to Centroptella Braasch and Soldan (1980), including the type species, 
are congeneric with Cloeodes, necessitating the following recombinations: Cloeodes penai 
(Morihara and Edmunds) n. comb., Cloeodes longisetosus (Braasch and Soldan) n. comb., 
and Cloeodes soldani (Miiller-Liebenau) n. comb. A new Neotropical genus, Bernerius n. 
gen., and its type species, B. incus n. sp., from Peru are described. Bernerius is a sister 
genus of Cloeodes. Species previously assigned to the ceylonensis-similis-pusilla species 
group of Centroptella are a monophyletic cluster unrelated to Cloeodes (= Centroptella 
Braasch and Soldan, in part) and are therefore placed in Chopmlla n. gen. that includes: 
Chopralla ceylonensis (Miiller-Liebenau) n. comb. (= type species), Chopralla similis 
(Miiller-Liebenau) n. comb., and Chopralla pusilla (Miiller-Liebenau) n. comb. 

Cloeodes was erected by Traver ( 1 938) to 
include several Puerto Rican species of Bae- 
tidae with adults very similar to Nearctic 
species assigned to Pseudocloeon Klapalek 
(both having paired marginal intercalaries 
of the forewings and no hindwings) and 
larvae distinct from any known Nearctic 
species assigned to Pseudocloeon {Cloeodes 
having a well-developed median terminal 
filament). Our revisionary study of Baetidae 
has shown Cloeodes to be a distinct genus 
possessing several apomorphic character 
states. We redefine and describe it herein 
along with describing a new sister genus of 
Cloeodes. The revision of Cloeodes has ne- 
cessitated the revision, and hence dissolu- 
tion, of the genus Centroptella Braasch and 
Soldan because certain of those species, in- 
cluding the type species of Centroptella, are 
actually Cloeodes. Other unrelated Cen- 
troptella are placed in an additional new and 
distinctive genus that is also described 

Genus Cloeodes Traver 

Cloeodes Traver, 1938: 32. Type species 
Cloeodes maculipes Traver, by original 

Notobaetis Morihara and Edmunds, 1980: 
606. Type species A^. penai Morihara and 
Edmunds, by original designation, n. syn. 

C6'«rra/?r£'//a Braasch and Soldan, 1980: 123. 
(in part). Type species C longisetosa 
Braasch and Soldan, by monotypy. n. syn. 

Larva. — Labrum slightly broader than 
long, clearly emarginate and with marginal 
shelf anteriorly. Left mandible (Fig. 3) with 
incisors fused apically; prostheca stout, dig- 
itate; no tuft of setae between incisors and 
molar area; thumb of molar area triangulate 
and elevated above plane of incisor base. 
Right mandible (Fig. 4) with incisors sep- 
arated apically; prostheca reduced, slender, 
and variably furcate. Labium with 3-seg- 
mented palps; terminal segment rounded to 
oblique; segment 2 with weakly developed 



Figs. 1-5. Cloeodes spp. l,Forewing. 
setal tuft. 

I, Metascutellar hump. 3, Left mandible. 4, Right mandible. 5, Ventral 

inner apical lobe; glossae subequal to para- 
glossae; paraglossae with subparallel mar- 

Femora parallel sided, without ventral se- 
tal patch and with short, dorsal bristles and 
similarly shaped distal bristles. Tibiae with 
subproximal arc of long, fine setae; no fine 
setae adjoining tibial seam. Foreleg with 
subtending bristle and tibial seam and with 
subproximal arc of long, fine setae. Claws 
(Fig. 17) ca. 0.33 X tarsal length, with or 
without microspines. 

Abdominal terga with broadly pointed, 
rectangulate-based scales and fine setae; 
scales with median length subequal to basal 
width; posterior marginal spines present on 
all terga; ventrally abdomen with scales and 
fine setae, with prominent tufts (i.e. with 
contiguous setal bases) of long, fine setae on 

segments 2-6 (Figs. 5, 18). Gills (similar to 
Fig. 13) 1-7, asymmetric, broadly pointed, 
without marginal spination or ciliation; each 
gill often 2 X or more length of associated 
tergum. Median terminal filament subequal 
to cerci. 

Adult male. — Forewings (Fig. 1) with 
paired marginal intercalaries in most cells; 
detached base of vein MA. extending well 
beyond 0.5 x distance between distal cross- 
vein and proximal (= MA, to MP;) cross- 
vein. Hindwings present or absent; when 
present with acute or distinctly hooked me- 
dian costal process. Posterior margin of 
metanotum deeply emarginate; metascutel- 
lar hump (Fig. 2) not flattened before apex 
but projecting dorsoposteriorly. Forceps 
four-segmented; segments 2 and 3 nearly 
fused; segment 3 ca. 2 x length segment 2, 



segment 2 subequal to segment 1 and with 
basal bulge bearing fine bristle-like setae; no 
median spine or protuberance between for- 
ceps bases. 

Adult female. — Marginal intercalaries of 
forewing paired or less often single. Poste- 
rior margin of metanotum not as deeply 
emarginate as in male; metascutellar pro- 
cess as in male. Subanal plate not devel- 

Material examined. — CI oeodes macu- 
lipes Traver: Holotype, 5 adult, Puerto Rico, 
Ludillo Mtns., camp lab 46 (107), VI- 14- 
1935. J. Garcia-Diaz. in alcohol, genitalia 
slide mounted in balsam (solvent: xylene); 
larval exuviae of holotype 5 slide mounted 
in euparal (solvent: abs. ale), C.U. Type 
No. 1402.1, Cornell University. Paratypes, 
S adult (genitalia missing), subimago exu- 
viae, larval exuviae (missing), same data and 
deposition as above; paratype (allotype), 9 
adult, in alcohol; wing slide mounted, Puer- 
to Rico, Trout's pool. El Yunque Trail, VI- 
12-1935. J. Garcia-Diaz. C.U. Type No. 
1402.2, same deposition as holotype. 
Cloeodes consignatus Traver: Holotype, $ 
adult, Puerto Rico, Yunez River, VI-21- 
1935, J. Garcia-Diaz. Cornell University 
Type No. 1403.1, in alcohol; one wing slide 
mounted. Notobaetis penai Morihara and 
Edmunds: Paratypes, 5 larva, Argentina, 
Tucuman N.W. of San Miguel de Tucuman, 
1-25-1969, W. L. and J. G. Peters, slide 
mounted in balsam (solvent: xylene), Pur- 
due Entomological Research Collection 
(PERC); 9 larva, Argentina, Cordoba Prov., 
Copina (ca. 25 km WNW Alta Garcia), elev. 
1650 m, IV-1 1/14-1969, L. Pena, slide 
mounted in balsam (solvent: xylene) 
(PERC); 9 adult, legs slide mounted in bal- 
sam (solvent: xylene), same data as above 
(PERC); 5 adult, wings slide mounted, same 
data as above (PERC); three whole larvae 
(in alcohol), same data as above, Florida 
A&M University Collection; two adult $ and 
one subimago $ and two associated larval 
exuviae (in alcohol), same data as above. 
University of Utah Collection. Centwptella 

longisetosa Braasch and Soldan: paratype 
larva (in alcohol). Peoples Republic of 
China, Liu Chui, Kuj Fon Shan River, 1 1 - 
12-1959, I. Hrdy, deposited Purdue Ento- 
mological Research Collection, originally 
from paratypes in the collection of T. Sol- 
dan. Centroptella soldani Muller-Liebenau: 
paratype larva (in alcohol), Ceylon, EC/ 
11/c Rasanawa-Fall, Ratnapura, XI- 19- 
1970. F. Starmiihlner deposited Florida 
A&M University. 

Remarks.— Although previously known 
only from Puerto Rico, a species revision 
of this genus by Waltz and McCafferty (in 
press) indicates that Cloeodes is a wide- 
spread, primarily tropical taxon of Neo- 
tropical origin, ranging from central Argen- 
tina to southwestern United States and 
southeastern Asia. Notobaetis Morihara and 
Edmunds and certain species of Centrop- 
tella Braasch and Soldan are clearly con- 
generic with Cloeodes Traver as indicated 
by the synapomorphic possession of ventral 
setal tufts on abdominal segments 2-6. 
Nominal species of the genus at this writing 
include Cloeodes consignatus Traver, C 
longisetosus (Braasch and Soldan) n. comb., 
C maculipes Traver, C. penai (Morihara 
and Edmunds) n. comb., and C. soldani 
(Muller-Liebenau) n. comb. We are describ- 
ing additional species in our revision of 
Cloeodes. Taxonomic and phyletic relation- 
ships o{ Cloeodes with the additional genera 
described herein is treated under those gen- 

Bernerius Waltz and McCafferty, 

New Genus 

Larva. — Labrum (Fig. 6) slightly broader 
than long, clearly emarginate and with mar- 
ginal shelf anteriorly. Left mandible (Fig. 7) 
with incisors fused apically; prostheca stout, 
digitate; no tuft of setae between incisors 
and molar area; thumb of molar area trian- 
gulate and not elevated above plane of in- 
cisor base. Right mandible (Fig. 8) with in- 
cisors separated apically; prostheca reduced, 
slender and furcate; no tuft of setae between 



Figs. 6-1 3. Bemerius incus larva. 6, Labrum (left, ventral; right, dorsal). 7, Left mandible. 8, Right mandible. 
9, Maxilla. 10, Labium (left, ventral; right, dorsal). 11, Foreleg (db-distal bristles, pa-proximal arc, ts-tibial 
seam, sb-subtending bristle). 12, Claw (ms-microspines). 13, Gill 4. 



incisors and molar area. Labium (Fig. 10) 
with three-segmented palps; terminal seg- 
ment obliquely truncate; segment 2 with 
weakly developed inner apical lobe; glossae 
subequal to paraglossae; paraglossae with 
weakly convex subparallel margins. 

Femora (Fig. 1 1 ) parallel sided, without 
ventral setal patch, and with short dorsal 
bristles and similarly shaped distal bristles. 
Tibiae with subproximal arc of long, fine 
setae; no fine setae adjoining tibial seam. 
Foreleg with subproximal arc of setae, sub- 
tending bristle, and tibial seam. Claws (Fig. 
12) ca. 0.33 X tarsal length, excavate and 
with microspines basally. 

Abdominal terga with rectangulate-based 
scales and fine setae; scales with median 
length ca. 2x basal width; posterior mar- 
ginal spines present on all terga; ventrally 
abdomen with scales and fine setae, no setal 
tufts. Gills (Fig. 13) 1-7 asymmetric, elon- 
gate, and broadly pointed apically; without 
marginal spination or ciliation. Median ter- 
minal filament subequal to cerci. 

Type species.— Bernerius incus Waltz and 
McCafferty n. sp. 

Etymology .— Bernerius is of masculine 
gender and is based on the sur-name of the 
American ephemeropterist Lewis Bemer. 

Remarks.— This genus is erected for a Pe- 
ruvian species previously treated as Baetis 
sp. B by Bemer (in Roback et al., 1 980) and 
redescribed as Bemerius incus n. sp. below. 
Adults are presently unknown. 

Bemerius is a sister genus to Cloeodes as 
demonstrated by the following synapomor- 
phies: the presence of a left mandible (Figs. 
3, 7) with incisors fused apically and a stout 
prostheca; a right mandible (Fig. 4, 8) with 
incisors partially fused but separated api- 
cally and a slender, furcate prostheca; the 
presence of an arc of long, fine setae sub- 
proximally on the tibia (Fig. 1 1-pa); simple, 
asymmetrically lamellate, and broadly 
pointed gills (Fig. 13). Bemerius retains the 
pleisomorphous condition of abundant long, 
fine setae on the venter of most abdominal 
segments, which in Cloeodes species form 

distinct tufts (Figs. 5, 1 8) on abdominal seg- 
ments 2-6. 

The elevation of the left mandible thumb 
above the plane of the incisor bases, the 
presence of clearly defined setal tufts on 
sterna 2-6, reductionist tendencies in spi- 
nation of the legs, increased stability in the 
length of posterior marginal row spines and 
differences in the degree of convexity of the 
exterior margin of the paraglossae (toward 
a more parallel and less convex condition) 
are phenoclines clearly differentiating lar- 
vae of the relatively more derived Cloeodes 
species from Bemerius. 

The similarity of Bemerius especially to 
plesiomorphic Cloeodes species suggests a 
Neotropical origin for Cloeodes. Species of 
Cloeodes we have studied that possess 
hindwing pads are apparently restricted to 
South America. Apomorphic species, in- 
cluding those to be described from North 
America and those transferred herein, C 
longisetosus (China) and C. soldani (Sri 
Lanka), have lost the hindwings. 

Bemerius incus Waltz and McCafferty, 

New Species 

Baetisp.) sp. B. Bemer, 1980: 190. 

Larva. — Body 10-11 mm long. Head cap- 
sule lightly pigmented, pale near oral mar- 
gin and frontal area below antennal bases. 
Antennae pale, tapered, ca. 3 x head capsule 
in length; scape subequal to pedicel in length; 
flagellar segments darkened apically. La- 
brum (Fig. 6) with 1 + 2-3 subapical setae. 
Maxillary palp (Fig. 9) exceeding galea-la- 
cinea, with evidence of third segment; palp 
with numerous fine setae. Segment 2 of la- 
bial palp (Fig. 10) ca. 1.25 x length of seg- 
ment 3 and with 6-8 dorsal setae; apex of 
segment 3 obliquely truncate with inner 
marginal stout bristles; paraglossa with 3 
apical rows of setae, 4-5 ventral intero-mar- 
ginal fine setae, and 5-6 dorsal intero-mar- 
ginal setae medioapically; glossa with ca. 20 
stout inner marginal setae. 

Prothorax lightly pigmented with medial 



dark hourglass marking; meso- and meta- 
thorax lightly pigmented, without pattern- 
ing; hindwing pads present. Legs (Fig. 1 1) 
lightly pigmented, tibiae and tarsi darker 
than femora; trochanters with 20 or more 
laterally serrated blade-like bristles and nu- 
merous fine setae, without scales; femora 
with many pointed, laterally serrate ventral 
bristles, with fine setae and scales, with short, 
blunt dorsal bristles, and with 6-7 stout, 
pointed, distal setae; tibiae with bristle-like 
setae dorsally and ventrally, (subtending 
bristle blade-like as other tibial setae); tarsi 
with scales, fine setae, and many bristle-like 
stout setae dorsally and ventrally, without 
dorsal row of fine setae. Claws (Fig. 12) ca. 
0.33 X tarsal length, with basal microspines. 

Dorsal abdomen lightly patterned with 
brown medially; segment 1 uniformly col- 
ored; segment 2 with pale brown trapezoidal 
medial patch; segments 3, 4, and 5 with pale 
broken coloration; segments 6 and 7 with 
dark trapezoidal medial patch; segments 8- 
10 pale to cream. Posterior marginal spines 
present on all tergites, basally with clear area 
as in Centwptilum sp. No. 4 Demoulin 
(1970), with base of each ca. 0.5 x length, 
subequal, evenly tapered; smaller spines sit- 
uated randomly throughout series. Abdo- 
men pale ventrally, with scales and long, 
fine setae randomly scattered over surface, 
without tufts of setae. Paraproct with spi- 
nous posteromedial margin (large and small 
spines alternating); surface with scales and 
fine setae. 

Type material. — Holotype, 9 larva, Peru, 
Llave River at Llave, VII-2-1977. 4000 m. 
Coll. #21. S. S. Roback, Academy of Nat- 
ural Sciences of Philadelphia. Slide 1, la- 
bium in balsam (solvent: xylene), other 
mouthparts mounted in euparal (solvent: 
abs. ale); slide 2, abdominal segments 4- 
10 mounted in euparal (solvent: abs. ale); 
slide 3, gills 3, 4, 7 mounted in balsam (sol- 
vent: xylene); slide 4, foreleg in balsam (sol- 
vent: xylene), abdominal segment 3 in eu- 
paral (solvent: abs. ale); slide 5, thorax, 
abdominal segments 1 and 2 mounted in 

euparal (solvent: abs. ale); head capsule 
(70% ETOH). 

Remarks. -Roback (Roback et al., 1980) 
described the Llave River where B. incus 
(= Baetis sp. B. Bemer) was collected as a 
large river ca. 200 meters wide and organ- 
ically enriched, presumably with the offal 
and raw waste from a local slaughter house. 
Photographs of the habitat indicate the river 
had slow to moderate current. 

Chopralla Waltz and McCafferty, 
New Genus 

Centwptella Braasch and Soldan, 1 980: 1 23 

(in part). 
Centwptella, Muller-Liebenau, 1984b: 96 

{ceylonensis-similis-pusilla species group). 
Genus No. 2 sp. 1 Muller-Liebenau, 1984a: 


Larva. — Labrum slightly broader than 
long, clearly emarginate and with marginal 
shelf anteriorly. Left mandible (similar to 
Fig. 3) with incisors fused apically; pros- 
theca stout and digitate; no tuft of setae be- 
tween incisors and molar area; thumb of 
molar area triangulate and elevated above 
plane of incisor base. Right mandible (sim- 
ilar to Fig. 4) with incisors separated api- 
cally; prostheca stout or reduced and fur- 
cate. Labium with 3-segmented palps; 
terminal segment rounded to broadly 
rounded apically; segment 2 with weakly 
developed inner apical lobe; glossae sub- 
equal to paraglossae; paraglossae with sub- 
parallel margins, not convex. 

Femora (Fig. 14) parallel sided, without 
ventral setal patch, and with long, dorsal 
bristles and similarly shaped distal bristles. 
Tibiae with subproximal arc (Fig. 1 4-pa) of 
long, fine setae and with long, fine setae ad- 
joining meso- and metatibial seams (Fig. 
14-tss). Foreleg with subproximal arc of long, 
fine setae, subtending bristle, and without 
tibial seam or with weakly developed seam 
only and with fine setae adjoining tract of 
tibial seam. Claws (Figs. 15, 16) ca. 0.33 x 
tarsal length, with 3-5 paired medial and 
apical denticles. 






Figs. 14-17. Chopralla ceylonensis larva. 14, Foreleg (db— distal bristles, pa — proximal arc, tss— tibial seam 
setae). 15, Claw (ventral) after Muller-Liebenau (1983). 16, Claw (oblique) after Muller-Liebenau (1983). 17, 
Cloeodes longisetosus larval claw (lateral). 

Abdominal terga with rectangulate-based 
and elongate, broadly pointed scales and 
with fine setae; scales with median length 
subequal to basal width or with median 
length ca. 2.0-2.5 x basal width; posterior 
marginal spines present at least on terga 9 
and 10; ventrally abdomen with broadly 
pointed, rectangulate-based scales and fine 
setae, no setal tufts. Gills 1-7 asymmetric, 
simple, and rounded apically; without mar- 
ginal spination, or ciliation. Median ter- 
minal filament subequal to cerci. 

Type species. — Centwptella ceylonensis 
Muller-Liebenau, 1983: 486. 

Etymology. — C/?o/?ra//« is of femine gen- 
der and is named in recognition of the In- 
dian ephemeropterist B. Chopra. 

Material examined. — C ceylonensis 
(Muller-Liebenau): eight larvae, Sri Lanka, 
Kitugala, Hal-oya, FC 35/a, XII-27-1970, 
det. L Muller-Liebenau, 1983. Chopralla sp.: 
four larvae, Viet Nam, Vinh Phu' Prov., 
stream nr. Tam-Dao, X- 1 0- 1 984, T. Soldan 

deposited in the Purdue Entomological Re- 
search Collection. 

Remarks.- Species herein assigned to the 
genus Chopralla are clearly separated from 
Cloeodes (= Centroptella in part) species by 
the absence of ventral tufts of setae on ab- 
dominal segments 2-6, the apically rounded 
gills (versus broadly pointed in Cloeodes 
species), the peculiar claw structure (unlike 
edentate claws of Cloeodes), and the pos- 
session of long, fine tibial seam setae (not 
present in Cloeodes species). 

Nominal species of Chopralla include the 
Sri Lankan species Chopralla ceylonensis 
(Muller-Liebenau) n. comb, and C similis 
(Muller-Liebenau) n. comb, and the East 
Malaysian species C pusilla (Miiller-Lie- 
benau) n. comb. The West Malaysian 
species. Genus No. 2 sp. 1 Muller-Liebenau 
( 1 984a) could be placed in Chopralla on the 
basis of the following apparent synapomor- 
phies: possession of Chopralla-hke claws 
with paired apical denticles, possession of 



Fig. 18. Cloeodes sp. larva. SEM of ventral setal 
tuft on left side of abdominal segment 4. 

long, fine tibial seam setae, asymmetric 
ovate gills, and fimbriate paraproct scales. 
This species differs from other known Cho- 
pralla species, however, by possessing the 
following inferred plesiomorphic charac- 
ters: 1) stout prostheca of the right mandi- 
ble; 2) posterior marginal spines present on 
segments 1-10; and 3) tergal scales with me- 
dian length subequal to basal width. Al- 
though this latter species appears to clearly 
belong to Chopralla, we do not name the 
species herein since we have not yet studied 
the single specimen described. 


We thank G. F. Edmunds, Jr., (Salt Lake 
City, Utah), W. Flowers, (Tallahassee, Flor- 
ida), J. K. Liebherr (Ithaca, New York), I. 

Miiller-Liebenau (Plon, West Germany), S. 
S. Roback (Philadelphia, Pennsylvania), T. 
Soldan (Ceske Budejovice, Czechoslova- 
kia), and Q. D. Wheeler (Ithaca, New York) 
for the loan of material; and J. Martin and 
A. V. Provonsha for Figs. 13 and 14 re- 
spectively. We especially with to thank T. 
Soldan for the gift of one paratype larva of 
Centwptella longisetosa now deposited in 
the Purdue Entomological Research Collec- 
tion (PERC), West Lafayette, Indiana. This 
paper has been assigned Purdue University 
Experiment Station Journal No. 10,436. 

Literature Cited 

Braasch,D. and T. Soldan. 1980. Centroptella n. gen., 
eine neue Gattung der Eintagsfliegen aus China 
(Baetidae: Ephemeroptera). Reichenb. 18: 123- 

Demoulin, G. 1970. Ephemeroptera des faunes 
Ethiopienne et Malgache. In South African Ani- 
mal Life, Results of the Lund University Expe- 
dition in 1950-1951. 14: 24-170. 

Morihara, D. K. and G. F. Edmunds, Jr. 1980. A^o- 
tobaetis: a new genus of Baetidae (Ephemerop- 
tera) from South America. Int. Rev. Gesampten 
Hydrob. 65: 605-610. 

Miiller- Liebenau, I. 1983. Three new species of the 
genus Centwptella Braasch and Soldan, 1980, from 
Sri Lanka (Insecta: Ephemeroptera). Arch. Hy- 
drob. 97: 486-500. 

. 1984a. New genera and species of the family 

Baetidae from West-Malaysia (River Gombak) 
(Insecta: Ephemeroptera). Spixiana 7: 253-284. 

. 1984b. Baetidae from Sabah (East Malaysia) 

(Ephemeroptera). Proc. Fourth Int. Conf Ephem- 
eroptera 1983: 85-99. 

Roback, S. S.. L. Bemer, O. S. Hint, Jr., N. Nieser, 
and P. J. Spangler. 1980. Results of the Cath- 
erwood Bolivian-Peruvian Expedition Part I. 
Aquatic insects except Diptera. Proc. Acad. Nat. 
Sci. Phila. 132: 176-217. 

Traver, J. R. 1938. Mayflies of Puerto Rico. J. Agric. 
Univ. P.R. 22: 5-42. 

Waltz, R. D. and W. P. McCafferty. 1987. Revision 
of the genus Cloeodes Traver (Ephemeroptera: 
Baetidae). Ann. Entomol. Soc. Am. (In press.) 


89(1), 1987, pp. 185-199 





W. A. Palmer 

North American Field Station, Queensland Department of Lands, 2714 Pecan Drive, 
Temple, Texas 76502. 

Abstract.— \ survey of the phytophagous insect fauna on Baccharis halimifolia and B. 
neglecta was undertaken between 1982 and 1986 as part of a program to find biological 
control agents for B. halimifolia in Australia. One hundred and thirty three phytophagous 
insect species were collected and these included 1 1 species that were considered monoph- 
agous. The monophagous group of species contained a high proportion of Lepidoptera 
and endophages, and all monophagous species were collected from B. halimifolia. Over 
50% of the monophagous group and 21% of the total number of species were found on 
both species of Baccharis. The relevance of the survey to the biological control program 
is discussed. 

The woody shrub Baccharis halimifolia 
L. (Family Asteraceae: Tribe Astereae: Sub- 
Tribe Baccharinae), which was introduced 
from North America in the latter part of the 
1 9th century (Bailey, 1 900), has become one 
of the most serious weeds in Queensland, 
Australia. The shrub invades cattle pas- 
tures, reforested areas, and disturbed sites 
and is a declared noxious weed under the 
Stock Routes and Rural Lands Protection 
Acts (1944-1967) of Queensland (Stanley 
and Ross, 1986). As part of its efforts to 
control this weed the Queensland Depart- 
ment of Lands has supported a long ranging 
research program by the Alan Fletcher Re- 
search Station to find suitable biological 
control agents from the New World where 
the Baccharinae are native. 

Although the genus Baccharis is best rep- 
resented in South America with over 300 
species, some 21 species including B. ha- 
limifolia are native to North America. The 
Alan Fletcher Research Station set up field 

stations in Lake Placid, Florida (1968) and 
Curitiba, Brazil (1974) to survey the phy- 
tophagous insect fauna on Baccharis and to 
determine which species were sufficiently 
stenophagous (i.e. having a limited host 
range) for introduction to Australia. A num- 
ber of insects were subsequently introduced 
(McFadyen, 1981). In 1982 the North 
American Field Station was established in 
Temple, Texas to survey B. halimifolia at 
the western margin of its range and the 
closely related species B. neglecta Britton 
which is found in central and western Texas. 
Various surveys of insects on Baccharis 
sp. have been reported. Bennett (unpub- 
lished) surveyed the fauna on B. halimifolia 
in southeastern United States and on var- 
ious species of Baccharis in Brazil. Tilden 
(1951), after a very comprehensive survey, 
listed the insects associated with the vege- 
tative parts ofB. pilularis in the area to the 
south of San Francisco, California. Kraft 
and Denno (1982) listed the major herbi- 



vores attacking B. halimifolia in Maryland. 
All the studies, with the exception of that 
of Kraft and Denno (1982) indicated the 
Bacchahs is associated with a considerable 
number of insect species and that a number 
of species were most probably monopha- 

The Phenology and Range of 
b. halimifolia and b. neglecta 

Both B. halimifolia and B. ncglecta are 
perennial, dioecious woody shrubs growing 
to a height of about 1 5 feet. Both species 
produce massive amounts of seed which are 
dispersed by air and so they are often found 
colonizing disturbed or denuded areas. They 
are typically found along watercourses, in 
neglected pastures, along roadsides and 
drainage ditches, and in vacant lots in towns. 
The plants usually maintain their foliage 
during winter but hard freezes can cause 
defoliation and stem dieback. 

New growth begins in late winter. Kraft 
and Denno ( 1 982) reported that the leaf bio- 
mass of B. halimifolia increased steadily 
throughout spring and summer and then 
dropped slowly during autumn in response 
to an increase in inflorescence biomass. 
However, possibly more important, they re- 
ported that the leaves became significantly 
tougher and thicker during the growing sea- 
son while the moisture content and nitrogen 
content declined. The maturing leaves also 
increased in the concentration of an acetone 
soluble secondary chemical that acted as a 
deterrent to herbivory (Kraft and Denno, 
1982). These parameters suggested to these 
authors that there was a general decrease in 
the quality and availability of the foliage to 
herbivorous insects, and they noted that no 
major herbivore was found to feed on B. 
halimifolia in Maryland after early summer. 

Both species flower in autumn. The sta- 
minate inflorescences, which are a rich 
creamy color, are first to bloom followed a 
couple of weeks later by the white pistillate 
inflorescences. By late autumn the very small 
achenes, each attached to a feathery pappus. 

are dispersed by air. The reproductive out- 
put of a stand of B. halimifolia has been 
estimated as high as 376,000 achenes per 
m-, a figure exceeding that of any other plant 
species reported in the literature (Panetta, 

B. halimifolia and B. ncglecta are mor- 
phologically very similar. Mahler and Wa- 
terfall (1964) separate the species on the 
characters of leaf shape and involucre length: 
B. halimifolia has elliptic to rhomboid leaves 
and involucres 4-6 mm in length and B. 
ncglecta, narrowly elliptic, linear or oblan- 
ceolate leaves and involucres 4-8 mm in 
length. In areas where the two species over- 
lap it is often difficult to separate them be- 
cause of the tremendous variation in leaf 
shape and because intermediate types exist. 
These intermediate types are a good indi- 
cation that the two species are very closely 
related indeed. 

The habitat range of B. halimifolia ex- 
tends along coastal areas from Massachu- 
setts to Texas (Correll and Johnston, 1979). 
In Texas it is found east of a line that could 
be drawn between Victoria, Bryan and Dal- 
las i.e. in higher rainfall areas with acid soil 
types. It is abundant in coastal areas, low 
lying and poorly drained areas, and in dis- 
turbed habitats in townships and oil drilling 
areas. It is not found in forested areas be- 
cause the large trees soon completely dis- 
place it. B. ncglecta on the other hand, is 
found throughout almost all of Texas to the 
west and south of that line (Correll and 
Johnston, 1979); in areas with moderate to 
low rainfall and alkaline soil types. It also 
is found along roadsides, creeks, vacant lots 
in townships and other disturbed areas. In 
some areas it is known as "New Deal weed" 
or "Roosevelt weed" (Correll and Johnston, 
1 979) because it became weedy in the 1 930s 
when farmers, for financial reasons, were 
unable to properly tend to their pastures. 
Specimens of B. ncglecta have been col- 
lected from Arizona through to North Car- 
olina and also into Mexico (Correll and 
Johnston, 1979). 



Fig. 1 . The area surveyed in Texas, Louisiana and northern Mexico. Cities that were often visited are shown. 

The Area and Methods of Survey 

The area covered by this survey is given 
in Fig. 1 . It might be described as consisting 
of southern Louisiana, the Gulf Prairies and 
Marshes of Texas, the Blackland Prairies of 
Central Texas, the southern Edwards Pla- 
teau of Texas, the South Texas Plains and 
northern Mexico (cf Correll and Johnston 
(1979) for descriptions of the vegetational 
areas of Texas). 

The survey was conducted over a four 
year period between 1982 and 1986. In 1983 
and 1984 regular inspections at about two 
weekly intervals were conducted at a num- 
ber of sites in close proximity to Temple, 
Texas with particular emphasis being placed 
on a site on Lake Stillhouse Hollow about 
1 5 miles west of Temple. The areas farther 
afield from Temple were visited on an ir- 
regular basis by 3-4 day trips to such cities 

as Lafayette, Beaumont, Conroe, Galves- 
ton, Brownsville, Del Rio and Monterrey, 
Mexico. Stands oiBaccharis were inspected 
along the roadside on these trips particularly 
where the plants looked to be unhealthy be- 
cause of possible insect attack. In addition, 
certain sites were established near all of the 
above cities and these were inspected at 
every visit to that city. 

Insects were collected by both visually in- 
specting the plant and by sweeping the fo- 
liage. When evidence of internal insect in- 
festation was present, plants were either 
removed from the ground and dissected or 
the appropriate limb sawn off and split. Any 
evidence of feeding by the insect was noted. 
When immatures were found without adults 
being present, the immatures were collected 
and reared through to maturity to obtain 
adults for identification. This applied par- 



Table 1 . Phytophagous insect species collected on either B. halimifolia or B. neglecta in Texas, Louisiana 
or northern Mexico. 











Stages Collected 


Baccharis Hosts 





Hesperotettix viridis viridis 

Melanoplus dijferentialis (Tho- 

Melanoplus sp. 

Schistocerca alutacea alholineata 

Schistocerca obscura (F.) 


Hyalymenus tarsatus (F.) 

Acanthocephala declivis (Say) 

Acanthocephala terminalis (Dal- 

Acanthocephala thomasi (Uhler) 

Leptoglossus phyllopus (L.) 

Merocoris typhaeus (F.) 

Mozena lurida (Dallas) 

Cohmelaena pulicaria (Germar) 

Pangaeus bilineatus (Say) 

Largus cinctus (Herrich-Schaef- 

Lygaeus kalmii StSl 

Melanopleuris belfragei (Stai) 

Neocoryphus bicrucis (Say) 

Nysius niger Baker 

Nysius raphanus Howard 

Ochrimnus mimulus (St^l) 

Oncopeltus fasciatus (Dallas) 

Oncopehus sexmaculatus St&l 

Lygus lineolaris (Palisot de 

Polymems basalts (Reuter) 

Pseudatomoscelis seriatus (Ren- 

Talorilygus pallidulus (Blan- 










foliage feeder B. neg 

foliage feeder B. neg 

foliage feeder B. hal 

foliage feeder B. neg 

foliage feeder B. neg 

R nymph, adult ectophagous B. neg; B. hal 




B. neg 




B. neg 




B. neg 




B. neg; B. 




B. neg 




B. neg 




B. neg 




B. neg 


nymph, adult 


B. neg 




B. neg 




B. neg 




B. neg 



foliage feeder 

B. neg 



foliage feeder 

B. neg; B. 



nymph, adult 

flower and seed 

B. neg; B. 



flower and seed 

B. neg 




B. neg 




flower feeder B. neg, B. hal 

ectophagous B. neg 

flower feeder B. neg; B. hal 

flower feeder B. neg; B. hal 




Table 1. Continued. 











Stages Collected 


Baccharis Hosts 




Euschistus servus (Say) 

Mecidea major Sailer 

Nezara viridula (L.) 

Thyanta accerra McAtee 

Galgupha sp. 

Acanalonia bivittata (Say) 

Acanalonia conica (Say) 

Acanalonia laticosta Doering 

Acanalonia pan'a Doering 

Aphis pr. baccharicola HRL 

Aphis coreopsidis (Thomas) 

Clastoptera xanthocephala Ger- 

Lepyronia quadrangidaris (Say) 

Aceratagallia calcaris Oman 

Balclutha sp. 

Chlorotettix viridius Van Duzee 

Empoasca fabae (Harris) 

Homalodisca coagidata (Say) 

Menosoma cinctum (Osbom & 

Oecleus productus Metcalf 

Oliaris aridus Ball 

Stobaera pallida Osbom 

Rhynchotnitra recurva (Metcalf) 

Anormenis septentrionalis (Spino- 

Metcalfa pruinosa (Say) 

Ormenis saucia Van Duzee 

Ormenis sp. 

Ormenoides venustus (Melichar) 

Poblicia fuliginosa (Olivier) 

Hysteropterum auroreum (Uhler) 











B. neg; B. hal 
B. neg 
B. hal 
B. neg 

ectophagous B. neg 

R adult ectophagous B. neg 

O nymph, adult ectophagous B. hal 

R adult ectophagous B. neg 

R adult ectophagous B. hal 

C nymph, adult ectophagous B. neg; B. hal 

C nymph, adult ectophagous B. hal 

C nymph, adult ectophagous B. neg; B. hal 

O adult ectophagous B. neg 

R adult ectophagous B. neg 

R adult ectophagous B. neg 

R adult ectophagous B. neg 

O nymph, adult ectophagous B. neg; B. hal 

C adult ectophagous B. neg; B. hal 

R adult ectophagous B. neg 

R adult ectophagous B. neg 

R adult ectophagous B. neg 

C nymph, adult ectophagous B. neg; B. hal 

O nymph, adult ectophagous B. hal 

O adult ectophagous 

O adult ectophagous 

O adult ectophagous 

R adult ectophagous 

O adult ectophagous 

R adult ectophagous B. neg 

R adult ectophagous B. neg 

B. hal 

B. neg; B. hal 
B. neg 
B. hal 
B. hal 



Table 1. Continued. 





Stages Collected 



Speci- nomic 

Bacchans Hosts ficity- Pests' 

Tortistilus abnormus (Caldwell) 
Vanduzeea segmentata (Fowler) 



Aristotelia ivae Busck 

Anacamptodes defectaria (Gue- 

Anavitrinelia pampinaha (Gue- 

Eupithecia miserulata Grote 

Itame varadaria (Walker) 

Pero meskearia Packard 

Pleuroprucha insulsaria (Gue- 

Bucculatrix ivella Busck 

Platysenta videns (Guenee) 

Spodoptera frugiperda (Smith) 

Spodoptera ornithogalli (Guenee) 

Oidaematophorus balanotes 

Oidaematophorus kellicotti 

Homoeosoma electellum (Hulst) 

Epiblema discretivana (Heinrich) 

Platynota sp. 

Sonia paraplesiana Blanchard 


Amniscus perplexus (Haldeman) 
Ancylocera bicolor (Olivier) 
Anelaphus sp. 
Dendrobias mandibularis Ser- 

Dorcasta cinerea (Horn) 
Eliphidion linsleyi Knull 
Eliphidionoides incertus New- 
Euderces pini (Olivier) 
Slenosphenus dolosus Horn 
Tragidion coquus L. 

R adult ectophagous 

O nymph, adult ectophagous 





foliage feeder 

foliage feeder 

foliage feeder 

foliage feeder 
foliage feeder 
foliage feeder 
flower feeder 


larva, pupa 


foliage feeder 
foliage feeder 
foliage feeder 

stem borer 
stem borer 

flower feeder 

C larva stem gall 

R pupa 

C larva, pupa root feeder 

B. neg 
B. neg 

B. neg; B. hal 

B. neg 

B. neg 

B. neg 
B. hal 
B. neg 
B. neg; B. hal 


C larva, pupa leaf miner B. neg; B. hal 

B. neg; B. hal 

B. neg 

B. neg; B. hal 

B. neg; B. hal 
B. neg 

B. neg 

B. hal 
B. neg 
B. hal 


larva, pupa 

stem borer 

B. hal 



B. neg 



B. neg 



B. hal 



B. neg 



stem borer 

B. neg 



stem borer 

B. neg 



B. neg 



B. neg 



B. neg 



Table 1. Continued. 




Stages Collected 


Bacchans Hosts 

Speci- nomic 
ficity- Pests' 


Alt lea sp. 




B. hal 


Calomicrus prob. blakeae Wilcox 



B. neg 


Colaspis planicostata Blake 



B. hal 


Colaspoides opacicollis Horn 



B. neg 


Cryptocephalus cribripennis Le 



B. neg 



Cryptocephalus nr. pumilis 




B. neg; B 




Diabrotica balteata Le Conte 




B. hal 

* * 

Diabrotica connexa Le Conte 




B. hal 


Diabrotica undecimpunctata ho- 




B. neg; B 


* * 

wardi Barber 

Diachus an rat us (Fab.) 



flower feeder 

B. neg; B. 



Exema elliptica Karren 


larva, adult 

foliage feeder 

B. hal 


Microtheca ochroloma StSl 




B. hal 


Monoxia sp. 




B. hal 


Nodonota rotundicollis Schaeffer 



foliage feeder 

B. neg 


Nodonota texana Schaeffer 




B. neg 


Nodonota tristis (Olivier) 




B. neg 


Systena blanda Melsheimer 



foliage feeder 

B. neg 

* * 

Ophraella sexvittata Le Conte 



foliage feeder 

B. hal 


Paria thoracica (Melsheimer) 




B. hal 


Trirhabda bacharidis (Weber) 


larva, adult 

foliage feeder 

B. neg; B. 




Baris sp. 



B. neg 


Compsus auricephalus (Say) 



B. neg 


Cophes texanus Sleeper 




B. neg 


Eudiagogus pulcher Fahraeus 




B. hal 


Isodacrys burkei Howden 



B. neg; B. 



Lixus scrobicollis Boheman 




B. neg 


Mitostylus setosus (Sharp) 




B. neg 


Prosaldius blanditus (Casey) 



B. hal 


Prosaldius deplanatus (Casey) 



B. neg; B. 




Cryptorhopalum uteanum Casey 



B. neg 



Melanotes indistinctus Quate 



B. hal 



Lucidota sp. 



B. neg 


Pyropyga sp. 



B. hal 



Cotinus mutabilis Gory and Per- 




B. hal 



Euphoria sepulchralis (F.) 




B. hal 



Bothrotes canaliculatus acutus 



B. neg 


(Le Conte) 



Table 1. Continued. 











Stages Collected 


Bacchans Hosts 





Phytobia sp. 



leaf miner 

B. neg 



Neolasioptera lathami Gagne 



stem galler 

B. neg; B. hal 



Acinia picturata (Snow) 



seed feeder 

B. hal 


Dioxyna picciola (Bigot) 



B. hal 


Euarestoides acutangulus 



B. neg 



Neaspilota dolosa Benjamin 



B. neg 


Neotephritis finalis (Loew) 



B. neg 


Tephritis new sp. 



stem galler 

B. neg; B. hal 


Tephritis subpura (Johnson) 



stem galler 

B. hal 


Trupanea nr. actinobola (Loew) 



B. neg 


Tomoplagia obliqua (Say) 



B. hal 


' R = rare, O = occasional, C = common. 

: *** = monophagous (host plants apparently restricted to the genus Baccharis); ** = oligaphagous (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. 

' * = pest species. 

ticularly to caterpillars, leafminers, and gall 
formers. Samples of inflorescences were also 
collected in the autumn and placed in emer- 
gence cages and any resulting insects col- 

All insect specimens were first submitted 
to the Biosystematic and Beneficial Insects 
Institute, Agricultural Research Service, 
USDA, Beltsville, Maryland for expert 
identification by specialists of that Insti- 
tute's Systematic Entomology Laboratory. 
When species could not be fully identified 
by this laboratory the specimens were later 
forwarded elsewhere to other taxonomists 
expert with the particular group in question. 

After the insects had been properly iden- 
tified, entomologists knowledgeable about 
the particular species or group and the lit- 
erature were consulted to determine the de- 
gree of stenophagy exhibited by the species. 
Species that appeared to be sufficiently ste- 
nophagous were then selected for formal host 
specificity testing in order to obtain per- 

mission to introduce the insect into Aus- 
tralia (e.g. Palmer, 1986). 


The phytophagous species (excluding pol- 
len and nectar gatherers) found on either 
species of Baccharis are shown in Table 1 . 
One hundred and thirty three species were 
collected representing six orders. The Or- 
thoptera, Hemiptera, Homoptera, Lepidop- 
tera, Coleoptera and Diptera were repre- 
sented by 5 (or 4% of the species), 27 (20%), 
27 (20%), 17 (13%), 46 (35%) and 11 (8%) 
species respectively. 

The insects were classified as monopha- 
gous if restricted to Baccharis, oligophagous 
if the host range was restricted to the Tribe 
Astereae and polyphagous if having a wider 
host range. Evidence of host range was ob- 
tained from formal host testing, observa- 
tions during the course of the survey, con- 
sultations with acknowledged experts on 
specific groups of insects, examination of 



major insect collections, and texts such as 
Amett (1985), Slater and Baranowski ( 1 978), 
Amett et al. (1980) and Borrer et al. ( 1 98 1 ). 
Eleven insects were considered monopha- 
gous, and interestingly, all were found on B. 
halimifolia. Six of these eleven were also 
found on B. neglecta but only one monoph- 
agous species, Tephritis new sp., was more 
commonly found on B. neglecta than on B. 
halimifolia. The monophagous fauna thus 
represented about 8% of the total phytoph- 
agous fauna. 

Six of the 1 1 monophagous insects (or 
approximately 55%) were endophagous for 
at least part of their lifecycle. Often species 
that were definitely endophagous on Bac- 
charis, six were monophagous, one was oli- 
gophagous, two were polyphagous, and the 
hosts of one were not known although it also 
was quite probably monophagous. A very 
high proportion (80%) of the endophages 
therefore had a limited host range. 

Only two insects were classified as oli- 
gophagous. The remaining 1 1 8 species were 
considered either polyphagous, host un- 
known or hosts unknown but probably po- 
lyphagous. The proportion of oligophagous 
to polyphagous species depends, of course, 
on what arbitrary criteria are set for oligoph- 

Five of 1 1 monophagous species were also 
Lepidoptera. This proportion is consider- 
ably higher than the proportion of Lepi- 
doptera found in the total number of species. 

While six of 1 1 (or 55%) monophagous 
species were common to both B. halimifolia 
and B. neglecta, only 28 of the total 133 
species (2 1 %) were common to both species. 
This perhaps indicates that many of the po- 
lyphagous insects did not have any sub- 
stantial relationship with these hosts but 
rather their occurance (or absence) was de- 
pendent primarily on other factors. 

A number of well known crop pests were 
collected on Baccharis. These included the 
differential grasshopper, Melanoplus differ- 
entialis Thomas; the lygus bug Lygus lineo- 
laris (Palisot de Beauvois); the cotton flea- 

hopper, Pseiidatomoscelis seriatus (Reuter); 
the brown stinkbug, Euschistus servus (Say); 
the southern green stinkbug, Nezara viri- 
dula (L.); the southern com rootworm, Dia- 
brotica undecimpunctata howardi Barber 
and the fall armyworm, Spodoptera frugi- 
perda (Smith). 

During the course of the study many non- 
phytophagous insects were collected. These 
included known predator and flower feeding 
species as well as many insects that were 
probably only casually associated with the 
plant. These species are listed in Table 2. 
The collection of these insects was only a 
very secondary aspect of the project and 
quite likely there were many more such 
species present on Baccharis than are listed. 

Notes on the More Important Species 

By far the most important phytophage was 
the chrysomelid Trirhabda bacharidis (We- 
ber) which was found throughout the survey 
area except for the lower Rio Grande Valley 
and northern Mexico. It is univoltine in the 
study area with larvae occurring in late win- 
ter and adults being found from late April 
to early August. However in one year, 1 984, 
following an unusually wet autumn, early 
instar larvae were found in mid-December 
but these were killed during winter. It there- 
fore appears that at least some individuals 
in the population do not have a diapause 
mechanism and that the regularity of emer- 
gence at the end of winter may be more a 
function of extreme mortality of early 
emerging individuals rather than a dia- 
pause. Both larvae and adults can occur in 
tremendous numbers and are capable of 
completely defoliating a bush. The effect is 
particularly destructive if late winter freezes 
occur while the bushes are regenerating their 
foliage following larval attack. In this situ- 
ation the stems are frequently killed. 

The case bearing chrysomelid Exema el- 
liptica Karren was common throughout the 
B. halimifolia area in spring and summer. 
Larvae were found in April and May and 
adults thereafter. Both stages fed on foliage 



Table 2. Non-phytophagous insect species collect- 
ed on either B. halimifolia or B. neglecta in Texas, 
Louisiana or Northern Mexico. 

Table 2. Continued. 




Phalacrus sp. 





Onus insidiosus (Say) 


Plecia nearctica Harcy 




Deraeocohs nebulosus (Uhler) 


Villa sp. 




Euthyrhynchus floridanus (L.) 


Chrysomya rufifacies (Macquart) 


Podisus maculiventris (Say) 


Cochliomyia macellaria (Fab.) 




Phymata americana Melin 


Prodadius bell us (Loew) 


Phymata sp. 




Apallates particeps (Becker) 


Zelus bilobus (Say) 


Conioscinella grisescens (Sa- 




Galgupha sp. 


Conioscinella nuda (Adams) 



Liohippelates pusio (Loew) 


Thaumatomyia glabra (Mg.) 



Mewbruchis major (Fall) 

pollen feeder 

Ditrichophora argyrostoma 




Cantharis sp. 

pollen feeder 

Ochthera lauta Wheeler 


Cantharis sp. 

pollen feeder 

Philygria debilis Loew 


Chauliognathus basalts Le 



pollen feeder 

Chauliognathus marginatus (F.) 

pollen feeder 

Musca domestica (L.) 


Chauliognathus scutellaris Le 



pollen feeder 

Palaeosepsis pusio (Schiner) 


Chauliognathus sp. 

pollen feeder 


Podabrus sp. 

pollen feeder 

Allograpta obi i qua (Say) 

pollen feeder 


Palpada agrorum (Fab.) 

pollen feeder 

Colliuris pennsylvanica (Linne) 


Palpada pusilla (Macquart) 

pollen feeder 


Palpada vinetorum (Fab.) 

pollen feeder 

Coleomegilla maculata fuscila- 
bris (Mulsant) 


Syrphus rectus O. S. 
Toxomerus politus (Say) 

pollen feeder 
pollen feeder 

Cycloneda sanguinea (L.) 



Hippodamia convergens Guerin 


Angiorhina sp. 


Olla V nigrum Mulsant 


Pseudomyothria nr. ancilla 

Scymnus loewii Mulsant 





Ptilodexia sp. 


Epicauta pennsylvanica (De- 






Apis mellifera L. 

pollen feeder 

Collops balteatus LeConte 



Collops quadrimaculatus F. 


Dasymutilla sp. 



Polistes apachus Saussure 


Carpophilus nr. transitans Sharp 


Polistes sp. 




and heavily infested small plants may ex- 
hibit damage to their terminals. Although 
the type series for E. elliptica was reported 
from Ivafrutescens L. (Karren, 1966), it was 
not found on /. frutescens in this survey 
even though this plant was growing in close 
proximity to infested B. halimifolia on many 
occasions. It is therefore considered that a 
misidentification of the morphologically 
similar plant species may have occurred and 
that E. elliptica may be specific to Bac- 
charis. Two other genera of chrysomelids 
are quite commonly found on Baccharis. 
Adults of Nodonota spp. were found on B. 
neglecta in the spring. Infestations oiN. w- 
tundicollis Schaeffer were often seen along 
the Rio Grande Valley. Damage was in- 
variably noticeable but of little significance. 
Adults of the three polyphagous Diabrotica 
species were taken quite commonly in the 
autumn but never in damaging numbers. 

The lepidopterous foliage feeders caused 
at most only minor damage to the plant. 
Bucculatrix ivella Busck was the most abun- 
dant of these; and in April populations of 
several hundred per plant were sometimes 
seen, particularly on B. halimifolia. The first 
three instars feed inside a serpentine mine 
while the last two instars are external feed- 
ers. The very characteristic ribbed pupal co- 
coons were also found on the plant. During 
the rest of the year only very occasional 
specimens were seen. Greater detail on the 
biology and host specificity is given by 
Palmer and Diatloff (in press). The leaf web- 
bing caterpillar, Aristotelia ivae Busck, was 
also quite commonly found in spring but 
there were rarely more than one or two per 
plant. These small greenish larvae feed un- 
der a web on the leaf and become explo- 
sively active when touched. The geometrid, 
Itame varadaria (Walker) was collected from 
B. halimifolia at a number of sites by sweep- 
ing the foliage. It had three generations per 
year with larvae being present in April, July 
and October. It was never very abundant: 
a collection of half a dozen larvae after an 
hour's sweeping was a typical result. 

The most abundant stem borer was the 
plume moth Oidaematophorus balanotes 
(Meyrick), which was found throughout the 
survey area. The phenology of this univol- 
tine species was clearly defined. Moths were 
active in late summer and early autumn. 
Early instar larvae were often seen in inflo- 
rescences placed in emergence cages. They 
were also found in damaged vegetative ter- 
minals. Later instar larvae bored into the 
woody tissue of the stem and created a char- 
acteristic gallery which were up to a meter 
in length. The exit hole was covered with 
woody frass which had been removed from 
the gallery. Pupation and eclosion of the 
moth occured in the gallery. Occasionally 
bushes were heavily infested with this 
insect and on one occasion 1 5 larvae were 
found in the one stem. However, it was much 
more common to find plants infested with 
just one or two larvae. The related species, 
O. kellicotti (Fish) was found in B. neglecta 
in northern Mexico. This is a new host rec- 
ord for this species that has previously been 
reported only from Solidago spp. (Cashatt, 

The cerambycid, Amniscus perplexus 
(Haldeman), was found to infest a large pro- 
portion of J9. halimifolia plants at just a few 
sites. It was also univoltine with adult ac- 
tivity in late spring and early summer. Eggs 
were oviposited under the bark, usually near 
the crown of the plant and within 30 cm of 
ground level. Larval feeding continued from 
summer to the following spring when both 
pupae and teneral adults were found in the 
larval galleries. A characteristic finely pow- 
dered frass was found at the base of infested 
plants. Both large and small plants were at- 
tacked and it was quite common to find 2- 
3 larvae in quite small plants. The larvae 
significantly weakened the stems and pre- 
disposed the plants to attack by disease or- 

Two cerambycids Eliphidion linsleyi 
KnuU and Eliphidionoides incertus were 
found in B. neglecta stems along the Rio 
Grande Valley. Both species were associ- 



ated with weakened or dying branches, but 
it was not ascertained whether they had 
caused this damage or whether they had at- 
tacked already dying material. 

Four gall forming species were commonly 
found. The cecidomyiid Neolasioptera la- 
thami Gagne was found on both Baccharis 
species throughout the survey area. It forms 
a soft globular gall on the stems and ter- 
minals. A large gall (3 cm in diameter) might 
contain as many as 15 larvae, each in an 
individual chamber. Isolated patches of 
Baccharis were infested with up to 10 galls 
per plant in spring. For the rest of the year 
only very occasional galls were found. Sig- 
nificant damage to the plant was not ob- 
served within this survey area. A more de- 
tailed account of this insect is given by 
Diatloffand Palmer (1987). 

The two tephritids, Tephritis subpura 
(Johnson) and T. new sp., appeared to oc- 
cupy very similar ecological niches on B. 
halimifolia and B. neglecta respectively. 
Both fed in the terminal stems in spring and 
caused characteristic swelling of the final 
few centimeters of the stem and ultimately 
terminal die-off. Flies of both species 
emerged in autumn from inflorescences 
placed in emergence cages. The autumn 
adults of both species were smaller and 
darker than the individuals emerging in the 

The tortricid Epiblema discretivana 
(Teinrich) occurred in elongate woody stem 
galls, approximately 3 cm in length, on B. 
halimifolia. This insect was also univoltine, 
with adults emerging in early spring and the 
insect overwintering as larvae. E. discreti- 
vana is generally distributed throughout the 
habitat of B. halimifolia. As many as ten 
galls have been found on plants, but damage 
attributable to them was not discerned. 

The delphacid Stobaera pallida Osbom 
was found to be quite abundant on B. ha- 
limifolia and much less abundant on B. ne- 
glecta. There appears to be three generations 
a year with population peaks occurring in 
May, July and September. It was possible 

to collect over 1 00 individuals by sweeping 
one large bush. The life cycle is similar to 
that of other species of Stobaera (McClay, 
1983; Reimer and Goeden, 1982). Eggs are 
oviposited into the pith of stems and both 
nymphs and adults remain on the plant. 

With the onset of flowering the lygaeid 
Ochrimnus mimulus (St^l) adults were pres- 
ent in tremendous numbers on both male 
and female inflorescences. Later in autumn 
nymphs were found by dissecting the female 
inflorescences. The insect overwintered as 
late instar nymphs or adults, which were 
quite commonly found throughout the 
spring and summer. A more detailed ac- 
count of this insect is given by Palmer ( 1 986). 
The coreid Leptoglossus phyllopus (Say) was 
also very commonly associated with these 
plant species while they were flowering. 

Prospects for Biological Control 

Four of the insects, Trirhabda bacharidis, 
Aristotelia ivae, Oidaematophorus bala- 
notes and Neolasioptera lathami had pre- 
viously been found elsewhere in the United 
States and had been proved host specific by 
various officers of the Queensland Depart- 
ment of Lands. T. bacharidis was released 
in Queensland where it now occurs in dam- 
aging populations in some localized areas. 
A. ivae became generally distributed but has 
only been found at low, non-damaging pop- 
ulation levels. O. balanotes is at present 
being released in the field and A^. lathami 
has not yet been reared in the laboratory in 

A further five insects have been tested at 
the North American Field Station, Temple 
and permission to introduce these species 
into quarantine in Australia has been ap- 
proved or is anticipated. The species are 
Tephritis new sp., Stobaera pallida, Buccu- 
latrix ivella, Itame varadaria and Amniscus 
perplexus. The remaining monophagous in- 
sects will be tested in the near future. 

These 1 1 insects were rated in two ways 
in an attempt to predict their eventual ef- 
fectiveness as biocontrol agents in Austra- 



lia. Ideally it would be highly desirable if a 
reliable quantitative formula were available 
for use by biocontrol researchers. Harris 
(1973) devised a formula that was later 
modified by Goeden (1983), and this latter 
formula is possibly the best available at this 
stage. All the insects were therefore scored 
by Goeden's formula (Table 3). The scores 
ranged from 34 to 53. In this system N. 
lathami, O. balanotes and B. ivella, by scor- 
ing more than 50 points, would be consid- 
ered superior prospects and the rest were 
predicted to be partially effective agents. 

The insects were also assessed subjec- 
tively by the author based on observations 
in the United States only and rated from 1 
(poor prospect) to 5 (superior prospect) after 
considering such aspects as damage ob- 
served, ecoclimatic similarity to Australia, 
potential reproductive rate and degree of 
parasitism observed. T. bacharidis, B. ivella 
and A. perplexus were considered to be the 
best prospects. 

It should be pointed out, however, that a 
general and reliable method for predicting 
eventual effectiveness of potential biocon- 
trol agents has not yet been devised and its 
reliability proved. Aspects of Goeden's for- 
mula have been criticized by both Palmer 
(unpublished) and Wapshere (1985) who 
questioned whether it was indeed possible 
to quantify potential effectiveness in a new 
habitat. Perhaps the point to be made is 
that, while it is highly desirable to attempt 
to predict the best possibilities, preferably 
by quantitative methods, all sufficiently 
stenophagous agents should ultimately be 
utilized when at all possible. 


Faunal richness of species inhabiting a 
plant species is determined by many factors 
but Strong et al. (1984) considered the two 
most important factors to be the size of the 
geographic range and the plant "architec- 
ture" (i.e. its size and growth form). Both 
Baccharis species are rather large, woody, 
perennial shrubs that occupy an extensive 

Table 3. The potential effectiveness of the mono- 
phagous species as biocontrol agents as predicted by 
the formula of Goeden (1983) and by the author's sub- 
jective assessment (with a poor candidate scoring 1 and 
a superior prospect scoring 5). 


Goeden's Assess- 

Formula menl 

Amniscus perplexus Al 

Trirhabda bacharidis 45 

Exema elliptica 34 

Aristotelia ivae 49 

Oidaematophorus balanotes 53 

Bucculatrix ivella 5 1 

Itame varadaria 44 

Epiblema discretivana 36 

Stobaera pallida 4 1 

Tephritis subpura 40 

Tephritis palmeri n. sp. 35 

geographic habitat and these factors should 
indicate a rich insect fauna such as was found 
in the survey. Perhaps it could also be ar- 
gued that as the geographic area occupied 
by B. halimifolia is much greater than that 
of J5. neglect a a greater number of monoph- 
agous insects might be associated with B. 
halimifolia, as was found in this study. 

Another factor influencing the number of 
species found is, of course, the length of time 
devoted to the survey. New species were still 
being found in the last year of this project 
and undoubtedly the faunal list would have 
been longer had the survey been continued 
for a longer period. Nevertheless, four years 
represents a very adequate time frame for 
such a survey. 

The number of insect species common to 
both species of Baccharis clearly indicates 
that these plant species are very similar 
chemically as well as morphologically. In 
fact, the association is even closer than the 
data indicate. In the laboratory A. perple- 
xus, I. varadaria, and E. elliptica, found only 
on B. halimifolia in the field, fed readily on 
B. neglecta. Furthermore, a number of ste- 
nophagous insects collected from B. pilu- 
laris D.C. have fed equally well on both 
local species of Baccharis and also B. sar- 



athoides Gray, which is found in Arizona. 
It is therefore very probable that the differ- 
ences in insect fauna found between the two 
Baccharis species in this survey are due to 
different cHmatic factors or factors other 
than intrinsic differences between the species 

A very close association between steno- 
phagy and endophagy was evident. Endo- 
phages by their very nature are specialized 
with adaptions for internal living and must 
develop a close relationship with their host. 
It is therefore not surprising that a signifi- 
cant proportion of them are highly stenoph- 
agous. The high proportion of endophages 
that were also monophagous in this survey 
highlights the need for those involved in 
biological control programs such as this to 
place great importance on searching for en- 


I particularly thank the expert taxono- 
mists who identified the insects. Most tax- 
onomists were associated with the System- 
atic Entomology Laboratory (SEL), 
Agricultural Research Service, USDA or the 
associated Department of Entomology, Na- 
tional Museum of Natural History (NMNH), 
Smithsonian Institution. I also thank the 
Biosystematics and Beneficial Insects Insti- 
tute for providing such a fine service for 
collectors in North America through these 
two institutions. 

Identifications were performed by the fol- 
lowing taxonomists many of whom also of- 
fered valuable advice on the biology and 
habits of the insects they had identified: R. 
L. Brown, Mississippi Entomological Mu- 
seum (Lepidoptera: Tortricidae); E. V. 
Cashatt, Illinois State Museum (Lepidop- 
tera: Pterophoridae); J. A. Chemsak, UC 
Berkeley (Coleoptera: Cerambycidae); D. R. 
Davis, NMNH (Lepidoptera: Lyonetiidae); 
D. C. Ferguson, SEL (Lepidoptera: Geo- 
metridae, Pterophoridae, Pyralidae); R. H. 
Foote, SEL (Diptera: Tephritidae); R. J. 

Gagne, SEL (Diptera: Cecidomyiidae, Cal- 
liphoridae); R. D. Gordon, SEL (Coleop- 
tera: Scarabaeoidae, Meloidae, Lampyri- 
dae, Coccinellidae); T. J. Henry, SEL 
(Hemiptera); R. W. Hodges, SEL (Lepidop- 
tera: Gelechoidae); F. Hovore, Placerita 
Canyon Nature Center (Coleoptera: Cer- 
ambycidae); J. Jenkins, Washington State 
University (Diptera: Tephritidae); J. M. 
Kingsolver, SEL (Coleoptera: Dermestidae, 
Bruchidae); L. Knutson, SEL (Diptera: 
Bombyliidae); J. P. Kramer, SEL (Homop- 
tera: Fulgoridae, Cicadellidae, Membraci- 
dae, Cercopidae); D. R. Miller, SEL (Ho- 
moptera: Coccoidae); D. A. Nickle, SEL 
(Orthoptera: Acrididae); R. W. Poole, SEL 
(Lepidoptera: Noctuidae); C. W. Sabrosky, 
SEL (Diptera: Chloropidae); F. C. Thomp- 
son, SEL (Diptera: Syrphidae); T. J. Spil- 
man, SEL (Coleoptera: Cerambycidae, Ela- 
teridae); M. B. Stoetzel, SEL (Homoptera: 
Aphididae); W. W. Wirth, SEL (Diptera: 
Chironomidae); R. E. White, SEL (Coleop- 
tera: Chrysomelidae, Cerambycidae); and 
D. R. Whitehead, SEL (Coleoptera: Cur- 

Literature Cited 

Amett, R. H. 1985. American insects. A handbook 
of the insects of America north of Mexico. Van 
Norstrand Reinhold Co. New York. 850 pp. 

Amett, R. H., N. M. Downie, and H. E. Jacques. 1 980. 
How to know the beetles. W. C. Brown Co. Du- 
buque, lA. 417 pp. 

Bailey, F. 1900. The Queensland flora. Part 3. A. J. 
Diddams & Co. Brisbane, Qld. 

Borrer, D. J., D. M. De Long, and C. A. Triplehom. 
1981. An introduction to the study of insects. 5th 
Edition. Saunders College Publishing. Philadel- 
phia, PA. 827 pp. 

Cashatt, E. D. 1972. Notes on the balanotes (Meyr- 
ick) group of Oidaematophoms Wallengren with 
description of a new species (Pterophoridae). J. 
Lepid. Soc. 26: 1-13.11 

Correll, D. S. and M. C. Johnston. 1979. Manual of 
the vascular plants of Texas. University of Texas 
at Dallas, Dallas, TX. 1881 pp. 

Diatloff", G. and W. A. Palmer. 1987. The host spec- 
ificity of Neolasioptera lathami Gagne (Diptera: 
Cecidomyiidae) with notes on its biology. Proc. 
Entomol. Soc. Wash. 89: 185-199. 



Goeden, R. D. 1983. Critique and revision of Harris' 
scoring system for selection of insect agents in bi- 
ological control of weeds. Prot. Ecol. 5: 287-301. 

Harris, P. 1973. The selection of effective agents for 
the control of weeds. Can. Entomol. 105: 1495- 

Karren, J. B. 1966. A revision of the genus Exema 
of America, North of Mexico. Univ. Kansas Sci- 
ence Bull. 46: 672. 

Kraft, S. K. and R. F. Denno. 1982. Feeding re- 
sponses of adapted and non-adapted insects to the 
defensive properties of Bacchahs halimifolia L. 
(Compositae). Oecol. 52: 156-163. 

Mahler, W. F. and U. T. Waterfall. 1964. Bacchahs 
(Compositae) in Oklahoma, Texas and New Mex- 
ico. Southwest. Nat. 9: 189-202. 

McClay, A. S. 1983. Biology and host specificity of 
Stobaem concinna (StSl) (Homoptera: Delphaci- 
dae), A potential biocontrol agent for Parthenium 
hvsterophorusL. (Compositae). Fol. Entomol. Mex. 

McFadyen,P. J. 1981. Current status of the biological 
control programme against groundsel bush (Bac- 
chahs halimifolia). Proc. 6th. Aust. Weeds Conf 
Vol. 1. 151-154. 

Palmer, W. A. 1986. Host specihcily of Ochrim?ius 
mimulus (StSl) (Hemiptera: Lygaeidae) with notes 
on its phenology. Proc. Entomol. Soc. Wash. 88: 

Palmer, W. A. In press. Host specificity of Biiccu- 

lathx wella Busck (Lyonetiidae): a potential bio- 
control agent for Bacchahs halimifolia L. in Aus- 
tralia. J. Lepid. Soc. 

Panetta, F. D. 1979. The effects of vegetation devel- 
opment upon achene production in the woody 
weed, groundsel bush (Bacchahs halimifolia L.). 
Aust. J. Agric. Res. 30: 1053-1065. 

Reimer, N. J. and R. D. Goeden. 1982. Life history 
of the delphacid planthopper Stobaera tricahnata 
(Say) on western ragweed. Ambrosia psilostachya 
DC, in southern California (Hemiptera-Homop- 
tera: Delphacidae). Pan-Pac. Entomol. 58: 105- 

Slater, J. A. and R. M. Baranowski. 1978. How to 
know the true bugs. W. C. Brown Co. Iowa. 256 

Stanley, T. D. and E. M. Ross. 1986. Flora of South- 
Eastem Queensland Vol. 2. Queensland Depart- 
ment of Primary Industries. Brisbane. Misc. Pub. 
QM84007. 623 pp. 

Strong, D. R., J. H. Lawton, and R. Southwood. 1 984. 
Insects on plants. Community patterns and mech- 
anisms. Harvard University Press. Cambridge, 
MA. 313 pp. 

Tilden, J. W. 1951. The insect associates of 5acc/?ara 
p/7j//am De Candolle. Microentomology 16: 149- 

Wapshere, A. J. 1985. Effectiveness of biological con- 
trol agents for weeds: present quandaries. Agric. 
Ecosystems Environ. 13: 261-280. 


89(1), 1987, p. 200 


Hedge Bindweed, Calystegia sepium (Convolvulaceae), an Adventitious Host of the 
Chrysanthemum Lace Bug, Corythucha marmorata (Heteroptera: Tingidae) 

The chrysanthemum lace bug, Corythu- 
cha marmorata (Uhler), ranges throughout 
most of the United States and southern 
Canada (Slater and Baranowski, 1978. How 
to know the true bugs (Hemiptera-Heter- 
optera). Wm. C. Brown, Dubuque, Iowa, 
256 pp.). As the common name suggests, 
this tingid feeds on Compositae (= Aster- 
aceae), particularly species of Ambrosia, As- 
ter, Chrysanthemum, Helianthus, and Sol- 
idago (Bailey, 1951. Entomol. Am. 31:1- 
140). Drake and Ruhoff (1965. U.S. Nat. 
Mus. Bull. 243: 1-634) listed Echinops, 
Rudbeckia, and Tanacetum spp. as addi- 
tional composite hosts, and Horn et al. 
(1979. No. Car. Agric. Exp. Stn. Tech. Bull. 
257: 1-22) added Silphium to the known 
hosts. In Pennsylvania, C marmorata is 
common on spotted knapweed, Centaurea 
maculosa Lam., and mugwort, Artemisia 
vulgaris L.; in herb gardens on the Cornell 
University campus, Ithaca, N.Y., it causes 
extensive foliar chlorosis on wormwood, 
Artemisia absinthium L., and southern- 
wood, A. abrotanum L. (unpubl. observa- 
tions). The record of this tingid from oak, 
the only noncomposite host given in the 
literature, probably should be referred to the 
oak lace bug, C arcuata (Say) (see Bailey, 
ibid.: 88). 

On 7 August 1986,1 observed a large pop- 
ulation of the chrysanthemum lace bug 
damaging hedge bindweed, Calystegia se- 
pium (L.) R. Br. (formerly in Convolvulus) 
at Milan (Bradford Co.), Pennsylvania. All 
life stages of the tingid were present; eggs 
were inserted near the midrib and lateral 
veins on abaxial and adaxial surfaces. The 
hedge bindweed, growing on a fence above 

a mugwort plant infested with the lace bug, 
showed severe chlorosis and accumulation 
of cast skins and black excrement. 

Rather than a host shift, or the adding of 
a convolvulaceous plant to the diet of C 
marmorata, my observations probably rep- 
resent adaptive behavior allowing the bug 
to cope with the apparent deterioration of 
an isolated mugwort plant. The hawthorn 
lace bug, C cydoniae (Fitch), has been re- 
ported to temporarily colonize and injure 
roses growing near heavily damaged plants 
of a preferred rosaceous host, Cotoneaster 
sp. (Wheeler, 1981. Great Lakes Entomol. 
14: 37-43). In the present case the adven- 
titious host, hedge bindweed, belongs to a 
different family (Convolvulaceae) and order 
(Polemoniales) from the typical hosts of C 
marmorata; the Convolvulaceae and Com- 
positae, however, are members of the same 
subclass (Asteridae) (Cronquist, 1968. The 
evolution and classification of flowering 
plants. Houghton Mifilin, Boston, 396 pp.). 
For C. morrilli Osborn and Drake, another 
species of the genus that feeds on herba- 
ceous composites (Silverman and Goeden, 
1979. Pan-Pac. Entomol. 55: 305-308), the 
only known noncomposite host also belongs 
to the Convolvulaceae: sweet potato, Ipo- 
moea batatas (L.) Lam. (Drake and Ruhoff, 
ibid.: 155). 

Adults and nymphs have been deposited 
in the insect collection of the Pennsylvania 
Department of Agriculture (PDA). 

A. G. Wheeler, Jr., Bureau of Plant In- 
dustry, Pennsylvania Department of Agri- 
culture, Harrisburg, Pennsylvania 17110. 

89(1), 1987, pp. 201-203 

Book Review 

Coevolution of Parasitic Arthropods and 
Mammals, edited by Ke Chung Kim. Wi- 
ley-Interscience, a division of John Wiley 
& Sons, Inc., 605 Third Avenue, New 
York, NY 10158. xvi + 800 pp. 1985. 
$69.95/cloth. ISBN: 471-08546-4. 

This collection of essays marks the latest 
effort to summarize and analyze the increas- 
ingly daunting literature on host associa- 
tions of parasitic arthropods. As such, it 
continues a genre begun with the First Sym- 
posium on Host Specificity among Parasites 
of Vertebrates (Neuchatel, Switzerland, 
1957) and recently elaborated by Adrian 
Marshall in his masterly survey. The Ecol- 
ogy^ of Ectoparasitic Insects (1981). The style 
of Kim's book is immediately revealed in 
his list of contributors, among whom are 
several renowned arthropod systematists, 
each the doyen of his field: Emerson (Mal- 
lophaga), Fain (Astigmata), Hoogstraal 
(Ixodoidea), Radovsky (Mesostigmata), 
Traub (Siphonaptera). Himself an expert on 
Anoplura, Kim is the sole author of four 
chapters (1, 5, 7, 13) and co-author of two 
more (4, 10). 

The text is divided into four parts. Part 
One (chapters 1-3) is an introduction to the 
broad spectrum of evolutionary relation- 
ships between arthropods and mammals. 
Topics covered include parasite and host 
anatomy and morphology, reproductive 
cycles, population dynamics, dispersal, and 
geographic radiation— particularly as af- 
fected by continental drift. Part Two (chap- 
ters 4-8) examines the host associations, 
evolution, and zoogeography of ectopara- 
sitic Insecta, chiefly Phthiraptera and Si- 
phonaptera. Part Three (chapters 9-12) 
covers the Acari, with emphasis on pro-, 
meso-, and astigmatid mites as well as the 
Ixodoidea (ticks). Part Four (chapter 1 3) is 
an overview of the evolutionary pathways 
detailed in Parts Two and Three. Each chap- 

ter comes with its own list of references, 
which together run to 79 pages. Following 
the text are two remarkable appendices: A, 
an alphabetical list by family and genus of 
the world's parasitic arthropods and their 
mammal hosts; and B, the reverse of the 
preceding, an alphabetical list of the orders 
and families of mammals and their arthro- 
pod parasites. The work concludes with a 
56-page index to arthropods, mammals, and 
all subject headings. Most chapters contain 
numerous illustrations, the best being those 
of flea morphology (chapter 8) and the his- 
tory of continental movements (chapter 3). 
However, several figures have not repro- 
duced well, either because the originals were 
crudely executed (chapter 4) or excessively 
reduced (chapter 1 0). Students of a partic- 
ular group will also quickly note a number 
of minor typographical errors, as on page 
664 where the amblyommine tick genus 
Aponomma is misspelled twice (as Apo- 
nemma and Aponema) in the same sen- 
tence. Such blemishes seldom impede un- 

My criticism of this work stems from its 
title, which unduly stresses coevolution. As 
Dan Janzen makes clear in his short but 
engaging essay "Coevolution as a Process: 
What Parasites of Animals and Plants Do 
Not Have in Common," this is not a col- 
lection of coevolutionary studies in the strict 
reciprocal sense advocated by many who 
contributed to Futuyma and Slatkin's sem- 
inal synthesis Coevolution (1983). Rather, 
it is a review of parallelisms, especially those 
illustrative of Fahrenholz's Rule and re- 
source tracking. In fact. Fain and Hyland 
refrain from even mentioning coevolution 
in their chapter, "Evolution of Astigmatid 
Mites on Mammals"; they prefer "parallel 
evolution." My concern over definitions 
may seem trivial, but it arises from a basic 
difference between Kim's text and that of 



Futuyma and Slatkin. The latter gathered 
authorities from a wide range of fields with 
the object of formulating coevolutionary 
concepts. Kim, on the other hand, has chief- 
ly collaborated with systematic specialists— 
taxonomists— each of whom has focused on 
a particular ectoparasite group in order to 
describe (as he sees it) evolution in that 
group. One result is that much of this book 
is unnecessarily Darlingtonian in tone: 
hundreds of pages are given over to lists of 
taxa, their hosts and distribution, often with 
minimal evolutionary follow-up. 

Another problem in working with spe- 
cialists is "expert opinion": anything said is 
automatically e.x cathedra. A striking ex- 
ample is Hoogstraal's depiction of evolu- 
tion in the Ixodoidea (pp. 508-5 1 6). Bearing 
in mind that no pre-Eocene fossil ticks have 
ever been found (a few forms resembling 
extant species are known from amber), we 
read that ancestral ticks were eyeless para- 
sites of large, "glabrous" reptiles living 
communally during the late Paleozoic or 
early Mesozoic eras. The argasid line was 
represented by Argas and Ornithodoros 
"partially as we know them today," but oth- 
er argasids "probably did not evolve until 
the Tertiar> ." "Modern" ixodids (the Hae- 
maphysalinae, Ixodinae, and Rhipicepha- 
linae) evolved from spiderlike amblyom- 
mines and were "probably as large as the 
largest extant Amblyomma.'" while xeroph- 
ilous Hyalomma "may have appeared later, 
close to the Cretaceous period of Mesozoic 
environmental stresses." This entirely con- 
jectural scenario, capped by a regrettable 
dendrogram reminiscent of something by 
Ernst Haeckel, is based on the author's ex- 
pert knowledge of tick morphology and host 
associations, but it does not address con- 
flicting evidence from other fields. For ex- 
ample, Hoogstraal believes that prostriate 
ixodids (i.e. the genus Ixodes) are advanced 
because of their smaller size, shorter palps, 
streamlined morphology, and presumably 
recent radiation with the Rodentia. How- 
ever, he overlooks the rich chaetotaxy of 

Ixodes larvae, occasional mating off'the host, 
and the absence in this genus of a cement 
feeding cone, all characters suggestive of the 
supposedly primitive argasids. Available 
karyotypes for Ixodes also are similar to 
those in the Argasidae, as are the systems 
of sex determination (XX-XY) and sper- 
matogenesis (males in both groups may re- 
main aphagous). 

How might these authors have injected 
greater objectivity into their arguments? The 
obvious answer is through cladistic (or phy- 
logenetic) analysis, the only repeatable 
method of biotic classification that enables 
its users to generate testable hypotheses of 
phylogeny. This is not to say that references 
to cladistics are missing from Kim's book. 
The now familiar terms (apomorphy, ple- 
siomorphy, etc.) are there, and we are even 
treated to a small cladogram (p. 272) for the 
five genera of Echinophthiriidac (Ano- 
plura). For the most part, however, these 
are evolutionary' "just-so" stories, products 
of traditional synthetic taxonomic proce- 
dures that are neither repeatable nor test- 
able and depend entirely on the opinions of 
experts. It follows that all the zoogeographic 
arguments would also have greatly benefited 
from application of modern vicariance 
methodology— the union of cladistics and 
Leon Croizat's track analysis— which is 
blissfully free of any a priori assumptions 
concerning dispersal or centers of origin. 

Though thin on evolutionary analysis, this 
book is a veritable encyclopedia of ecto- 
parasitology, worthy of a prominent posi- 
tion in any entomologist's library. Kim's 
compilation is not likely to be superseded 
in our time, which should be comforting to 
those who must purchase it. But the flip side 
of literary immortality is the sad realization 
that contemporary society no longer seems 
interested in a sequel. As one painfully aware 
of this problem, Frank Radovsky deserves 
the last word (p. 496): "A pattern of . . . 
evolution has emerged, and I have attempt- 
ed to interpret it here. However, forms that 
are significant in understanding this pattern 



continue to be discovered. Our inventory 
of these is at best sketchy; the gaps in knowl- 
edge of bionomics and basic host-parasite 
relationships are especially glaring. Follow- 
ing a period of considerable interest in the 
1950s and early 1960s, there has been a 
decline in biological studies of vertebrate- 
associated [arthropods]. This important area 
of research should be revitalized." 

Richard G. Robbins, Department of 
Health and Human Services, Public Health 
Service, National Institutes of Health, Na- 
tional Institute of Allergy? and Infectious 
Diseases, % Department of Entomology, 
Museum Support Center, Smithsonian In- 
stitution, Washington, D.C. 20560. 

89(1), 1987, pp. 203-204 

Book Review 

The Pleasures of Entomology. Portraits 
of Insects and the People Who Study 
Them. By Howard E. Evans. Smithsonian 
Institution Press, Washington DC. 1985. 
238 pp., illus. Price: $14.95, paperbound 

The title says it all — studying insects is a 
pleasure. Not only studying them, reading 
about the studies of others is a pleasure. 
That's how Howard Evans, high expert on 
the Psammocharidae and other wasps at 
Colorado State University, feels about it. 
And he makes others feel the same, what 
with his low-key enthusiasm for the subject 
and his easy style of writing. He isn't a ran- 
ter, nor does he want us to act as if we were 
entering a cathedral of science. Put simply, 
he simply likes our animals. 

He devotes 12 chapters to 12 different 
insects, things such as gypsy moth, love bug, 
killer bee, boll weevil, and some less no- 
torious beasts. Each life history portrait is 
"painted" in a most engaging way, as is the 
story of the scientist who worked out that 
life history. There are chapters on garden 
insects, past entomologists, and himself. 

These are not dry accounts of insects; they 
are accounts of the positive or negative im- 
pacts that insects have made on the public 
and on the people who studied them. Evans 

knew or knows many scientists who worked 
out life histories or complicated control 
measures. Even when he discusses those he 
didn't know, usually long before his time, 
he does it with appreciation and freshness. 
He stresses the personal involvement of the 
entomologist with the insect, putting a hu- 
man touch in the science of entomology; he 
loves the lover (amateur), sometimes even 
more than the professional, and rues his 
passing. Because many entomologists came 
accidentally to their calling and suffered and 
enjoyed much along the way, the evolution 
of each is almost as interesting as the life 
history of the insect. 

You don't have to be a good bug to be 
interesting to Howard Evans, for he some- 
times welcomes pests into his home garden 
for the sheer pleasure of watching them. He 
watches for what he has read and looks for 
something new. A few extra plants, so the 
crop yield is still adequate, is a small price 
to pay for a pest's presence. He even wishes 
he could attract a few more that sound in- 
teresting, like the harlequin cabbage bug or 
imbricated snout beetle. And as for control, 
he admits to the pleasure of popping and 
oozing tobacco homworms between his toes, 
just as he did as a boy. 

Evans often implies or states emphati- 
cally that our society is too materialistic, 



that we are too much concerned with the 
GNP (gross national product) when we 
should be concerned with the PGN (pro- 
founder grasp of nature). Surely this book 
and others like it will help toward a better 
balance of our priorities. 

I have recently become interested in the 
history of entomology and the Americans 
who made it, so I was happy to read his 
chapters devoted to Abbot, Say, Peck, Har- 
ris, Cockerell, Williams, and Dietrich. As 
in all history, there is no set formula for 
success. Different personalities, such as 
Cockerell and Williams, with very different 
methods of attack can make tremendous 
contributions. Evans tells us that the only 
requirements for success in our field are great 
amounts of inquisitiveness, patience, and 
love of insects. 

Not only are insects interesting to ento- 
mologists, they provide a source of liveli- 
hood. Therefore, for all the information that 
Drosophila has supplied and for the careers 

that it has made possible, Evans awarded 
those fruitflies the Evans Prize: a bunch of 
bananas. I'll turn that line back on him, so 
to speak; for giving pleasure with this book, 
I award Howard E. Evans the Spilman Prize: 
a sharp nib, a full jar of ink, and an ento- 
mological author's rating of 9.0 (in a scale 
of 1-10). (He's in good company; I once 
awarded Shakespeare a 9.9 and the author 
oftheBiblea 10). 

It is my duty as reviewer to find an error. 
On page 164, the Colorado Potato Beetle 
reached Illinois by 1864, not 1964. Thus 
have I justified receiving this book free. Even 
so, I feel guilty; now that I have read it, I 
almost want to pay for it. It's that good. 

T. J. Spilman, Systematic Entomology 
Laboratory, BBII, Agricultural Research 
Service, USD A, % National Museum of 
Natural History, NHB-168, Washington 
B.C. 20560. 



Miscellaneous Publications 

Cynipid Galls of the Eastern United States, by Lewis H. Weld $ 5.00 

Cynipid Galls of the Southwest, by Lewis H. Weld 3.00 

Both papers on cynipid galls 6.00 

Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman 1.00 

Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. 

Traver 1.00 

A Short History of the Entomological Society of Washington, by Ashley B. Gumey 1.00 

Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. 

Steyskal 1.50 

Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. 

Steyskal 2.00 

Memoirs of the Entomological Society of Washington 

No. 1. The North American Bees of the Genus Osmia. by Grace Sandhouse. 167 pp. 1939 $15.00 

No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. (out of 
95 pp. 1942 print) 

No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. 

253 pp. 1949 15.00 

No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 15.00 

No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 

1957 15.00 

No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi 

Takahasi. 230 pp. 1969 15.00 

No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. 

1976 11.00 

No. 8. The North American Pifedaceous Midges of the Genus Palpomyia Meigen (Diptera: Cera- 

topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 12.00 

No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 

pp. 1981 10.00 

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 1 1.00 

No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. 

1983 18.00 

No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 

67 pp. 1984 „ 5.00 

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. 


(Continued from front cover) 

PETERS, T. M.— A new Nearctic Dixa (Diptera: Dixidae) from Pennsylvania 137 

SCHAUFF, M. E.— Taxonomy and identification of the egg parasites (Hymenoptera: Platygas- 
tridae, Trichogrammatidae, Mymaridae, and Eulophidae) of citrus weevils (Coleoptera: 
Curculionidae) 31 

SCHWARTZ, M. D. andG. M. STONED AHL—Oaxacacom, a new plant bug genus and three 

new species of Orthotybni from Mexico (Heteroptera: Miridae) 15 

SHARKEY, M. J.—Agathis thompsoni n. sp., a Nearctic species of Agathidinae (Hymenoptera: 

Braconidae) parasitic on Greya subalba (Braun) (Lepidoptera: Incurvariidae) 47 

SMITH, D. R. and E. M. BARROWS— Sawflies (Hymenoptera: Symphyta) in urban environ- 
ments in the Washington, D.C. area 147 

SPANGLER, P. J. and R. C. FROESCHNER- Distributional data, illustrations, and habitat of 

the South American water-strider Microvelia ayacuchana (Hemiptera: Veliidae) 167 

STARK, B. P. and B. C. KONDRATIEFP— A new species of Peltoperla from eastern North 

American (Plecoptera: Peltoperlidae) 141 

TURNER, C. E., R. W. PEMBERTON, and S. S. ROSENTHAL- Host range and new host 
records for the plume moth Platyptilia carduidactyla (Lepidoptera: Pterophoridae) from 
California thistles (Asteraceae) 132 

WALTZ, R. D. and W. P. McCAFFERTY-New genera of Baetidae (Ephemeroptera) from 

Africa 95 

WALTZ, R. D. and W. P. McCAFFERTY— Generic revision of Cloeodes and description of 

two new genera (Ephemeroptera: Baetidae) 177 

WHARTON, R. A.— Changes in nomenclature and classification of some opiine Braconidae 

(Hymenoptera) 61 

WOODLEY, N. E.— The Afrotropical pachygastrine genera Ashantina Kertesz and Meristo- 

meringina James, with two new generic synonyms (Diptera: Stratiomyidae) 103 

WOOLLEY, J. B. and H. W. BROWNING— Morphometric analysis of uniparental Aphytis 
reared from chaff scale, Parlatoria pergandii Comstock, on Texas citrus (Hymenoptera: 
Aphelinidae; Homoptera: Diaspididae) 77 


HALSTEAD, J. A.—Brachymeria discretoidea, a new junior synonym oi Brachymeria discreta 

(Hymenoptera: Chalcididae) 131 

WHEELER, A. G., JR.— Hedge bindweed, Calystegia sepium (Convolvulaceae), an adventitious 

host of the chrysanthemum lace bug, Corythucha marmorata (Heteroptera: Tingidae) . . . 200 


ROBBINS, R. G. — Coevolution of Parasitic Arthropods and Mammals 201 

SPILMAN, T. J. — The Pleasures of Entomology. Portraits of Insects and the People Who Study 

Them 203 


\ ' VOL. 89 

APRIL 1987 

NO. 2 

(ISSN 0013-8797) 


of the 





ADAMSKI, D. and R. L. BROWN— A new Nearctic Glyphidocera with descriptions of all stages 

(Lepidoptera: Blastobasidae: Symmocinae) 329 

BURROWS, W. L.— A new species of Ameletus (Ephemeroptera: Siphlonuridae) from eastern 

North America 284 

DAVIS, D. R.— Neotropical Tineidae, IV: Three new Acrolophus species from Cuba and the 

rediscovery of Acrolophus niveipunctatus Walsingham (Lepidoptera) 275 

DONNELLY, T. W.— Structural variation of Ophiogomphus mainensis: description of a new 

subspecies and relationship to sibling species (Odonata: Gomphidae) 205 

FENNAH, R. G.— A new subfamily of Nogodinidae (Homoptera: Fulgoroidea) with the de- 
scription of a new species of Gastrinia 363 

GOEDEN, R. D. — Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in south- 
em California 269 

GRISSELL, E. E. and L. DE SANTIS— A new species of Erixestus (Hymenoptera: Pteromalidae), 

an egg parasitoid of Calligrapha polyspila (Coleoptera: Chrysomelidae) in Argentina .... 264 

HOEBEKE, E. R., Q. D. WHEELER, and R. L. GILBERTSON- Second Eucinetidae-Conio- 
phoraceae association (Coleoptera: Basidiomycetes), with notes on the biology of Eucinetus 
oviformis LeConte (Eucinetidae) and on two species of Endomychidae 215 

HURYN, A. D.— A new species of Notiphila (Notiphila) (Diptera: Ephyridae) from Ohio 322 

KROMBEIN, K. V.— Synonymic notes on the Bethylidae described by V. de Motschulsky 

(Hymenoptera: Aculeata) 356 

MASON, W. R. M.— Discovery of female Apozyx (Hymenoptera: Apozygidae) and comments 

on its taxonomic position 226 

MASON, W. R. M.— Vadum. a new genus of Nearctic Braconidae (Hymenoptera) 325 

MILLER, J. S.— A revision of the genus Phryganidia Packard, with description of a new species 

(Lepidoptera: Dioptidae) 303 

{Continued on back cover) 




Organized March 12, 1884 

Thomas E. Wallenmaier, President 
F. Eugene Wood, President-Elect 
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Second Class Postage Paid at Washington, D.C. and additional mailing office. 


89(2), 1987, pp. 205-214 




Thomas W. Donnelly 

2091 Partridge Lane, Binghamton, New York 13903. 

Abstract.— Ophiogomphus mainensis mainensis Packard, new status, consists of a large, 
stable population (New England and adjacent parts of New Jersey, Pennsylvania, New 
York, Quebec, and New Brunswick) and a disjunct population in the high mountains of 
North and South Carolina. An allopatric population in central Pennsylvania and West 
Virginia is named as a new subspecies fastigiatus. Structurally intermediate specimens 
from the vicinity of the boundary of the two ranges have been found in Pennsylvania and 
New York. Ophiogomphus incurvatus Carle may have been derived from O. m. fastigiatus, 
with the subspecies O. i. alleghaniensis Carle the bridging taxon. Ophiogomphus acumi- 
natus Carle occurs southwest of these two species; its relation to the other two species is 
uncertain. The geographic pattern of variation of O. mainensis sspp. is unusual for odo- 
nates of the eastern United States, and is reminiscent of "leapfrog" variation of Andean 

The gomphid Ophiogomphus mainensis 
was described by Packard (1863) from a fe- 
male collected in Maine. A male was de- 
termined as mainensis by Hagen (Selys, 
1878; Howe, 1918); however, Howe's (1918) 
figure shows that it is not this species, and 
Garman (1927) identified it as carolus 
Needham. Needham (1897) reared a male 
of mainensis but described it as O. johan- 
nus. The two species johannus and mainen- 
sis were considered to be distinct until Gar- 
man (1927) estabUshed the synonymy. 

The name Ophiogomphus incurvatus was 
proposed by Carle (1982) to replace the in- 
correctly named O. carolinus Hagen, a 
species of the piedmont of Georgia to Mary- 
land. Carle also described the subspecies O. 
i. alleghaniensis, which occurs to the west 
of the nominate subspecies. 

Ophiogomphus acuminatus was de- 
scribed by Carle (1981) from specimens from 
Tennessee. O. bouchardi, which was de- 

scribed by Louton (1982), is a synonym. 
Although placed by Carle with O. edmundo 
Needham, it seems to be closer to O. mai- 
nensis s. lat. and incurvatus s. lat. 

The present study began with the recog- 
nition of a distinct taxon occurring in cen- 
tral and western Pennsylvania south to high 
elevations in West Virginia (Fig. 1). Al- 
though this taxon has some resemblance to 
O. i. alleghaniensis, it is closely related to 
mainensis. Preliminary studies suggested 
that it might merit specific status, but a few 
specimens of this taxon and typical mai- 
nensis close to their geographic boundary in 
northeastern Pennsylvania and southern 
New York show intermediate characters. 
Thus, the new taxon is considered a sub- 
species of mainensis. 

Abbreviations for collectors' names in the 
descriptions are: TD = T. Donnelly; CS = 
C. Shiffer; HW = H. B. White III; FC = F. 
Carle; SD = S. Dunkle; CU = Cornell Univ.; 



Fig. I. Map showing distribution of Ophiogomphus m. mainensis (U.S. specimens examined for this study 
only; "m" for Canadian localities reported in Walker, 1958); O. m. fastigiatus\ O. i. incurvatus (localities from 
Carle, 1982 and from my collection); O. i. alleghaniensis (localities from Carle, 1982, with additional localities 
from specimens in J. Louton Collection); and O. acuminatus. Arrows show localities with structurally inter- 
mediate specimens. 



OF = O. S. Flint, Jr., U.S. Nat. Mus.; SB = 
S. W. Bromley, U.S. Nat. Mus.; ED = E. 
M. Davis, Mus. Comp. Zool.; EW = E. B. 
Williamson, Univ. Mich. Mus. Zool. 

Ophiogomphus mainensis fastigiatus 

Donnelly, New Subspecies 

Figs. 2-5 

Ophiogomphus mainensis, Ahrens et al., 
1968: 107. Butler Co. specimen included 

Ophiogomphus mainensis, Beatty et al., 
1969: 131. In part this subspecies. 

Holotype male.— Head: pale yellow green, 
very dark brown as follows: thin marginal 
line on labrum, vertex except for a pale sub- 
basal medial spot. 

Thorax: proepistemum very dark brown 
with pale lateral spots, proepimeron yellow 
brown. Pterothorax yellow green, with a very 
dark brown mid-dorsal line, which does not 
meet the anterior margin. Mesepistemum 
with a dark curved line extending to % the 
distance to rear and not touching meso- 
pleural suture. Mesopleural and metapleu- 
ral sutures and margins of mesepimeron and 
metepistemum dark brown. Legs black ex- 
cept for obscure pale of coxae and interior 
surface of pro femora. 

Abdomen: Each segment mainly black, 
green as follows: sides of 1 and 2; basal- 
lateral and medial-lateral spots on 3 to 7, 
decreasing in size rearwards; fused lateral 
spots on 8 and 9; medial-basal lines on 1 to 
8, thin and short on 3 to 8; tiny middorsal 
spot on 9; 10 with postero-lateral line and 
small posterior middorsal spot. Appendages 
brown, superiors indistinguishable from m. 
mainensis, in lateral view each appendage 
straight, tapering, divergent in dorsal view 
(Fig. 5, no. 11), sub-parallel sided in lateral 
view with prominent, small posteriorly di- 
rected apical point. Inferior appendage in 
lateral view with lateral spine low and less 
prominent than apical spine, located about 
Vi the distance from base (Fig. 2, no. 9); in 

ventral view (Fig. 3, no. 9) the appendage 
is distinctly wedge-shaped, in sharp contrast 
to typical m. mainensis, in which the ap- 
pendage is obtusely truncated (Fig. 3, nos. 
1 to 4). 

Allotype female.— Coloration as in male, 
with pale colors slightly more extensive. The 
occiput with paired, anteriorly directed 
spines as in m. mainensis. The vulvar lam- 
ina resembles that of m. mainensis; in lat- 
eral view the tips are elevated and then de- 
flected caudally, forming a sigmoid bend 
(Fig. 5, no. 10). A reliable distinction be- 
tween females of m. fastigiatus and m. mai- 
nensis has not been found. 

Types. -Holotype 3, PENNSYLVANIA: 
Sullivan Co., Loyalsock Cr. 3 mi. N. of 
LaPorte, 4 July 1983 (TD). Allotype 9, same 
loc, 19 June 1983 (TD). Paratypes: PENN- 
SYLVANIA: same loc. as holotype: 1 9 June 
and 4 July, 5 <3 2 9 1983 (TD and CS); Clin- 
ton Co., Kettle Cr., 2 mi. E. of Hammersley 
Fork, 21 June 1969, 21 June 1975, 22 June 
1977, 25 June 1983, 5 3 4 9 (CS); Centre 
Co., Black Moshannon Creek nr. Black 
Moshannon Dam, 4 Aug. 1983, 1 6 (CS); 
Somerset Co., Rockwood, 29 June 1900, 1 
6 (EW); WEST VIRGINIA: Nicholas Co., 
Cranberry R. 5 mi. N. of Richwood (2100'), 
25 June 1969, 1 6 (TD); Randolph Co., 
Shaver Fork of Cheat R. at highway 250 
(3600'), 23 June 1973, 1 6 (HW). The ho- 
lotype and allotype are deposited in the 
Florida State Collection of Arthropods. 

Dimensions. —The holotype male has ab- 
domen and appendages 34 mm long, and 
hind wing 26 mm long. The allotype female 
has these dimensions 32 and 27 mm. The 
total group examined (14 males, 7 females) 
has these dimensions 32.7 mm (.85) and 
25.65 (.5) mm for the males and 31.8 (.4) 
and 27.1 (.7) for the females. The figure in 
parentheses is standard deviation. 

Variations in the type series.— The color 
patterns show Httle variation. In about half 
the specimens the posterior end of the mes- 
epistemal dark stripe is free, and in half it 



Fig. 2. Profiles of appendages of male Ophiogomphus m. mainensis, 1-5, and O. m. fastigiatus, 6-12. 
Specimens 5 {mainensis) and 6-8 {fastigiatus) show intermediate characteristics. Localities are 1 = Essex Co., 
Vt.; 2 = Perry Co., Pa.; 3 = Pike Co., Pa.; 4 = Haywood Co., N.C.; 5 = Sullivan Co., N.Y.; 6 and 7 = Sullivan 
Co., Pa.; 8 = Centre Co., Pa.; 9 = Sullivan Co., Pa. (holotype); 10 = Somerset Co., Pa.; 1 1 = Nicholas Co., W. 
Va.; 12 = Randolph Co., W. Va. 

is fused with the humeral stripe. The three 
pale spots on abdominal segment 10 are 
fused in many specimens, forming a pos- 
terior band. The appendages show some 
variation, as seen in Figs. 2, 3, and 4. In 
three specimens (two from Sullivan Co., 
Pennsylvania, and one from Centre Co., 
Pennsylvania; Figs. 2 and 3, nos. 6 to 8) the 
lateral spine on the inferior appendage is set 
posterior to the midpoint, somewhat as in 
m. mainensis. Although these specimens are 
more like m. fastigiatus than m. mainensis 
in the ventral view of the appendages (Fig. 

3, nos. 6 to 8) and in the relatively small 
medial spine of the inferior appendage, their 
intermediate character is significant. 

Remarks.— The subspecies m. fastigiatus 
is distinguishable from the nominate sub- 
species by the form of the inferior append- 
age, which has a low, blunt lateral spine set 
at about the midpoint of the appendage (Fig. 
2, nos. 9 to 12). In m. mainensis this ap- 
pendage is variable, but always has a very 
prominent, pointed lateral spine located 
distinctly caudal to the mid point of the 
inferior appendage (Fig. 2, nos. 1 to 4). In 



9 10 11 

Fig. 3. Ventral views of male appendages of same specimens shown in Fig. 2. 

ventral view (Fig. 3, nos. 9 to 12) the wedge 
of the inferior appendage is distinctive (the 
name fastigiatus refers to a gable). 

Ophiogomphus mainensis mainensis 

Packard, New Status 

Figs. 2-4 

Ophiogomphus mainensis, Packard, 1863. 

Holotype 9. 
Ophiogomphus johannus, Needham, 1897. 

Holotype 6. 
Ophiogomphus johannus, Howe, 1918. 
Ophiogomphus mainensis, Garman, 1927. 

Established synonymy of mainensis and 

johannus. Descr. $ and 9, figs. 

Ophiogomphus mainensis, Walker, 1958. 
Descr. 3 and 9, figs. 

Ophiogomphus mainensis. Carle 1982. De- 
scriptive notes, figs. 

The holotype is a female, which, as has 
been the case with many other gomphids, 
has resulted in continuing confusion. The 
male of the nominate subspecies has the 
thorax marked as follows: yellow green, with 
a dark brown mid-dorsal stripe ending pos- 
terior to anterior margin; dark brown stripes 
on mesopleural (humeral) and metapleural 
sutures, the latter covering the anterior half 
of suture; dark curved stripe on mesepi- 
stemum adjacent to mesopleural suture and 



Fig. 4. Inclined views of male appendages of O. m. mainensis, 1-2, and O. m. fastigiatus, 3-4. Specimen 3 
shows intermediate characteristics. Localities are 1 = Perry Co., Pa.; 2 = Pike Co., Pa.; 3 = SuHivan Co., Pa.; 
4 = Sullivan Co., Pa. (holotype). 

extending % distance towards wing bases. 
The distinctive inferior appendage is shown 
in Figs. 2, 3, and 4. 

Needham's holotype of O. johannus was 
examined and found to be m. mainensis. 
However, the holotype is a reared male pre- 
served in alcohol, and the specimen has sub- 
sequently turned to a jellied mass, of which 
the sclerotized abdominal appendages are 
among the few portions recognizable as parts 
of an insect. 

Material examined. — MAINE: Aroos- 
took Co., 18 mi. W. Ashland, 19 July 1982, 
1 $ (TD); Washington Co., E. Machias R. 3 
mi. N. of Wesley, 11 July 1982, 2 <5 (TD); 
Penobscot Co., Seboeis R. west of Shin 
Pond, 28 July 1959, 2 5 1 9 (TD). MAS- 
SACHUSETTS: Franklin Co., West Haw- 
ley, 23 June 1971, 1 <5 (HW); Middlesex Co., 

Ashby, 9 July 1939, 1 $ (ED). NEW YORK: 
Hamilton Co., nr. L. Durant, 3 July 1980, 
13 19 (HW); Herkimer Co., Wilmurt, n.d., 
1 $ (holotype of O. johannus) (CU); Dela- 
ware Co., Beaver Kill, nr. Roscoe, 31 July 
1967, 1 6 (TD); SulHvan Co., Beaver Kill at 
Lewbeach, 31 July 1967, 2 6 (somewhat 
intermediate to fastigiatus) (TD). VER- 
MONT: Essex Co., nr. Island Pond, 14-15 
July 1982, 5 <5 3 9 (SD); Brunswick, 7 July 
1985, 1 3 1 9 (HW); Outlet Dennis Pond, 
24 June 1983, 2 $ (FC); Caledonia Co., Pea- 
cham, 27 June 1983, 2 $ (FC). NEW 
HAMPSHIRE: Hillsborough Co., Wilton, 
22 June 1969, 1 <3 (HW). CONNECTICUT: 
Fairfield Co., Stamford, 16 June 1940, 1 <5 
(SB). NEW JERSEY: Morris Co., Hackle- 
barney St. Pk., 21 June 1959, 1 $ (TD). 
PENNSYLVANIA: Pike Co., and Wayne 



Fig. 5. Structural details of Ophiogomphus incurvatus alleghaniensis, 1-4, 9, O. mainensis fastigiatus, 10- 
12, and O. acuminatus, 5-8. 1 and 5 are lateral, 2 and 6 ventral, and 3, 7, and 11 are ventral views of male 
appendages. 9 and 10 are inclined views of female vulvar laminae. 4, 8, and 12 are the vesicle of the penis. 

Co., nr. Greentown, 4-5 July 1985, 2 <3 (TD); 
Perry Co., nr. New Bloomfield, 23 June 
1965, 3-10 July 1966, 3 June 1975, 4 3 1 2 
(CS). NORTH CAROLINA: Haywood Co., 
nr. Sunburst, 15 June 1984, 2 <5 (TD and 
CS); Yancey Co., nr. Busick, 1 1 June 1958, 
6 S (TD); Avery Co., nr. Minneapolis, 13 
June 1958, 2 <5 (TD). SOUTH CAROLINA: 
Oconee Co., Burrell's Ford, Chattooga R., 
19 May 1970, 1 <3 (OF). 

Variations in Ophiogomphus m. mainen- 
sis.— Specimens from three disjunct popu- 
lations were examined: a northern, extend- 
ing from Maine to New Jersey and 
northeastern Pennsylvania (26 males, 6 fe- 
males), a small group from Perry Co., Penn- 
sylvania (4 males, 1 female), and a group 
from western North Carolina and South 
Carolina (1 1 males). Sizes of the three pop- 
ulations are virtually the same, and I de- 

tected no distinction in color pattern nor 
structure between the northern and south- 
em populations. The Perry Co. specimens 
were slightly larger but identical in color- 
ation. The mean dimensions for males of 
the three groups of m. mainensis are as fol- 
lows: northern group abdomens 32.4 (.6) 
mm and hind wings 25.7 (.6) mm; for the 
southern group 33.3 (.9) and 25.9 (.45) mm; 
for Perry Co., Pennsylvania 34.25 (.3) and 
26.25 (.3) mm. 

Remarks.— The two subspecies are indis- 
tinguishable in color pattern, and both show 
parallel variations in the mesepistemal dark 
stripe and pale color of segment 10. The 
sizes are very similar, except for the slightly 
larger Perry Co. population of m. mainen- 

The inferior appendages show some vari- 
ation in the size of the lateral spine, but in 



all specimens this spine is prominent, point- 
ed, and located posterior to the mid point 
of the appendage. In ventral view the ap- 
pendage varies from a nearly oblique ter- 
mination to a more acute wedge shape. Two 
specimens from Sullivan Co., New York 
(Figs. 2, 3, no. 5) are extreme in this regard 
and appear to vary in the direction of m. 
fastigiatus, while still retaining a predomi- 
nantly m. mainensis appendage. 

In summary, the two subspecies m. mai- 
nensis and m. fastigiatus are allopatric with 
a tendency for structurally intermediate 
specimens to occur near the boundary be- 
tween their ranges. The disjunct southern 
population is separated from West Virginia 
m. fastigiatus by a gap occupied by Ophio- 
gomphus incurvatus alleghaniensis. 

Distinction Among Taxa of the 
Ophiogomphus mainensis Group 

The Ophiogomphus mainensis group 
contains three species {mainensis, incurva- 
tus, and acuminatus), two of which have 
subspecies. Males of the three species are 
most readily identified by the form of the 
superior appendage (cerci), as shown in 
Fig. 5. 

Ophiogomphus mainensis and incurvatus 
(Fig. 5, nos. 1-4) are distinguished by 
the pale femora, and rounded, blunt-tipped 
superior appendages of incurvatus. The 
inferior appendage viewed ventrally is wed- 
ge-shaped as in m. fastigiatus but has a 
prominent lateral spine as in m. mainensis. 
The abdomen is paler in incurvatus than in 
mainensis. The vesicle of the penis is black 
and broader in mainensis and narrower and 
centrally pale in incurvatus. The two sub- 
species of incurvatus have been character- 
ized by Carle (1982). The similarity in the 
inferior appendages of /. alleghaniensis and 
m. fastigiatus is noteworthy and might have 
a bearing on their relationship. Ophiogom- 
phus acuminatus is a lesser known species 
found in Tennessee. The male is readily dis- 
tinguished by its broad superior appendage 
with a distinctively pointed tip (Fig. 5, nos. 

5-7). The vesicle of the penis has narrower 
paired cylindrical processes than either in- 
curvatus or mainensis. 

Females of mainensis and incurvatus are 
best distinguished by the paler colors of the 
latter, as discussed by Carle ( 1 982). The vul- 
var lamina of i. alleghaniensis (Fig. 5, no. 
10) is similar to that of m. fastigiatus (Fig. 
5, no. 9) but appears on the basis of one 
specimen to be distinguishable by its tip 
which is deflected ventrally rather than sig- 
moidally ventrally and caudally. However, 
this distinction may be a poor one; this 
sclerite is subject to both in vivo and post 
mortem changes in shape. The female of 
acuminatus has not been described. 

Ophiogomphus m. mainensis and m. fas- 
tigiatus are allopatric and are distinguished 
from each other only by the form of the male 
inferior appendages (epiproct). Although this 
appendage shows some variability through- 
out both populations, all specimens exam- 
ined are referrable to one or the other sub- 
species by size of the lateral spine of the 
inferior appendage and by shape of this ap- 
pendage in ventral view, as discussed above 
and shown in Figs. 2, 3 and 4. The existence 
in the boundary area between the subspecies 
of some specimens with appendages of in- 
termediate shape shows that genetic isola- 
tion has not been achieved and reinforces 
the conclusion that the two taxa are of sub- 
specific rank. 

The Systematic Relationships and 
Putative Derivation of the Taxa in 


Ophiogomphus mainensis occurs from 
Quebec and New Brunswick southward in 
the high Appalachian Mountains to South 
Carolina, a distribution shared with many 
other Odonata. However, the subspecies 
mainensis occurs as two disjunct popula- 
tions: the northern is widespread in New 
England, extending southward to Harris- 
burg at high elevations. The southern oc- 
cupies the high mountains of western North 
and South Carolina. The subspecies fasti- 



giatus occupies much of the intervening area, 
occurring from Pennsylvania south to the 
New River valley, which cuts through the 
Appalachians in southern West Virginia and 
southwestern Virginia. I suggest that m. 
mainensis is the ancestral form and origi- 
nally had a continuous distribution 
throughout the entire range. This type of 
distribution is well known among Andean 
birds and has been called "leapfrog varia- 
tion" by Remsen (1984), who suggested that 
the very young tectonics of the Andes might 
provide opportunities for restriction in gene 
flows, with the consequent appearance of 
derivative forms in the center of a once- 
continuous range. 

The tentative explanation offered here be- 
gins with late Cenozoic uplift of the Ap- 
palachian mountains which was concen- 
trated in the Carolinas along the Cape Fear 
arch. The uplift and accompanying west- 
ward tilting caused major tributaries of the 
Mississippi River system (Tennessee, 
French Broad, Holston, and New rivers) to 
penetrate the mountains and capture former 
eastward drainages. Continued develop- 
ment of these river systems left the Appa- 
lachian chain divided into segments. The 
New River created the broadest low-ele- 
vation division of the mountain chain. Sub- 
sequent to Pleistocene glaciation, odonate 
species in a putative southern Appalachian 
refugium spread northward along the 
mountain chain. Continued amelioration of 
the climate drove many species into higher 
areas, and the gaps created by cross-cutting 
river systems left some of these populations 
genetically isolated. The originally contin- 
uous population ofOphiogomphus mainen- 
sis was broadly divided by the New River 
gap. One possible result was the appearance 
of a relatively low-elevation form (O. mai- 
nensis fastigiatus) in this gap. Further cli- 
matic amelioration drove the boundary be- 
tween mainensis and fastigiatus further 
north to its present position, where the two 
subspecies face each other across a relatively 
short boundary region. 

At a still later time, a new form, O. in- 
curvatus, supplanted /fl^r/'^/a/wj at lower el- 
evations in this gap. It subsequently divided 
into two forms, /. incurvatus and /. alle- 
ghaniensis, across the drainage divide be- 
tween the New and Roanoke River systems 
in southwestern Virgina. At present /. in- 
curvatus is confined to Atlantic drainages, 
and /. alleghaniensis occupies Gulf and Mis- 
sissippi drainages. The origin of acuminatus 
remains elusive, but a derivation from in- 
curvatus is plausible. The similarity be- 
tween the inferior appendages of m. fasti- 
giatus and /'. alleghaniensis is consistent with 
these taxa being annectant forms between 
the two species but is not compelling. Other 
schemes might be proposed; a final reso- 
lution of this question would best be ap- 
proached through an assessment of protein 


I am most grateful for the loan or gift of 
specimens by O. S. Flint, Jr., J. Louton, H. 
B. White III, S. Dunkle, F. Carie, C. Shiffer, 
and L. K. Gloyd. I am further indebted to 
Frank Carle for extended discussions on 
speciation in Ophiogomphus and related 
genera and to J. Louton, S. Dunkle, R. Gar- 
rison, F. Carle, and C. Shiffer for stimulating 
criticism of this paper. 

Literature Cited 

Ahrens, C, G. H. Beatty, and A. F. Beatty. 1968. A 
survey of the Odonata of western Pennsylvania. 
Proc. Pa. Acad. Sci. 42: 103-109. 

Beatty, G. H., A. F. Beatty, and C. Shiffer. 1969. A 
survey of the Odonata of central Pennsylvania. 
Proc. Pa. Acad. Sci. 43: 127-136. 

Carle, F. C. 1981. A new species of Ophiogomphus 
from eastern North America, with a key to the 
regional species (Anisoptera: Gomphidae). Odon- 
atologica 10: 271-278. 

Carle, F. L. 1982. Ophiogomphus incurvatus: A new 
name for Ophiogomphus carolinus Hagen (Odo- 
nata: Gomphidae). Ann. Entomol. Soc. Am. 75: 

Garman, P. 1927. The Odonata or Dragonflies of 
Connecticut. Conn. State Geol. Nat. Hist. Surv. 
Bull. 39. 331 pp. 

Howe, R. H. Jr. 1918. Manual of the Odonata of 



New England, part III. Mem. Thoreau Mus. Nat. 
Hist: II, pp. 25-40. 

Louton, J. A. 1982. A new species of Ophiogomphus 
(Insecta: Odonata: Gomphidae) from the western 
highland rim of Tennessee. Proc. Biol. Soc. Wash. 
95: 198-202. 

Needham, J. G. 1 897. Preliminary studies on North 
American Gomphinae. Can. Entomol. 29: 181- 

Packard, A. S. 1863. In Walsh, B. D., ed.. Obser- 
vations on Certain N.A. Neuroptera. Proc. Ento- 
mol Soc. Phila. 2: 167-272. 

Remsen, J. V. Jr. 1984. High incidence of "leapfrog" 

pattern of geographic variation in Andean birds: 

Implications for the speciation process. Science 

224: 171-173. 
Selys Longchamps, E. de. 1878. Quatriemes addition 

au synopsis de gomphines. Bull. Acad. R. Belg. 

(2)46: 408-698. 
Walker, E. M. 1958. The Odonata of Canada and 

Alaska, volume 2. Univ. of Toronto Press. 318 



89(2), 1987, pp. 215-218 





E. Richard Hoebeke, Quentin D. Wheeler, and Robert L. Gilbertson 

(ERH, QDW) Department of Entomology, Cornell University, Ithaca, New York 14853; 
(RLG) Department of Plant Pathology, University of Arizona, Tucson, Arizona 85721. 

Abstract. —A breeding population (adults and larvae) of the eucinetid beetle, Eucinetus 
oviformis LeConte, was found in association with basidiocarps of the wood-rotting fungus 
Coniophora arida (Fr.) Karst. var. arida (Basidiomycetes, Coniophoraceae) in the Finger 
Lakes region of New York in late summer 1986. Larvae of £". oviformis, maintained in 
the laboratory, aggregated upon the substrate and pupated in one closely-packed group. 
Pupae hung inverted from the last larval exuvia which were attached to the substrate by 
the caudal end. In addition, adults and larvae of the endomychids, Mycetina perpulchra 
(Newman) and Aphorista vittata (F.), also were found in association with the same wood- 
rotting fungus. The Coniophoraceae-^". oviformis association and the two breeding species 
of Endomychidae on a host in the Coniophoraceae provide additional evidence for the 
diversity of Coleoptera feeding on minute Basidiomycetes. 

Wheeler and Hoebeke (1984) recently re- 
ported the discovery of adults, larvae and 
pupae of Eucinetus oviformis LeConte in 
association with basidiocarps of the wood- 
rotting fungus Coniophora olivacea (Pers.) 
Karst. near Highlands, North Carolina (Ma- 
con County). This represented only the sec- 
ond breeding record for a eucinetid beetle 
on a Basidiomycete, although fungus-feed- 
ing habits have been suggested for more than 
a century (Perris, 1851). The first substan- 
tiated record was for E. punctulatus Le- 
Conte, which breeds in boletes (Bruns, 
1984), although fungus associations were 
also reported by Klausnitzer (1971, 1975). 

In this paper we report the discovery of 
a breeding population of E. oviformis near 
Trumansburg, New York (Tompkins Coun- 
ty) also on a host in the Coniophoraceae, 
and report the association of the endomy- 

chids Mycetina perpulchra (Newman) and 
Aphorista vittata (F.) with the same fungus. 


Until our previous report on E. oviformis 
(Wheeler and Hoebeke, 1984), no Eucine- 
tidae were known to feed on wood-rotting 
Basidiomycetes (Gilbertson, 1984). The 
collection of another breeding population 
(including adults and larvae) of this euci- 
netid beetle on a related species of the same 
genus of fungus, Coniophora, provides fur- 
ther confirmation of this association. 

On August 26, 1986, one of us (ERH) 
found a large fallen trunk of hemlock (Tsuga 
canadensis (L.) Carr.) with a very extensive 
development of basidiocarps of Coniophora 
arida (Fr.) Karst. var. arida. Found on the 
surface of the fungal fruiting bodies were 



adults and larvae of Eucinetus oviformis. 
Mature larvae and some adults were still 
present as late as October 18, 1986. The 
collection site was in a large virgin woodlot 
(known as the Henry A. Smith Woods, 
founded in 1909), located on the southern 
village limit of Trumansburg, New York. 
This woodlot is a small (ca. 30 acres) but 
impressive stand of old growth forest. The 
principal soil type is an Arkport fine sandy 
loam (deep, well-drained and acidic 
throughout the profile). The forest is dom- 
inated by beech (Fagus grand (folia Ehr.), 
sugar maple {Acer saccharum Marsh.), tulip 
tree {Liriodendron tulipifera L.), and hem- 
lock (Tsuga canadensis (L.) Carr.). The 
woodland floor is shadowed by an extensive 
canopy and moisture content is relatively 
high. These conditions support an under- 
story dominated by elder (Sambucus), 
spikenard (Aralia racemosa L.) and numer- 
ous herbaceous plants. In addition to the 
various life stages of the eucinetid, the ba- 
sidiocarps of C arida were supporting large 
breeding populations of two species of En- 
domychidae that are discussed below. 

Until its association with Coniophora oli- 
vacea, the hosts of Eucinetus oviformis were 
unknown. Although other members of the 
Eucinetidae feed on either fungi or slime 
molds, there is no clear evidence to suggest 
that any eucinetids have particularly broad 
feeding habits. Our two records of Conioph- 
ora hosts may suggest that E. oviformis has 
a restricted host range— within the Conio- 
phoraceae or related taxa. This hypothesis, 
of course, remains to be tested with future 
field work. 

Both species of Coniophora reported as 
hosts for eucinetid beetles have a wide range 
in North America. Coniophora arida var. 
arida is circumglobal in the North and South 
Temperate zones, and C. olivacea is circum- 
global in the North Temperate Zone (Ginns, 
1982). They occur primarily on dead co- 
nifers but may also decay dead hardwoods, 
particularly those in coniferous forest eco- 
systems. Their basidiocarps are annual, and 

in most parts of North America would de- 
velop primarily in the summer and early 
fall and would deteriorate rapidly with the 
advent of low temperatures. They overwin- 
ter as mycelia in the wood. Coniophora 
species cause a brown rot, selectively re- 
moving the cellulose and hemicelluloses 
from wood and leaving a stable residue of 
slightly modified lignin. 

Orientation of Eucinetid pupae 

We maintained live Eucinetus oviformis 
larvae in the laboratory, taken from the Tru- 
mansburg population. These larvae suc- 
cessfully pupated, and we noticed an inter- 
esting phenomenon that has not been 
previously reported in the Eucinetidae. Lar- 
vae pupated in the rearing container gre- 
gariously, and pupae hung inverted by an 
attachment to the last larval exuvium (Fig. 
1 ). The same attachment was apparent, upon 
reexamination, in the material collected in 
North Carolina. We are uncertain how 
widespread this pupation behavior is in oth- 
er Coleoptera, but have noted a similar case 
(involving gregarious behavior and inverted 
hanging in the larval exuvium) in a species 
of Neotropical Nilio (Nilionidae or Tenebri- 
onidae) (Wheeler, unpublished data). 

Endomychidae Associated with 

Relatively few specific fungal associa- 
tions are recorded for species of Endomy- 
chidae; those that are recorded include 
various Basidiomycetes (Scheerpeltz and 
Hofler, 1948; Benick, 1952). The lack of 
much specific host data suggested to Crow- 
son (1984) that many endomychids might 
be associated with Ascomycetes. Our ob- 
servations on two genera (below), however, 
add another possible explanation for the lack 
of published host records. At the Trumans- 
burg collecting site and intermingled with 
the eucinetids, breeding populations of two 
species of Endomychidae were also found 
on the same host, Coniophora arida. On 
August 26, 1986, numerous larvae of My- 



Figs. 1,2. 1, Aggregation of pupae o{ Eucinetus oviformis LeConte; pupae hang inverted from the last larval 
exuvia which are attached by the caudal ends to the substrate. Scale line = 4 mm. 2, Mature larvae ofAphorista 
vittata (F.) (4 large, dark larvae) and Mycetina perpulchra (Newman) (2 small, pale larvae) grazing on basidiocarps 
of the wood-rotting fungus Coniophora arida var. arida. Scale line = 1 cm. 



cetina perpulchra (Newman) and Aphorista 
vittata (F.) were found moving over basi- 
diocarps of the wood-rotting fungus (Fig. 2). 
Adult pairs of both species were found in 
copula upon the felled hemlock trunk. Lar- 
val identification was made by association 
with adults. Considerably more larvae and 
adults of A/, perpulchra than A. vittata were 
present on this first collecting date. On later 
visits to the same site (September 27 and 
October 18, 1986), ERH found larvae o{ A. 
vittata in much greater abundance. Also on 
October 18, adults of M perpulchra were 
again extremely numerous. So far as we 
know, this is the first published record of 
Endomychidae on a host in the Conio- 
phoraceae. Voucher specimens of larvae and 
adults of both endomychids are in the Cor- 
nell University Insect Collection along with 
samples of the wood-rotting fungus. 


Many wood-rotting Basidiomycetes nev- 
er produce large fruiting bodies (Gilbertson, 
1984) and yet occur with such abundance 
and frequency that they must pose a sizable 
resource for mycophagous insects. This 
confirmation of the Coniophoraceae-Euci- 
netidae association and the new breeding 
reports of two species of Endomychidae on 
a host in the Coniophoraceae provide added 
evidence for the diversity of Coleoptera 
feeding on minute Basidiomycetes, a fauna 
that also includes Dasyceridae (Wheeler, 
1984). Similar field studies are needed to 
further elucidate relationships between bee- 
tles and these wood-rotting fungi. 


This research was supported, in part, by 
NSF Grant No. BSR-83 15457 and Hatch 
Project no. NY(C) 139426 to Q. D. Whee- 

ler. We thank Peter Marks (Cornell Univer- 
sity) for providing us with information about 
the abiotic and biotic communities of the 
Henry Smith Woods. 

Literature Cited 

Benick, L. 1952. Pilzkafer und Kaferpilze. Acta Zool. 
Fennica, no. 70, 250 pp. 

Bruns, T. D. 1984. Insect mycophagy in the Boletales: 
Fungivore diversity and the mushroom habitat, 
pp. 9 1-1 29. /« Wheeler, Q. and M. Blackwell, eds., 
Fungus-Insect Relationships. Columbia Univer- 
sity Press, New York. 

Crowson, R. A. 1 984. The associations of Coleoptera 
with Ascomycetes, pp. 256-285. In Wheeler, Q. 
and M. Blackwell, eds. Fungus-Insect Relation- 
ships. Columbia University Press, New York. 

Gilbertson, R. L. 1984. Relationships between in- 
sects and wood-rotting Basidiomycetes, pp. 130- 
165. In Wheeler, Q. and M. Blackwell, eds., Fun- 
gus-Insect Relationships. Columbia University 
Press, New York. 

Ginns, J. 1982. A monograph of the genus Co/j/op/zo- 
ra (Aphyllophorales, Basidiomycetes). Opera Bot. 
61: 1-61. 

Klausnitzer, B. 1971. Zur Biologic einheimischer Ka- 
ferfamilien, 7. Eucinetidae. Entomol. Ber. 15: 73- 

. 1975. Beitrage zur Insektenfauna der DDR: 

Coleoptera-Eucinetidae. Beitr. Entomol., Bed. 
25(2): 325-327. 

Perris, E. 1851. Quelques mots deles metamorphoses 
de Coleopteres mycetophages, le Triphyllus punc- 
tatus. Fab.; le Diphyllus lunatus. Fab.; YAgathi- 
dium seminulum, Linn., et VEucinetus (Nycteus 
Latr.) meridionalis de Castelnau. Ann. Soc. Ento- 
mol. France (2)9: 39-53. 

Scheerpeltz, O. and K. Hofler. 1 948. Kafer und Pilze. 
Verlag fur Jugend und Volk, Wien. 351 pp. + IX 

Wheeler, Q. D. 1 984. Notes on host associations and 
habitats of Dasyceridae (Coleoptera) in the south- 
em Appalachian Mountains. Coleopt. Bull. 38: 

Wheeler, Q. D. and E. R. Hoebeke. 1984. A review 
of mycophagy in the Eucinetoidea (Coleoptera), 
with notes on an association of the eucinetid bee- 
tle, Eucinetus oviformis, with a Coniophoraceae 
fungus (Basidiomycetes: Aphyllophorales). Proc. 
Entomol. Soc. Wash. 86: 274-211. 


89(2), 1987, pp. 219-225 




Paul J. Spangler 

Department of Entomology, National Museum of Natural History, Washington, D.C. 

Abstract.— A new species of psephenid beetle, Pheneps cursitatus Spangler, from Cerro 
de la Neblina, Venezuela, is described, illustrated with line drawings and scanning electron 
micrographs, and compared to its closest relative, Pheneps antennalis Spangler & Steiner 
(1983), from Surinam. A photograph and notes on the habitat are provided. 

Insect collections made during a biolog- 
ical survey of the plants and animals of the 
tepui Cerro de la Neblina, in southern Ven- 
ezuela, yielded numerous new taxa, distri- 
bution records, and biological data. The new 
aquatic beetle described in this paper is 
another example of the rich and relatively 
unstudied fauna of the Amazon basin. 

The genus Pheneps and two species, P. 
gracilis from Haiti and P. cubanus from 
Cuba, were described by Darlington (1936); 
a third species, P. antennalis from Surinam, 
was described more recently by Spangler and 
Steiner (1983). Although the male of P. cu- 
banus is unknown, the males of P. gracilis, 
P. antennalis, and P. cursitatus, new species, 
are known and all have exceptionally long 
antennae (Figs. 1, 2) as described for the 
genus by Darlington. 

Pheneps cursitatus Spangler, 
New Species 
Figs. 1-12, 15 

Holotype male.— Form and size: Body 
flattened; thorax narrowed anteriorly; elytra 
diverging slightly posteriorly, widest at about 
posterior three-fourths, apices rounded. 
Length, 2.47 mm; width, 1.31 mm. 

Coloration: Covered with fine, dense, 

short, recurved, golden pubescence. Head, 
antennal segments 3-11, pronotum, and 
elytra dark brown; basal two antennal seg- 
ments, labium, and labial palpi yellowish 
brown. Ventral surface light yellowish brown 
except maxillary palpi, prostemum, sides of 
metastemum, apices of femora, and tibia 
gray brown. Abdomen dark gray brown ex- 
cept slightly yellowish medially. Tarsi yel- 
lowish brown. 

Head: Almost flat behind eyes, decurved 
between eyes; finely microreticulate and 
finely punctate, more densely so anteriorly; 
labroclypeal suture distinctly depressed. 
Clypeus with anterior margin subtruncate. 
Labrum narrow, broadly subrectangular, 
and indistinctly arcuately emarginate on an- 
terior margin. Eyes prominent, hemispher- 
ical. Antenna (Fig. 5) densely pubescent; fi- 
liform; exceptionally long, as long as length 
of body. Maxillary palpus (Fig. 2) 4 seg- 
mented, filiform; about three-fourths the 
length of antennal segments 1-3 combined; 
first segment shortest, about a fifth as long 
as second segment; second segment longest, 
only slightly longer than fourth segment; 
third segment about three-fourths as long 
as second segment; fourth segment slightly 
longer than third segment, slightly com- 



Figs. 1-4. Pheneps ciirsitatus, new species, male. 1, Habitus, dorsal view. 2, Habitus, ventral view. 3, 
Maxillary palpus, apical segment. 4, Protarsus. 

pressed laterally (Fig. 3). Labial palpus very 
small, 3 segmented; first segment about a 
third as long as second segment; second seg- 
ment swollen and slightly longer than third 
segment; third segment partially com- 
pressed apically. 

Thorax: Pronotum widest at base; discal 

area densely, finely punctate; discal punc- 
tures separated by Vi to 1 times their di- 
ameter; disc moderately convex, moderate- 
ly depressed adjacent to posterolateral 
angles; finely and indistinctly margined 
ventrolaterally; anterolateral angles strong- 
ly rounded; posterolateral angles forming 



right angles but slightly rounded; with apical 
and basal margins moderately bisinuate. 
Scutellum slightly wider than long, flat; fine- 
ly and densely punctate; rounded poste- 
riorly. Elytra about 3.5 times longer than 
pronotum and about a fourth wider at wid- 
est point than pronotum; humeri slightly 
gibbous; apices broadly rounded; each ely- 
tron with 5 indistinct striae on discal area 
paralleling elytral suture; punctures fine and 
very dense. Prostemum (Fig. 2) moderately 
short in front of procoxae; covered by plas- 
tron setae. Prostemal process narrow, keel- 
like; apex extending into mesostemum. 
Mesostemum narrow between mesocoxae, 
about an eighth as wide as mesocoxal width 
(Fig. 8); with a very narrow, median, lon- 
gitudinal cleft. Metastemum behind me- 
socoxae abruptly and strongly raised above 
plane of mesostemum; with a very fine, me- 
dian, longitudinal line extending along 
length of raised portion; surface microretic- 
ulate and finely, densely punctate. Femora 
robust and swollen. All tibiae very slender 
and each with a low but distinct carina pos- 
terolaterally (Figs. 2, 3, arrows). Mesotibiae 
and metatibiae each with a bifurcate carina. 
Protibiae and mesotibiae grooved laterally 
on apical three-fifths. Protarsal segments 
(Fig. 4) and mesotarsal segments 1 and 2 
much broader than segments 3, 4, and 5 and 
densely pubescent ventrally; metatarsi un- 
modified. Tarsal claws small, slender, and 
sharp; each with a subbasal tooth. 

Abdomen: All sterna with surface sculp- 
ture as on metastemum and covered with 
plastron setae. Second visible sternum 
strongly emarginate in medial third. Fifth 
visible sternum broadly emarginate along 
posterior margin. Sixth visible stemum 
broadly triangularly incised medially, re- 
sulting in 2 subtriangular lateral lobes. Sev- 
enth visible stemum rounded at apex. 

Male genitalia: Bulbous basally and tri- 
lobate apically as illustrated (Figs. 11, 12). 

Female.— The female differs from the 
male as follows. Larger size, 3.90 mm long, 
1.51 mm wide. Antennae (Fig. 6) only 

slightly longer than length of pronotum and 
segments submoniliform. Maxillary palpi 
short; about as long as combined length of 
basal 2 antennal segments and about half as 
long as palpi of male. Mesostemum wide 
between mesocoxae, about half as wide as 
mesocoxal width (Fig. 7). Visible abdomi- 
nal stema 1-5 subequal in length (Fig. 9); 
only fifth stemum shallowly emarginate ap- 
icomedially; sixth stemum broadly round- 
ed. Tarsal segments all equally very slender. 

Variation.— The specimens in the type se- 
ries are very similar except for the slightly 
variable extent of the yellowish color of the 
abdominal stema and variations in length 
from 2.47 to 2.90 mm and width from 1.20 
to 1.51 mm. 

Comparative notes.— The new species is 
very similar to Pheneps antennalis Spangler 
& Steiner (1983). All specimens in the type 
series of P. antennalis have a reddish-brown 
head and pronotum which contrasts with 
the dark brown elytra; P. cursitatus is com- 
pletely dark brown dorsally. Also, P. cur- 
sitatus is a smaller species (male length, 2.47 
to 2.97 vs 3.1 to 3.8 mm). The shapes of 
the median lobe and parameres of the male 
genitalia are distinctive (Figs. 11-14). Fe- 
males of P. antennalis are unknown. 

Type-data. — Holotype $ and allotype: 
AMAZONAS: Departamento Rio Negro: 
Cerro de la Neblina, Base camp, Rio Baria, 
00°50'N66°10'W, 140 m, 11 Febmary 1985, 
P. J. and P. M. Spangler, R. A. Faitoute, W. 
E. Steiner; deposited in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution. Paratypes: Same data as holotype, 
90 (5, 3 9; same data except as follows: 27 
January 1985, 2 9; 10 February 1985, 5 <5; 
12 Febmary 1985, 15 <5; 14 February 1985, 
1 (5; 20 February 1985, 1 <5; 5 February 1985, 
W. E. Steiner, 1 9; 6 February 1985, W. E. 
Steiner, 1 $. 

Paratypes will be deposited in the collec- 
tions of the Instituto de Zoologia Agricola, 
Facultad de Agronomia, Maracay, Vene- 
zuela; American Museum of Natural His- 



"^f^ ® 

Figs. 5-10. Pheneps cursitatus, new species. 5, Antenna, male. 6, Antenna, female. 7, Mesostemum, female. 
Mesostemum, male. 9, Abdomen, female. 10, Abdomen, male. 




Figs. 11-14. 11, 12, Pheneps cursitatus. new species. 11, Male genitalia, dorsal view. 12, Male genitalia, 
lateral view. 13, 14. Pheneps antennalis Spangler and Steiner. 13, Male genitalia, dorsal view. 14, Male genitalia, 
lateral view. 



Fig. 15. Biotope; riffle area in Rio Baria, T. F. Amazonas, Venezuela. 

tory, New York; California Academy of Sci- 
ences, San Francisco; Canadian National 
Collection, Ottawa; Institut Royal de His- 
toire Naturelle de Belgique, Bruxelles; Mu- 
seum National de Histoire Naturelle, Paris; 
Museo Argentina de Ciencias Naturales, 
Buenos Aires; Zoologische Sammlung Bay- 
erischen Staates, Munchen; and the collec- 
tion of Harley P. Brown, Norman, Okla- 

Etymology.— The trivial name, cursita- 
tus, is derived from Latin meaning "run 
hither and thither" in reference to the rapid, 
erratic movements of these beetles over the 
rocks in riffles in their habitat. 

Habitat. — Specimens of Pheneps cursi- 
tatus were collected with aerial nets or oc- 
casionally in a seine from bare emergent 
rocks in the riffles; others were attracted to 
black lights operated on the bank of the Rio 
Baria or in a clearing in the rainforest near 
the river. 

In bright sunlight on February 1 1 and 12, 
we found specimens of P. cur sit at us in rath- 
er large aggregations running over rocks in 
an area of shallow riffles in the Rio Baria 
(Fig. 1 5). The water level of the river was 
dropping rapidly during that time and tem- 

perature differences were distinctly evident 
as we waded through the warm, shallow 
water into the cool, deeper water and faster 
current. The beetles were difficult to capture 
because they ran rapidly and took flight 
quickly when attempts were made to cap- 
ture them in an aerial net. The decreasing 
water level and increasing temperature of 
the exposed rocks seems to have triggered 
eclosion from the pupal stage because 66% 
of the specimens obtained were collected 
during those two days. An additional 15 
specimens were collected at our blacklight 
operated on shore beside the riffle on the 
evening of February 1 1 . 

Colorimetric water chemistry analyses 
provided the following data: pH, 4; oxygen, 
12-15 ppm; hardness, 0. When these anal- 
yses were made, the day was sunny, the air 
temperature was 28.5°C, the water temper- 
ature was 24.5°C, and the water around the 
rocks where the beetles congregated was 
about 1 5 cm deep but kept getting shallower 
throughout the day. As the water level 
dropped and the rocks became exposed, the 
psephenid beetles kept moving to other rocks 
still surrounded by water at their bases. The 
elevation at the stream was about 120 m. 




The specimens of this new species were 
collected during a biological survey of Cerro 
de la Neblina. The expedition to Cerro de 
la Neblina Park was organized and directed 
by the Foundation for the Development of 
Physics, Mathematics, and Natural Sci- 
ences of Venezuela, with the patronage of 
the following Venezuelan organizations: The 
Ministry of Education, The Ministry of the 
Environment, the National Council of Sci- 
entific and Technological Research, the 
Venezuelan Air Force, and the National In- 
stitute of Parks. The entire project was co- 
ordinated by Dr. Charles Brewer-Carias and 
was conducted in collaboration with the Na- 
tional Science Foundation of the United 
States, the American Museum of Natural 
History, the Field Museum of Natural His- 
tory, the Missouri Botanical Garden, the 
New York Botanical Garden, and the 

Smithsonian Institution. Biologists from 
several universities and other institutions 
also participated. I thank all of the above 
organizations and their administrators for 
the extensive contributions to the survey. 

I also thank Young T. Sohn for the line 
drawings, Robin A. Faitoute and Heidi Wolf 
for the scanning electron micrographs, Roy 
W. McDiarmid for reviewing the manu- 
script; and Phyllis M. Spangler for typing 
the manuscript. 

Literature Cited 

Darlington, p. J. 1936. A list of the West Indian 
Dryopidae (Coleoptera) with a new genus and eight 
new species, including one from Colombia. Psyche 
43: 64-83. 

Spangler, P. J. and W. E. Steiner. 1983. New species 
of water beetles of the genera Elmopamus and 
Pheneps from Suriname (Coleoptera: Dryopidae: 
Psephenidae). Proceedings of the Entomological 
Society of Washington 85: 826-839. 


89(2), 1987, pp. 226-229 


W. R. M. Mason 

Biosystematics Research Centre, Agriculture Canada, Ottawa, Ontario KIA 0C6, Can- 

Abstract.— Freviously unknown, the female of Apozyx penyai Mason (Apozygidae) is 
found to be very similar to the male. Several more males from south-central Chile are 
reported. A further examination of characters of Apozyx shows it to be undoubtedly a 
member of Ichneumonoidea but not of Microhymenoptera nor Aculeata. Differences 
between Apozygidae and Braconidae are tabulated. 

In the eight years since I described Apozyx 
penyai Mason (Mason, 1978) several more 
males and a female have been collected in 
Chile. Additional males have been taken in 
Nuble (Las Trancas, 5 km E.S.E. of Recinto 
at 1250 m) in a Malaise trap operated by 
A. Newton and M. Thayer in Dec. 1982 in 
Nothofagus forest. Later a male and a female 
were taken in Nahuelbuta National Park, 
50 km W. of Angol (12-1500 m, flight in- 
tercept trap operated from Dec. 1984 to 17 
Feb. 1985 by S. and J. Peck), the site being 
in Nothofagus-Araucaria forest. 

Apozyx penyai Mason, 
Supplementary Description 

Description of male, see Mason (1978). 

The sexes are very similar except for the 
large ovipositor (Fig. 4), the stouter flagellar 
articles, and smaller eyes of the female (Figs. 

Abdomen of 9 (Fig. 4) resembling that of 
$ but hypopygium large and triangular in 
profile. Ovipositor sheaths hairy, slightly 
deeper apically, about as long as metasoma; 
ovipositor weakly and evenly curved up- 
ward, uniformly deep except subapically 
where it is slightly deeper, the apex sharply 

pointed and bearing a few small teeth ven- 

Ragellum of 9 with 1 7 articles, about 20% 
shorter than that of 6\ antipenultimate ar- 
ticle 1.4 times as long as wide (in $ 2.3-2.8); 
second article of female 3 times as long as 
wide (in <3 2.5-3.5). Compound eye in 9 con- 
siderably smaller and less protuberant than 
in 3 (Figs. 1, 2); thus in dorsal or anterior 
aspect outer part of the eyes slightly closer 
than temples (in $ eyes bulging distinctly 
farther apart than temples) and the vertex 
elevated above top of eyes for a distance 
almost equal to eye height (in $ vertex ele- 
vated for a distance about half eye height). 

In both sexes second abdominal (first 
metasomal) sternum including 2 parts: 1, 
an anterior heavily sclerotized section fused 
with the tergite into a tubular structure with 
component parts not easily distinguishable; 
2, a small lenticular posterior section formed 
of 2 subtriangular plates separated medially 
by narrow membranous area (Figs. 5, 6). 
During flexion of the part of the abdomen 
behind tergum 2 this pair of plates overlaps 
externally the tubular fused tergite and ster- 
nite 2. Metasomal terga 1 and 2 articulate 
laterally by pair of submarginal condyles 



Figs. 1-4. Apozyx penyai. 1, 2, Head of female, frontal and lateral view. 3, Paramere and volsella, inner 
side. 4, Female abdomen. 

(Fig. 5). Sterna not involved in articulation. 
Because sterna of holotype could not be 
counted, original description fails to men- 
tion that the metasomal stemites 2 and 3 
are fused exactly as their respective tergites. 
On p. 609, Hne 9, the terms (2r-m) and (3r- 
m) should be corrected to (Ir-m) and (2r- 
m) respectively. 

Male genitalia (Fig. 3) with weakly ex- 
panded and rounded parameres. Cuspis 
broad, truncate and very short, digitus ta- 
pered, extending far beyond cuspis. 


In 1978, 1 was uncertain about the place- 
ment of this family but studies during in- 
tervening years and recent knowledge of the 

female anatomy have confirmed my first 
opinion. Apozyx shares the following char- 
acter states with other Ichneumonoidea: 1 , 
flagellar articles with large longitudinal 
placodes; 2, mandibles with two apical teeth; 
3, pronotum much shorter medially than 
laterally, the posterodorsal comers touching 
the tegulae; 4, pronotum (plus prepectus) 
rigidly attached to the mesoscutum and 
mesopleuron; 5, posterodorsal comer of 
pronotum developed into a lobe behind 
which lies the mesothoracic spiracle; 6, pre- 
pectus fused to the posterior vertical pro- 
notal margin, the pit that indicates exter- 
nally the site of the apodeme bearing the 
origin of the spiracular occlusor muscle lying 
very near the posterior margin of the fused 



Figs. 5, 6. Apozyx penyai. junction of abdominal segments 2 and 3, showing abdominal tergites 2 and 3 (T2, 
T3), sternites 2 anterior (S2a) and posterior (S2p), stemite 3 (S3), and condyle (C). 5, Anterolateral view. 6, 
Ventrolateral view. 

pronotum plus prepectus (for definition of 
prepectus and discussion of characters 4-6, 
see Gibson, 1985); 7, costa and radius of 
fore wing fused; 8, sternum of metasoma 
one divided into a heavily sclerotized an- 
terior section and comparatively weakly 
sclerotized posterior section (Mason, 1981, 
1983); 9, metasomal segments 1 and 2 ar- 
ticulated by a pair of sublateral condyles on 
the hind margin of tergum 1 and anterior 
margin of tergum 2, the sterna playing no 
part in the articulation. 

The presence in the female of complete 
metasomal terga 7 and 8, the latter bearing 
large cerci, eliminates /1/7c»zva' from the Acu- 
leata (Koenigsmann, 1978). The presence of 
spiracles on each of abdominal tergites 1 to 
8 as well as the step-like posterior deflection 
of fore wing Cu at its junction with m-cu 
eliminates Apozyx from Microhymenop- 
tera because all Microhymenoptera have a 
comparatively straightened Cu (if it is not 
reduced to invisibility) and their abdomen 
has only 2 pairs of spiracles on terga 1 and 

8, or only one pair on tergum 1 (propo- 
deum), the other spiracles being absent or 

The general aspect of Apozyx is similar 
to that of a braconid but there are important 
differences: 1 , in Apozyx abdominal sterna 
3 and following are rounded and sclerotized 
as strongly as the terga, a basic condition 
shared with most other Hymenoptera, 
whereas the same sterna of Braconidae are 
extensively soft, membranous and usually 
divided into several weakly sclerotized 
plates, a derived condition found only in 
Braconidae, Ichneumonidae and Paxylom- 
matidae; 2, vein 2r-m (2nd recurrent) is 
present as a strong tubular vein in the fore 
wing, a basic condition (Mason, 1981), 
whereas in Braconidae this vein is missing, 
a derived condition. 


My thanks go to the technicians of the 
Electron Microscope Center (Canada Ag- 
riculture) and to H. E. Bisdee, B.R.C., for 



assistance in making the illustrations. I also 
thank my colleagues J. R. Barron and M. J. 
Sharkey for suggestions and review of the 

Literature Cited 

Gibson, G. A. P. 1 985. Some pro- and mesothoracic 
structures important for phylogenetic analysis of 
Hymenoptera, with a review of terms used for the 
structures. Can. Entomol. 1 17: 1395-1443. 

Koenigsmann, E. 1978. Das phylogenetische system 

der Hymenoptera, 4. Dtsch. Entomol. Zeit ^5- 

Mason, W. R. M. 1978. A new genus, species and 
family of Hymenoptera (Ichneumonidae) from 
Chile. Proc. Entomol. Soc. Wash. 80: 606-610. 

. 1981. Paxylommatidae: The correct family- 
group name for Hybrizon Fallen (Hymenoptera: 
Ichneumonoidea), with figures of unusual anten- 
nal sensilla. Can. Entomol. 113: 433-439. 

. 1983. A new South African subfamily related 

to Cardiochilinae (Hymenoptera: Braconidae). 
Contrib. Am. Entomol. Inst. 20: 49-62. 


89(2), 1987, pp. 230-241 




A. J. Ritchie and J. D. Shorthouse 

(AJR) Department of Biology, Carleton University, Ottawa, Ontario (Current address: 
47 Deepwood Drive, Hamden, Connecticut 06417); (JDS) Department of Biology, Lau- 
rentian University, Sudbury, Ontario P3E 2C6, Canada. 

Abstract.— Three new species ofSynergus Hartig (Hymenoptera: Cynipidae) from Gua- 
temala arc described and Synergus ftliconiis Cameron is redescribed. A key to S. cultratus 
new species, S. kinseyi new species, S. mesoamericanus new species, and S. filicornis 
Cameron is presented. Biological notes, when available, are given for each species. Cynips 
guatemalemis Cameron is transferred to Andricus and is a senior synonym of A ndricus 
me.xicana Bassett, A. ?m'xicaniis Kinsey new synonym, and Synergus dorsalis Cameron 
new synonym. The nomenclature for Synergus dorsalis Cameron and S. dorsalis (Pro- 
vancher) is discussed. 

During a visit to the American Museum 
of Natural History, one of us (AJR) discov- 
ered a series of undescribed inquiline gall 
wasps belonging to the genus Synergus Har- 
tig (Hymenoptera: Cynipidae) from Gua- 
temala. These species are of interest for sev- 
eral reasons. Guatemala is the southernmost 
point of the range of inquiline cynipids in 
the New World and is isolated from other 
cynipid habitats by the lowland Isthmus of 
Tehuantepec. Secondly, some of the most 
primitive oak gall wasps come from Gua- 
temala and southern Mexico. Finally, two 
of the three new species are extremely large 
for inquiline Cynipidae. 

Very little is known about the gall wasps 
of Guatemala. Cameron (1883) described 
the woody oak-stem galls of Cynips cham- 
pioni and C. imitator and the oak-leaf galls 
of C guatemalensis. Cameron described also 
two species of Synergus from Guatemala: 
S. filicornis from an unknown gall, and S. 
dorsalis from galls of C. guatetualensis. 

KJnsey (1920) cut insects from galls that he 
considered to be made by C. guatetualensis. 
Kinsey (1936) later described Acraspis fu- 
giens, Atrusca lucaris, and Atrusca lumi- 
naris, all of which form leaf galls on oaks. 
Weld (1913) described Synergitsfurnessana 
from Mexico but Weld (1930) later decided 
that this was synonymous with S. filicornis 
Cameron. The only other inquiline known 
from Mexico is S. dugesi Ashmead (holo- 
type in USNM; examined), which has been 
reared from galls of Andricus championi 

Kinsey made important contributions to 
our knowledge of the taxonomy and biology 
of gall wasps. Although his taxonomic con- 
cepts and nomenclature were radical and 
have not been followed since, his collection 
of Cynipidae, now in the American Mu- 
seum of Natural Histor>', is one of the most 
important in the world. He was an avid col- 
lector, and some estimates of the size of his 
collection reach 5 million specimens (Weld, 



1952). Kinsey never described or reported 
any of the inquilines or parasites that he 
reared from galls. He exhaustively labelled 
them with locality, collection and emer- 
gence dates, host plant and host gall, but 
curated them under the name of the host 
gall inducer. Unfortunately, he assigned 
manuscript names to many species of gall 
inducer which he never described. This is 
apparently the case with the Guatemalan 
gall wasps, as two of the four galls from 
which he reared inquilines cannot be posi- 
tively identified; one (brelandi) is a Kinsey 
manuscript name (Weld, 1952), and the 
other (ruginos.) is abbreviated. Tentative 
host identifications are discussed in the 
remarks for S. cultmtus. Collection dates 
were given in the following form, 
"month. day. year," and the symbol © was 
used to indicate the collection date of the 
gall and "inq." for the date of emergence 
for an inquihne. Kinsey usually put a num- 
ber and letter after the locality; these indi- 
cate the number of miles and direction from 
the nearest town (i.e. Sacapulus 9S means 
9 miles south of Sacapulus). This has been 
followed in the present paper. 

Morphological terms are those of Ritchie 
and Peters (1981) and Shorthouse and Rit- 
chie (1984), with the following exceptions: 
supracoxal carina = carina on propodeum 
running from metacoxal socket dorsally to 
anterior of the flap over propodeal spiracle; 
metanotal median fovea = the small me- 
dian fovea posterior and between the lateral 
foveae, and above the area between the pro- 
podeal carinae. 

The majority of specimens examined in 
this study, including the holotypes of the 
new species, are deposited in the American 
Museum of Natural History, New York, 
New York (AMNH) except as noted. The 
holotypes of Cynips guatemalensis Cam- 
eron and Synergus dorsalis Cameron are de- 
posited in the British Museum (Natural 
History) (BMNH), the holotype of Cer- 
optres dorsalis Provancher is in the Public 

Museum of Quebec, Quebec City, Quebec 
(PMQ), and the holotype of Synergus fur- 
nessana Weld is in the Philadelphia Acad- 
emy of Natural Sciences, Philadelphia, 
Pennsylvania (PANS). Kinsey collected all 
specimens and made all host-gall determin- 
ations except as noted. All Kinsey material 
was collected from galls on Quercus pili- 
caulis. Representative specimens of the four 
Guatemalan species of Synergus have been 
retained by both authors. 

Key to the Species of 
Synergus from Guatemala 

1 . Female metasomal tergite 2 + 3 (T2 + 3) deeply 
excavated along posterodorsal margin, poster- 
ior third sparsely and weakly punctate (Figs. 
9, 10); meso- and metasoma black and white, 
black and yellow, or entirely yellow 2 

- Female T2 + 3 at most weakly excavated along 
posterodorsal margin, posterior third entirely 
smooth (Figs. 11, 12); mesosoma entirely black, 
metasoma black and rufous 3 

2. Crossvein r-m in hind wing recurved (Fig. 1 7); 
radial sector (Rs) 1 and Rs2 nearly parallel (Fig. 
19); third male antennomere (A3) elongated 
beyond notch (Fig. 13); mesopleuron entirely 

yellow above coxal bases 

cultratus Ritchie and Shorthouse, n. sp. 

- Crossvein r-m in hind wing normal (as in Fig. 
18); Rsl and Rs2 divergent (Fig. 20); male A3 
not elongated beyond notch (Fig. 14); meso- 
pleuron with some black above coxal bases . 
filicornis Cameron 

3. Female with 1 5 antennomeres; radial cell length 
3.0 times width; Rs2 straight (at most weakly 
bent distally); areolet absent or elongate (Fig. 

21); usually over 5.0 mm long 

kinseyi Ritchie and Shorthouse, n. sp. 

- Female with 14 antennomeres; radial cell length 
usually 2.0 times width; Rs2 distinctly bent; 
areolet equilateral (Fig. 22); under 4. 1 mm . . 

. . mesoamericanus Ritchie and Shorthouse, n. sp. 

Synergus cultmtus 
Ritchie and Shorthouse, 

New Species 
Figs. 1, 5,9, 13, 17, 19 

Female (holotype). — Length 7.0 mm. 
Yellow; tip of mandible, frons above anten- 
nal socket and between frontal ridges, me- 
dian third of vertex and occiput, pronotal 





plate, scutum between anterior lines, meso- 
pleuron between coxal sockets, anterodorsal 
margin of T2 + 3, posterodorsal margin of 
T7 black; distal half of mandible except tip, 
A1-A7, tarsal claw, Tl, posterodorsal mar- 
gin of T2 + 3, exposed dorsal margins of T4- 
T6 piceous brown. Head slightly narrower 
than thorax (slightly wider in all other 
species); malar space 0.56 of eye height; 
frontal ridge weak, reaching ectal margin of 
posterior ocellus; LOL = 0.8D, POL = 1 .4D, 
OOL = 1.4D; antenna with 14 antenno- 
meres, 5.29 mm long; A1-A14 (mm): 0.33, 
0. 1 7, 0.56, 0.57, 0.58, 0.56, 0.46, 0.40, 0.35, 
0.29, 0.25, 0.25, 0.21, 0.31. Mesoscutum 
(Fig. 1) transversely rugulose, punctate be- 
tween anterior lines; notauli percurrent, ir- 
regularly interrupted by transverse rugulae; 
median groove represented by weak crease 
extending about one third of mesoscutum; 
anterior lines slightly divergent, extending 
about one fourth of mesoscutum. Meso- 
scutellum (Fig. 1) rugose, rugosity stronger 
posteriorly, median depression absent; up- 
turned posterior carina strong, complete; 
fovea large, shallow, rugulose. Mesopleuron 
(Fig. 5) entirely striate. Metanotal median 
fovea with longitudinal carina. Propodeum 
with an extra weak carina parallel to and 
between supracoxal carina and propodeal 
carina; propodeal carinae very strong, thick- 
ened, strongly coriaceous. Metasoma (Fig. 
9) longer than head plus mesosoma. T2 + 3 
covering about two thirds of metasoma (lat- 
eral view), strongly excavated along pos- 
terodorsal margin, posterior third sparsely, 
weakly punctate. Forewing (Fig. 1 9) 6.0 mm 
long; radial cell length 3.0 times width; Rsl 
nearly parallel to Rs2 (divergent in all other 
species); Rs2 very weakly bent near wing 
margin, almost straight; areolet normal; hind 
wing 4.0 mm long, r-m strongly recurved 

(Fig. 17) (weakly recurved in all other 
species; Fig. 18). Tarsal claw with distinct 

Male (allotype). — Differs from holotype 
as follows: length 4.75 mm; antenna 4.9 mm 
long, with 1 5 antennomeres, A 1-A 1 5 (mm): 
0.38, 0. 15, 0.50, 0.50, 0.50, 0.50, 0.42, 0.33, 
0.31, 0.26, 0.23, 0.21, 0.21, 0.18, 0.22; A3 
weakly bent and notched (Fig. 13), elon- 
gated beyond notch; T2 + 3 covering entire 
metasoma (dorsal view), T7 (posterior view) 
with setigerous punctures; forewings 5.6 mm 
long, hind wings 3.4 mm long. 

Variation. — F^'m^/^'." Length 5.0-7.25 
mm, average of 50 specimens = 6.3 mm; 
mesoscutum from yellow with small black 
area between anterior lines to black with 
small yellow areas at posterior ends of no- 
tauli; scutellum sometimes amber or light 
brown; propodeal carinae yellow or black, 
black area sometimes extending ectal of ca- 
rinae; mediodorsal black area of metasoma 
sometimes expanded to occupy entire dor- 
sal margin. 

Male: Length 3.3-5.1 mm, average of 50 
specimens = 4.4 mm; coloration as in fe- 
male except propodeum, mesoscutum, 
mesoscutellum, and metapleuron some- 
times entirely black. 

Type material. — Holotype, 2 (AMNH): 
Huehuetenango 3S, Guate[mala], 7300', ® 
12.22.35, inq. 8.18.37, ex gall of brelandi. 
Allotype, 6: with same data as holotype. 
Paratypes (142 9, 145 <5; AMNH): 71 9, 74 
S with same data as holoypte; 13 9, 16 3 with 
same data as holoytpe but emerged 7.1- 
9.10.36; 20 9, 26 6 with same data as ho- 
lotype but emerged 7.1-9.1.38; 13 9, 3 3 
with same data as holotype but emerged 
8.22-8.27.37, oxgall of ruginos.\ 5 9, 10 S, 
Sacapulus 9S, Guate., 6000', 12.28.35, 
inq. 7.3-10.22.37, oxgall of brelandi; 1 9, 

Figs. 1-8. 1-4, Mesonota of Synergus spp., dorsal view. 1, Synergus cultratus. 2, Synergus filicornis. 3, 
Synergus kinseyi. 4, Synergus mesoamericanus. 5-8, Mesopleura of Synergus spp., lateral view. 5, Synergus 
cultratus. 6, Synergus filicornis. 7, Synergus kinseyi. 8, Synergus mesoamericanus. 



1 $ with same data as preceding scries but 
emerged 9.1.38; 1 9 with same data as pre- 
ceding series but emerged 7.1.39; 2 9, 2 <5, 
Quiche 2N, Guate., 7500', © 12.27.35, inq. 
8.10-9.10.37, ^.v gall of hrelandi: 4 9, 2 5 
with same data as preceding series but 
emerged 7.1-7.15.38; 1 9, Guatemala City 
3W. Guate., 6300', © 12.30.35. inq. 8.3.37, 
e.\ gall of brelandi\ 11 9, 1 6 (Label lost), 
Guatemala. 1935, inq. 9.20.37, ^.v gall of 

Excluded from type series: 89 specimens 
from the above localities and emergence 
dates, in various but poor condition; ten- 
tatively identified as 5". cidtratus. 

Etymology. — The species name is based 
on the Greek word for knife in reference to 
the strongly compressed metasoma. 

Diagnosis.— 5'r/?£'/'g//5 cidtratus can be 
distinguished easily from other Guatemalan 
species by the following characters: r-m in 
the hind wing strongly recurved; Rsl nearly 
parallel to Rs2; mesopleuron entirely yel- 
low; male A3 elongate beyond notch; and 
the extra carina between the supracoxal and 
propodeal carinae. Some specimens of this 
species are the largest known inquiline cyni- 
pids. being over 7.0 mm in length. 

Remarks.- Host relationships for S. cul- 
t rat us are unclear, as neither of the reported 
hosts on the labels can be definitely iden- 
tified. One labelled host, brelandi. is a Kin- 
sey manuscript name for a species of An- 
dricus (Weld, 1952). The other labelled host. 
ruginos.. may be an abbreviation for An- 
dricus ruginosus Bassett. which produces a 
hard polythalmous stem gall (Weld. 1957) 
or more likely, a variety of ruginosus (see 
Kinsey. 1930. 1936 for his taxonomic con- 
cepts and use of varietal names). The emer- 
gence dates for this species are also unusual, 
as most species ofSynergus emerge after one 
or two winters in the gall (Eady and Quin- 
lan. 1963). Synergus cultratus appears to 
have a one to four year variable life cycle, 
although it is possible that first year emer- 
gents were from older galls and that the life 
cycle is from two to four years in length. 

Most individuals (65% of types) emerged 
after two winters in the gall. 

Synergus filicornis Cameron 

Figs. 2, 6, 10, 14, 20 

Synergus filicornis Cameron, 1883: 71 (9, 
$). TYPE: Holotype, 9 (BMNH). Exam- 

Synergus furnessana Weld, 1913: 134(9,3); 
synonymized by Weld, 1930: 143. TYPE: 
Holotype, + (PANS). Examined. 

Female. — 2.0-3.6 mm. Black and yellow; 
Head yellow; frons above antennal socket 
and between frontal ridges, ocellar area, me- 
dian third of vertex, median third of occiput 
usually black (rarely entirely yellow); anter- 
ior of pronotum, anterior of pronotal plate 
piceous brown to black; posterior of pro- 
notal plate yellow to black, remainder of 
pronotum yellow to amber; mesoscutum 
usually yellow or amber, with black or pi- 
ceous brown between notauli (rarely entire- 
ly yellow or mostly piceous brown); meso- 
scutum usually yellow to amber, rarely dark 
piceous brown; mesopleuron black, fre- 
quently with median yellow area, rarely en- 
tirely yellow; metanotum and metapleuron 
usually black, sometimes piceous brown, 
rarely yellow; propodeum piceous brown to 
black, rarely yellow; spiracular flap almost 
white, always lighter than surrounding col- 
oration; metasoma yellowish, with broad 
dorsal band piceous brown to black, band 
sometimes reduced to anterior half of T2 + 3; 
legs yellow, metatarsus amber to brown; 
wings hyaline, veins yellow to light brown. 
Head as wide or slightly wider than thorax; 
frons below antennal socket and beneath 
eye with strong radiating striae (relatively 
stronger than in other species), striae weaker 
between anterior tentorial pits; malar space 
about 0.54 of eye height; postero ventral 
margin of gena with weak short carina; fron- 
tal ridge present, moderately strong, not 
quite reaching posterior ocellus; LOL = 
0.91D. POL = 2.0D. OOL = 1.36D; anten- 
na with 14 antennomeres; A3-A6 relatively 



thin, subsequent antennomeres stouter; rel- 
ative length of antennomeres (A1-A14): 
0.25, 0. 10, 0.32, 0.32, 0.32, 0.29, 0.25, 0.21, 
0. 1 7, 0. 1 5, 0. 1 3, 0. 1 2, 0. 1 2, 0.20. Mesoscu- 
tum (Fig. 2) transversely rugulose; notauli 
percurrent, grooves sometimes interrupted 
by transverse rugulae, bottom of grooves 
weakly rugulose; median groove, lateral lines 
absent; anterior lines parallel, extending 
about one fourth of mesoscutum. Mesoscu- 
tellum (Fig. 2) rugose, median depression 
absent, weak posterior carina not upturned; 
fovea large, shallow, sometimes weakly ru- 
gulose, not well defined. Mesopleuron (Fig. 
6) entirely striate, striations stronger ven- 
trally. Metanotal median fovea broadened, 
broader than in other species. Propodeum 
with moderately weak transverse carinae 
betwen supracoxal and propodeal carinae, 
no extra longitudinal carina; propodeal ca- 
rinae very strong, thick, strongly coriaceous. 
Metasoma (Fig. 1 0) slightly longer than head 
plus mesosoma; T2 + 3 covering about two 
thirds of metasoma, posterior third punc- 
tate, posterodorsal margin deeply excavat- 
ed; T4-T7 exposed along posterodorsal 
margin, more densely punctate than T2 + 3. 
Forewing (Fig. 20) about 1.0-1.1 length of 
body; radial cell length 3.0 times width; Rs2 
very weakly bent near wing margin, almost 
straight; areolet of normal shape, posterior 
margin very weak to missing. Tarsal claw 
with a distinct tooth. 

Male. — Differs from female as follows: 
length 2.0-2.5 mm; frontal ridge sometimes 
complete, running to anterior margin of 
posterior ocellus; antenna with 15 anten- 
nomeres, about equal to body length, A3 
prominently notched and bent (Fig. 14), rel- 
ative lengths of antennomeres (A1-A15): 
0.12, 0.11, 0.11, 0.10, 0.10, 0.09, 0.13; 
mesoscutum sometimes entirely black, with 
stronger transverse rugosity; mesopleuron 
sometimes entirely black, with stronger 
striae, propodeal carinae not as strong or as 
thickened, not as strongly coriaceous; meta- 
soma sometimes entirely black; T2 + 3 cov- 

ering entire metasoma, not posterodorsally 
exavated; posterior third of T2 + 3 usually 
with band of punctures interrupted by pos- 
terodorsal smooth area, punctures some- 
times stronger and band not interrupted. 

Type material examined.— Synergus fili- 
cornis Cameron. Holotype, 9 (BMNH, Type 
No. 7.1 10), labelled: "San Geronimo, Gua- 
temala, Champion," '"Synergus filicomis." 
Synergus furnessana Weld. Holotype, 9 
(PANS), "Michoacan, Mex.," ""Synergus 
furnessana,'''' "Holotype" (red label). Allo- 
type, 3 (PANS), with same data as holotype. 

Other material examined (AMNH).— 18 
9, 1 <5, Huehuetenango 14S, 7500', Guate., 
e 12.23.35, inq. 1.7-1.14.36, ex gall of 
mexicanus; 1 9, 1 3, as previous series but 
emerged 7.1.36; 23 9, 1 S, Sacapulus 9S, 
6000', Guate., e 12.28.35, inq. 1.7-1.14.36, 
oxgall of mexicanus; 10 9, 16, Quiche 2N, 
7500', Guate.,® 12.27.35, inq. 1.7-1.22.36, 
ex gall of mexicanus; 1 6, Guatemala City 
5W, 6500', Guate., ® 12.30.35 (no emer- 
gence date), ex gall of mexicanus. 

Diagnosis. — Synergus filicornis Cameron 
is most similar to S. cultratus and can be 
distinguished from it and other species by 
the following characters: under 4.0 mm; 
posterior third of T2 + 3 punctate, postero- 
dorsal margin of female T2 + 3 deeply ex- 
cavated; A3-A5 elongate and thin. The type 
of Synergus filicornis is in poor condition; 
the following structures are missing: left A2- 
A14, right A8-A14, and the metasoma. 

Remarks.— This species is the most vari- 
able of the Guatemalan species of Synergus, 
especially in coloration. Very small females 
(~2.0 mm) are almost entirely yellow and 
have much finer sculpture of the mesopleu- 
ron and punctures on T2 + 3, whereas some 
of the larger females have much more brown 
or black and have stronger sculpturing. Sim- 
ilarly, two of the larger males have the meso- 
and metasoma entirely black, very strong 
mesopleural striations, and have a complete 
band of punctations in the posterior third 
of T2 + 3. Unlike the other species, S. fili- 
cornis has been reared from both a leaf gall, 









Andricus guatemalensis (Cameron), and a 
stem gall, A.furnessana Weld. The life cycle 
of this species appears to be limited to one 
year and the rapid emergence of some spe- 
cimens may indicate that S. filicornis has 
more than one generation per year. 

Synergus kinseyi Ritchie and 
Shorthouse, New Species 
Figs. 3, 7, 11, 15, 18, 21 

Female (holotype). — Length 6.25 mm. 
Head yellow; frons above antennal sockets 
median third vertex, and median third of 
occiput black; antenna and mandible brown. 
Mesosoma black, legs except metacoxal 
bases yellow, wings hyaline, veins brown. 
Metasoma rufus except for dorsal margin of 
T2 + 3 and exposed dorsal margins of T4- 
T7 black. Head slightly wider than meso- 
soma; malar space 0.73 of eye height; pos- 
teroventral margin of gena with short, weak 
carina; frontal ridge absent; LOL = 0.8D, 
POL = 1.7D, OOL = 2.0D; antenna with 
15 antennomeres, 3.88 mm long; A1-A15 
(mm): 0.42, 0.21, 0.49, 0.31, 0.33, 0.39, 
0.27, 0.25, 0.20, 0.20, 0.20, 0. 1 7, 0. 1 7, 0. 1 7, 
0.20. Mesoscutum (Fig. 3) with weak trans- 
verse rugulae; notauli percurrent, grooves 
shining smooth, not interrupted by trans- 
verse rugulae, distinctly widened posterior- 
ly; median groove, lateral lines absent; an- 
terior lines slightly convex, extending about 
one third of mesoscutum. Mesoscutellum 
(Fig. 3) weakly rugose, median longitudinal 
depression and posterior upturned carina 
well developed; fovea well developed, dis- 
tinctly closed posteriorly. Mesopleuron (Fig. 
7) aciculate above, weakly striate below. 
Metanotal medial fovea without longitu- 
dinal carina. Propodeum ruglose; supracox- 

al carina strong; propodeal carinae strong, 
not thickened, finely coriaceous. Metasoma 
(Fig. 1 1) longer than head plus mesosoma; 
T2 + 3 covering about three fourths of meta- 
soma, weakly excavated along posterodor- 
sal margin, posterior third glabrous. Fore- 
wing (Fig. 21) 5.25 mm long; radial cell 
length 3.0 times width; Rs2 weakly but dis- 
tinctly bent near wing margin; areolet ab- 
sent; hind wing (Fig. 18) 3.6 mm long. Tar- 
sal claw with a distinct tooth. 

Male (allotype). — Differs from holotype 
as follows: length 4.25 mm; malar space 0.53 
of eye height; antenna 3.41 mm long; Al- 
A15 (mm): 0.32, 0.17, 0.42, 0.25, 0.25, 0.25, 
0.2 1 ; A3 (Fig. 1 5) slightly notched and bent, 
elongate beyond notch; anterior lines more 
distinct; median groove reduced to notch 
between raised ental margins of notauli; 
mesopleuron evenly, finely striate; supra- 
coxal carina weaker; propodeal carinae not 
as strong; T2 + 3 covering entire metasoma, 
dorsal black area larger, yellowish ventrally; 
radial cell length about 2.7 times width. 

Variation. — Fe-m^/e.- Length 4.75-6.7 
mm, average of 50 specimens = 5.8 mm; 
head sometimes amber; areolet absent or 
elongate, posterior margin very weak (both 
sexes); metasoma from rufus to dark rufus, 
dorsal black area sometimes expanded ven- 

Male: Length 3.5-4.5 mm, average of 34 
specimens = 4.2 mm; coloration similar to 
female except metasoma ventrally dark ru- 
fus to dark amber, dorsal black area some- 
times expanded. 

Type material. — Holotype, 2, Huehue- 
tenango 3S, Guate., 7300', 12.22.35, Q. 
pilicaulis, Kinsey coll., ex gall of ruginos., 

Figs. 9-18. 9-12, Metasoma of Synergus spp., lateral view. Scale bar = 1.0 mm. 9, Synergus cultratus. 10, 
Synergus filicornis. 11, Synergus kinseyi. 12, Synergus mesoamericanus. 13-16, Male antennomere 3 (A3) of 
Synergus spp., lateral view. Scale bar = 0.1 mm. 13, Synergus cultratus. 14, Synergus filicornis. 15, Synergus 
kinseyi. 16, Synergus mesoamericanus. 17, 18, Hind wings of Synergus spp. Scale bar = 1.0 mm. 17, Synergus 
cultratus. 18, Synergus kinseyi. 



Figs. 19-22. Forewings (radial cell) of Synergus spp. 19, Synergus cultratus. 20, Synergus filicornis. 21, 
Synergus kinseyi (inset shows variant without areolet). 22, Synergus mesoamericanus. 

Kinsey det. Allotype, 5, with same data as 
holoytpe. Paratypes (86 9, 34 $: AMNH): 
36 2, 12 3 with same data as holotype; 45 
$, 22 (5, Sacapulus 9S, Guate., 6000', © 
12.28.35, inq. spr. '37, ex gall oi brelandi; 
5 9, (Label lost), Guatemala, 12.36, fall '36, 
ex gall o{ peredurus. Excluded from type se- 
ries: 133 specimens from above localities, 
© 12.22-28.35, inq. fall '36-spr. '37; in var- 
ious but poor condition; tentatively iden- 
tified as S. kinseyi. 

Etymology.— This species is named for 
Dr. A. C. Kinsey to commemorate his con- 
tribution to our knowledge of the Cynipi- 

Y)'\2ignos\s.— Synergus kinseyi is easily 
distinguished from other Guatemalan 
species by the following characters: areolet 
absent or elongate with the posterior margin 
very weak; mesopleuron entirely black; fe- 
male antenna with 15 antennomeres. This 
species is also very large, exceeded only by 
S. cultratus in size. 

Ktmdir^is.— Synergus kinseyi has been 
reared from three hosts and appears to be 
restricted to woody stem galls, two of which 
(brelandi and ruginos. ) are the same as those 
for S. cultratus (see remarks for 5". cultratus 
for problems in identifying the hosts). The 
third recorded host, Andricus peredurus 
Kinsey, produces a hard polythalmous stem 
gall. The presence of /I. peredurus in Gua- 
temala is a major range extension, as this 
species previously known only from San 
Louis Potosi, Mexico (Kinsey, 1920). 

There is a contradiction in the labelling 
of the five specimens reared from the galls 
of /4. peredurus, in that the label shows them 
being collected in December of 1936 and 
emerging in the fall of 1936. It is known 
that Kinsey collected in Guatemala during 
1935 (Kinsey, 1936). Therefore, it is prob- 
able that these insects were collected in 1 935 
and emerged in the fall of 1936. This agrees 
with the dates of some of the specimens 
excluded from the type series. Less is known 



about the life cycle of this species than for 
other Guatemalan Synergus spp., as only 
one series has emergence dates. These data 
indicate that S. kinseyi has a one to two year 
life cycle. 

Synergus mesoamericanus 

Ritchie and Shorthouse, 

New Species 

Figs. 4, 8, 12, 16,22 

Female (holotype). — Length 3.4 mm. 
Coloration similar to S. kinseyi. Head 
slightly wider than mesosoma; malar space 
0.57 of eye height; posteroventral margin of 
gena with moderately long carina; frontal 
ridge absent; LOL = 0.9D, POL = 1.7D, 
OOL = 2. 3D; antenna with 14 antenno- 
meres, 2.75 mm long; A1-A14 (mm): 0.29, 
0.17, 0.17, 0.15, 0.15, 0.23. Mesoscutum 
(Fig. 4) with strong transverse rugulae 
(stronger than in other species); notauli 
strong, percurrent, bottom of grooves 
smooth, not interrupted by transverse ru- 
gae; median groove, lateral lines absent; an- 
terior lines convergent, extending about one 
third of mesoscutum. Mesoscutellum (Fig. 
4) strongly rugulose, posterior of disc de- 
pressed, posterior upturned carina moder- 
ately developed; fovea large, deep anterior- 
ly, open posteriorly. Mesopleuron (Fig. 8) 
entirely striate, striations stronger ventrally. 
Metanotal median fovea without longitu- 
dinal carina. Propodeum with supracoxal 
carina weak ventrally, stronger dorsally; 
propodeal carinae moderately strong, not 
thick, finely coriaceous. Metasoma (Fig. 1 2) 
longer than head plus mesosoma; T2 + 3 
covering about three fourths of metasoma, 
weakly excavated along posterodorsal mar- 
gin, posterior third smooth. Forewing (Fig. 
22), 3.8 mm long; radial cell length 2.0 times 
width, Rsl very weak along anterior margin 
of wing (distinct in other species); Rs2 
strongly bent near wing margin; areolet nor- 
mal; hind wing 2.4 mm long. Tarsal claw 
with a distinct tooth. 

Male (allotype). — Differs from holotype 

as follows: length 3.7 mm; antenna 3.05 mm 
long, with 1 5 antennomeres, A 1 -A 1 5 (mm): 
0.21, 0.17, 0.17, 0.17, 0.15, 0.15, 0.17; A3 
(Fig. 1 6) notched and bent, not elongate af- 
ter notch; upturned carina on posterior mar- 
gin of scutellum not as strong; T2 + 3 cov- 
ering entire metasoma, black dorsally, dark 
rufus ventrally; forewing 3.8 mm long, hind 
wing 2.5 mm long; radial cell length 2.4 
times width. 

Variation. — Female length 3.1-4.0 mm, 
average of 5 specimens = 3.7 mm; male 
length 2.75-3.7 mm, average of 2 specimens 
3.2 mm; head from yellow to amber; meso- 
scutum and mesoscutellum from black to 
dark rufus; mesoscutellum sometimes with 
fovea partly closed posteriorly, a weak me- 
dian depression, and posterior upturned ca- 
rina stronger; radial cell sometimes more 
elongate with Rs2 less strongly bent, ante- 
rior margin sometimes indistinctly closed. 

Type Material. — Holotype, 9, Sacapulus 
9S, Guate., 6000', 12.28.35, inq. spr. '37, 
Q. pilicaulis, Kinsey coll., oxgall of brelan- 
di, Kinsey det. Allotype, <5, with same data 
as holotype. Paratypes: 14 $, 4 3 with same 
data as holotype (AMNH). Excluded from 
type series: 3 specimens from type locality 
in various but poor condition; tentatively 
identified as S. mesoamericanus. 

Etymology.— The specific name means 
"middle America" and refers to this species 

T>'\^gnos\s. — Synergus mesoamericanus 
can be distinguished from all other Guate- 
malan species by the following characters: 
female antenna with 1 4 antennomeres, A3 
1.5 times length of A4; radial cell length 2.0 
times width, Rs2 distinctly bent; female 
metasoma rufus below, male metasoma en- 
tirely black. The radial cell appears to be 
open in many specimens as Rs 1 is often very 
weak along the anterior margin of the wing 
(Fig. 22). 

Remarks.— This species is one of the two 
smaller species of Synergus occurring in 
Guatemala. All the specimens examined 



were reared from galls of ''brelandr and 
emerged during the second spring after col- 

Andricus guatemalensis (Cameron), 
New Combination 

Cynips guatemalensis Cameron, 1883: 71, 
gall. TYPE: Holotype, gall (BMNH). Ex- 

Andricus? mexicana Bassett, 1890: 78 (gall). 

Andricus mexicanus Bassett, Dalla Torre, 

Andricus mexicanusYAnsty , 1920: 308, figs. 
25-27 (9, (5, gall). New Synonymy. 

Synergus dorsalis Cameron, 1883: 72 (not 
'Provancher, 1888: 398). (9, <5). New Syn- 
onymy. TYPE: Holotype, 9 (BMNH). 

Remarks.— Cameron (1 883) described the 
galls of C. guatemalensis which were sent 
to him from Guatemala. Bassett (1890) de- 
scribed the galls oi Andricus? mexicana from 
galls collected in Mexico. Neither Cameron 
nor Bassett described what they thought was 
the gall inducer, although Cameron (1883) 
described Synergus dorsalis, which he 
thought was an inquiline in galls of C gua- 
temalensis (see below). Kinsey (1920) was 
the first to describe the adult gall former, 
which he obtained by cutting from galls that 
had been sent to the American Museum of 
Natural History. Kinsey recognized that the 
galls he had were the same as those of C. 
guatemalensis Cameron and also those of 
Andricus mexicanus Bassett. However, 
Kinsey did not believe that names based on 
only the galls were valid as he described 
Andricus mexicanus as a new species while 
listing C. guatemalensis Cameron 1883 and 
Andricus mexicanus Bassett 1890 as syn- 
onyms. Thus, A. mexicanus Kinsey is a ju- 
nior synonym of both C guatemalensis 
Cameron and A. mexicanus Bassett, and a 
junior homonym of /I. mexicanus Bassett. 
A new name {or Andricus mexicanus Kinsey 
1920 would needlessly burden the already 
confused nomenclature. Both Bassett and 

Kinsey were correct to assign the species to 
Andricus and C. guatemalensis is hereby 
transferred to this genus. 

The holotype of 5". dorsalis (in the British 
Museum (Natural History)) belongs to the 
genus Andricus and is a synonym of Cynips 
guatemalensis Cameron. Although the type 
of guatemalensis is a gall while the type of 
dorsalis Cameron is an insect and it is de- 
sirable to base species of Cynipidae on the 
insect rather than the gall, we retain gua- 
temalensis as the valid name in the interest 
of stability. There are two reasons for doing 
this. First, the name guatemalensis has been 
associated with a gall-inducing species while 
dorsalis Cameron has been associated with 
an inquiline species. Second, retention of 
guatemalensis simplifies a problem of sec- 
ondary homonymy between S. dorsalis 
Cameron 1883 and 5". dorsalis (Provancher) 

When Weld (1951) transferred Ceroptres 
dorsalis Provancher to Synergus, this species 
became a secondary homonym of S. dor- 
salis Cameron. However, Weld did not re- 
name S. dorsalis (Provancher) either be- 
cause he did not know that it was a 
homonym, or because he knew that S. dor- 
salis Cameron should not have been placed 
in Synergus. Weld visited the British Mu- 
seum (Natural History), after which he syn- 
onymized S. filicornis Cameron and S. fur- 
nessana Weld (Weld, 1 930). It is quite likely 
that he also examined the type of 5*. dorsalis 
Cameron as well. A new name for S. dorsalis 
(Provancher) is unnecessary as S. dorsalis 
Cameron is now considered a junior syn- 
onym oi Andricus guatemalensis Cameron 
and therefore the two species are no longer 


We thank the following curators for mak- 
ing types and other material available for 
study: D. Azuma, Philadelphia Academy of 
Natural Sciences, Philadelphia, Pennsyl- 
vania; A. Menke, Systematic Entomology 
Laboratory, Agricultural Research Service, 



USDA, United States National Museum, 
Washington, D.C.; J. Perron, Laval Uni- 
versity, Quebec, Quebec; J. Quinlan, British 
Museum of Natural History, London, 
United Kingdom; and R. T. Schuh, Amer- 
ican Museum of Natural History, New York, 
New York. The scanning electron micro- 
graphs were taken by Lewis Ling (Carleton 
University). This research was supported in 
part by an operating grant from the Natural 
Sciences and Engineering Research Council 
of Canada (No. A0230) awarded to J. D. 

Literature Cited 

Bassett, H. 1890. New species of North American 
Cynipidae. Trans. Entomol. Soc. Am. 17: 59-72. 

Cameron, P. 1883. Hymenoptera. Biologi Centrali- 
Americana, vol. 1, 497 pp., 120 Plates. 

Dalla Torre, K. W. 1893. Cynipidae. Catalogus Hy- 
menoptorum II. 140 pp. 

DallaTorre.K.W.andJ.J. Kieffer. 1910. Cynipidae. 
DasTierreich 24: 1-891. 

Eady, R. D. and J. Quinlan. 1963. Hymenoptera: 
Cynipoidea. Handbooks for the Identification of 
British Insects, VIII (la). Royal Entomological So- 
ciety, London. 86 pp. 

Kinsey, A. C. 1920. New species and synonymy of 

American Cynipidae. Bull. Am. Mus. Nat. Hist. 
42: 293-317. 

. 1 930. The gall wasp genus Cynips. Ind. Univ. 

Studies 84-86: 1-577. 

. 1936. The origin of the higher categories in 

Cvnips. Indiana Univ. Pubs., Sci. Ser. No. 4: 1- 

Provancher, L. 1888. Additions et Corrections au 
Volume II de la Faune de la Province de Quebec. 
Hymenoptera. Family VI. Cynipidae, pp. 397-398. 

Ritchie, A. J. and T. M. Peters. 1981. The external 
morphology of Diplolepis rosae (L.). Ann. Ento- 
mol. Soc. Am. 74: 191-199. 

Shorthouse.J.D.andA. J. Ritchie. 1984. Description 
and biology of a new species of Diplolepis Fourcroy 
(Hymenoptera: Cynipidae) inducing galls on the 
stems of Rosa aciciilahs. Can. Entomol. 1 16: 1623- 

Weld, L. H. 1913. A new oak gall from Mexico. In- 
secutor Inscitiae Menstruus 1: 132-134. 

. 1 930. Notes on types (Hymenoptera: Cynipi- 
dae). Proc. Entomol. Soc. Wash. 32: 137-141. 

. 1951. Cynipoidea. 7^7 Muesebeck, C. F. W., 

K. V. Krombein, and H. K. Townes, eds., Hy- 
menoptera of America north of Mexico, United 
States Department of Agriculture, Monograph 2. 
1420 pp. 

. 1952. Cynipoidea. Ann Arbor, Michigan. 351 


— . 1957. Cynipid Galls of the Pacific Slope. Ann 
Arbor, Michigan. 80 pp. 


89(2), 1987, pp. 242-243 




Curtis W. Sabrosky 

Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % 
U.S. National Museum NHB 168, Washington, D.C. 20560. 

Abstract.— Leptometopa nilssoni Sabrosky, new species, is described from Madagascar. 
Apparently this is the first published record of the genus from that island. 

A new species of the milichiid genus Lep- 
tometopa is described to make the name 
available for L. Anders Nilsson of the In- 
stitute of Systematic Botany, Uppsala Uni- 
versity, Sweden, who investigated the pol- 
lination of Ceropegia albisepta Jum. & H. 
Perr. (Asclepiadaceae) in Madagascar. Pol- 
linaria from this plant were found attached 
to the proboscis of flies entrapped in the trap 
flowers. This paper appears to be the first 
published record of Leptometopa from 

Leptometopa nilssoni Sabrosky, 
New Species 

Polished black species, with black hal- 
teres and a pteropleural (anepimeral) bris- 

Female.— Almost entirely shining black, 
except for white epistomal triangle on lower 
part of face, palpus, proboscis, and stalk of 
halter yellowish to brownish, halter knob 
dull black, mid and hind tarsi yellowish, and 
wing pale with yellowish to whitish-yellow 

Head with frons highly polished but mi- 
croscopically striate; 2 interfrontal rows with 
weak interfrontal hairs in tiny punctures, 
otherwise scarcely evident; cheek polished, 
its height V^ that of an eye and less than 
breadth of third antennal segment, lower 

margin with a row of 5 long setae that in- 
crease slightly in length toward a slightly 
longer and stronger vibrissa; chaetotaxy as 
usual for genus: inner and outer vertical, 
postocellar, and ocellar pairs of bristles, and 
on each side of frons 2 lateroclinate orbital 
(upper) and 2 mesoclinate frontal (lower) 
bristles, all approximately same length and 

Mesoscutum only moderately covered 
with hairs, in about 1 2 rows; scutellum bare; 
mesopleuron (anepistemum) with fine hairs 
on posterior portion; chaetotaxy: 1 humer- 
al, 1 + 1 notopleural, 1 supra-alar, 1 post- 
alar, 1 dorsocentral and 1 pteropleural (ane- 
pimeral) pairs of bristles, and 1 basal and 1 
apical pairs of scutellar bristles. 

Hind tibia broadening from slender base 
to apex, at its widest about three times as 
broad as basal portion, distal half postero- 
dorsally with broad yellowish tibial organ. 

Wing as usual for genus, with two costal 
breaks, costa extending to 4th vein(M) 
slightly beyond apex of wing, crossveins well 
separated, and anal vein not extended be- 
yond anal cell; vein 4(M) converging slightly 
toward preceding vein (R 4 + 5), narrowing 
cell toward apical margin of wing and mak- 
ing the 4th section of costa slightly shorter 
than the 3rd. 

Length.— 1.25-1.5 mm. 



Holotype and two paratypes. — All fe- 
males, Madagascar Centre; Mandraka, April 
6, 1985 (L. A. Nilsson), in flowers of Cer- 
opegia albisepta. Type series deposited in 
the U.S. National Museum of Natural His- 
tory, by courtesy of the collector. 

The polished black head and body of L. 
nilssoni, so unlike the densely gray micro- 
tomentose type species, Leptometopa la- 
tipes (Meigen), at first glance suggest a species 
of the genus Madiza. However, there are 
polished black species in the genus, notably 
the Nearctic Leptometopa halteralis (Co- 
quillett). Moreover, some intermediate 
species neatly bridge the gap between pol- 
ished and microtomentose species. Lepto- 
metopa beardsleyi Hardy and Delfinado 
from the Hawaiian Islands has a polished 
black head but gray microtomentose thorax, 
although the latter is not as densely gray as 
in latipes and therefore appears darker. Also, 
Leptometopa albipemiis (Lamb) from the 
Seychelles has the head and most of the tho- 
rax polished black but the scutellum is 

brownish microtomentose. The latter com- 
bination of characters is also found in an 
undescribed species from New South Wales, 
Australia, but its relationship to albipennis 
has not been studied. Incidentally, L. albi- 
pemiis shows other differences from nils- 
soni, such as having the frons reddish an- 
teriorly, knob of halter whitish yellow, and 
wing very milky white. 

Males are not available, unfortunately, 
because the peculiarly broadened hind tibia 
in that sex is peculiar to the genus Lepto- 
metopa. However, the head of the female is 
characteristic of Leptometopa, with the lu- 
nule extending ventrad as a wedge-shaped 
divider between deeply-set antennae, its 
apex nearly touching the apex of the trian- 
gular epistomal area of the lower face. Fur- 
thermore, a pteropleural (anepimeral) bris- 
tle is present, a common, although not uni- 
versal, feature of Leptometopa. 

The specific name, a noun in the genitive 
case, is dedicated to L. Anders Nilsson, the 
botanist who collected the specimens. 


89(2), 1987, pp. 244-249 


A. G. Wheeler, Jr. and Jonathan E. Fetter 

Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn- 
sylvania 17110. 

Abstract. — Chilacis typhae (Perris), an Old World lygaeid restricted to cattails (Typha 
spp.), is reported new to the Western Hemisphere based on collections from Delaware (1 
county), Maryland (3), New York (3), and Pennsylvania (25). The known Nearctic dis- 
tribution is mapped, notes on seasonal history and habits in Pennsylvania are given, and 
morphological characters facilitating recognition of this immigrant heteropteran are pro- 

Chilacis typhae (Perris) is an Old World 
artheneine lygaeid occurring throughout 
England but uncommon in Wales and Ire- 
land (Southwood and Leston, 1 959). On the 
Continent it ranges from France east to Bul- 
garia, Rumania, and the European U.S.S.R., 
including Caucasus to the south, and north 
to Scandinavia; in Asia it has been recorded 
from Jordan (Slater, 1964). As the specific 
name implies, C typhae is a specialist on 
plants of the genus Typha (Typhaceae), par- 
ticularly common cattail, T. latifolia L. 
(called reedmace or bulrush in Britain). Jor- 
dan ( 1 935) and Cobben ( 1 95 3) discussed the 
habits of this seed-feeding bug; see Slater 
(1964) for additional references to papers 
containing biological information on C 0^- 

The discovery of this lygaeid in Pennsyl- 
vania and its subsequent collection in Del- 
aware, Maryland, and New York represent 
the first Nearctic records for the species. The 
only other artheneine known from the New 
World is Polychismeferruginosus (St^l). This 
Neotropical bug, recorded from Colombia 
and Venezuela, was recently removed from 
the Ischnorhynchinae and transferred to the 
Artheneinae, thus providing the first West- 

em Hemisphere record and making the 
subfamily known from all major zoogeo- 
graphic areas except the Nearctic (Slater and 
Brailovsky, 1986). 

Here, we give collection data and map the 
known North American distribution of C 
typhae\ give notes on its seasonal history, 
relative abundance, and habits on cattail in 
Pennsylvania; and list adult and nymphal 
characters that allow recognition of this ad- 
ventive lygaeid in the Nearctic fauna. 

North American distribution.— The first 
known collection of C typhae in the New 
World was made on 4 June 1986 (by JEF) 
near Lickdale, Pennsylvania. Further col- 
lecting showed that this species is well es- 
tablished in eastern Pennsylvania (but ap- 
parently scarce in the western counties) and 
present in southern and central New York 
and northern Delaware and Maryland. We 
did not encounter C. typhae during limited 
collecting in Ohio, western New York, and 
in more southern areas of Maryland (Fig. 
1), or in Indiana or Virginia (negative sites 
not shown in Fig. 1). The following records 
are available for C typhae in eastern United 
States. All collections were made in 1986 
from Typha latifolia; voucher specimens 



Fig. 1 . Map of the Middle Atlantic and New England States showing known distribution of Chilacis typhae 
in North America. Closed circles represent 1986 collection sites; open circles, sites where cattail heads did not 
yield C. typhae. 

have been deposited in the insect collections 
of Cornell University, Pennsylvania De- 
partment of Agriculture, U.S. National Mu- 
seum of Natural History, and University of 

DELAWARE: New Castle Co., e. of New- 
ark, 5 Nov., F. G. Steams. MARYLAND: 
Allegany Co., nr. Flintstone, 30 July, A. G. 
Wheeler, Jr.; Frederick Co., Emmitsburg, 
18 June, AGW; Washington Co., nr. Han- 
cock, 30 July, AGW. NEW YORK: Broome 
Co., s. of Kirkwood and nr. Castle Creek, 
26 June, AGW; Tioga Co., e. of Caroline 
Center, 26 June, AGW; Tompkins Co., Be- 
semer, 26 June and s. of Ithaca, 29 June, 
AGW; Town of Ulysses, 9 mi. n. of Ithaca, 
7 July, E. R. Hoebeke. PENNSYLVANIA: 
Adams Co., s. of East Berlin, 16 June, AGW 

and nr. Gettysburg and Hunterstown, 18 
June, AGW; Berks Co., nr. Frystown and 
Rehrersburg, 1 7 June, AGW and nr. Ham- 
burg, 24 June, J. F. Stimmel; Bradford Co., 
Towanda, 7 Aug. AGW and n. of Stevens- 
ville, 7 Aug., R. J. Gallagher; Bucks Co., s. 
of Quakertown, 24 June, J. F. Stimmel; Car- 
bon Co., Nesquehoning, 3 July, AGW; 
Centre Co., 1-80 e. of Clearfield Co. line nr. 
Lanse, 10 June, AGW and nr. Martha Fur- 
nace, 20 July, JEF; Chester Co., Longwood 
Gardens, 5 Nov., F. G. Steams; Columbia 
Co., e. of Bloomsburg, 5 Aug. and 29 Oct., 
AGW; Cumberland Co., nr. Wertzville, 16 
June, AGW; Dauphin Co., nr. Harrisburg 
Area Community College, 9 June, AGW and 
Dauphin, 12 June, JEF; Delaware Co., Me- 
dia, 5 Nov., F. G. Steams; Juniata Co., nr. 



Port Royal, 12 June, JEF; Lackawanna Co., 
nr. Clarks Summit, 24 June, R. J. Gallagher 
and e. of Factoryville, 26 June, AGW; Leba- 
non Co., 1-81 at Exit 30, w. of Lickdale, 4 
June and Rts. 22 and 934, nr. Harper Tav- 
ern, 1 1 June, JEF; e. of Fredericksburg, 1 7 
June, AGW; Lehigh Co., e. of Fogelsville, 
3 July, AGW; Luzerne Co., Frances Slocum 
St. Pk., 24 June, R. J. Gallagher and 1-80 
at June. Rt. 309, nr. Freeland, 26 June, 
AGW; Monroe Co., Sciota, 3 July, AGW; 
Montour Co., n. of Danville, 5 Aug., AGW; 
Schuylkill Co., 1-81 at junc. Rt. 125, nr. 
Ravine, 6 June and Barnesville, 3 July, 
AGW; Susquehanna Co., Great Bend, 26 
June, AGW; Washington Co., nr. Bulger, 

17 June, L. L. Garrett; Wayne Co., Beach 
Lake and s. of Hoadleys, 1 9 June, K. Valley; 
Westmoreland Co., Delmont, 2 Dec, L. L. 
Garrett; Wyoming Co., Overfield Twp. n.e. 
of L. Winola, 24 June, R. J. Gallagher and 
e. of Dixon, 26 June, AGW; York Co., s.w. 
of Kralltown, 1 6 June and s. of Dillsburg, 

18 June, AGW. 

Chilacis typhae appears to be a relatively 
recent introduction in North America, even 
though its principal host, Typha latifolia, is 
naturally Holarctic (Gleason and Cronquist, 
1963; Everett, 1982). The tribe to which C 
typhae belongs, Artheneini, is considered a 
Palearctic element (Slater, 1964; Slater and 
Brailovsky, 1986). Claassen (1921) collect- 
ed the lygaeid Kleidocerys resedae (Panzer) 
during his work on cattail insects, and it is 
unlikely he would have overlooked C. ty- 
phae in his thorough survey of Typha, which 
was conducted mainly in the Ithaca, New 
York area where this bug is now established. 

Biological notes.— We observed C. ty- 
phae in the field and brought back cattail 
heads to the laboratory so that nymphs and 
adults could be extracted with a Berlese fun- 
nel. In early June we found adults and 
nymphs of all stages in heads of the previous 
season. The bugs were restricted to old heads 
fluffed out by the feeding of the cosmopte- 
rygid Limnaeciaphragmitella Stainton (Fig. 
2) (see Claassen, 1921). Four or five heads 

collected in Schuylkill Co. on 6 June yielded 
29 first, 27 second, 35 third, 67 fourth, and 
28 fifth instars. Some old heads harbored 
several hundred adults and nymphs; cast 
skins often were conspicuous on heavily in- 
fested heads. 

The ischnorhynchine K. resedae also was 
present in old heads taken in several cattail 
colonies and occasionally occurred in the 
same head with C. typhae. Three such heads 
contained 200 individuals of C. typhae and 
107 of A^. resedae. The latter species feeds 
on seeds of numerous plants, particularly 
birch, rhododendron and other ericaceous 
shrubs, and cattail (see Wheeler, 1976). The 
interaction and possible competition be- 
tween these lygaeids of similar feeding hab- 
its (Sweet, 1960), although difficult to as- 
sess, seem worthy of study. 

By mid-July to early August fewer old 
heads contained large numbers of adults; 
nymphs, nearly all late instars, became 
scarce. During this time adults were ob- 
served in newly formed staminate spikes 
where they usually were concealed beneath 
the stamens. The staminate or upper por- 
tion of the fruiting head is not persistent, 
the stamens soon being shed, and adults are 
more likely to be detected on the new pis- 
tillate spikes. Mating occurred on the sur- 
face, the pairs typically clustered near white 
fluffy patches that mark tunnels of L. phrag- 
mitella (Fig. 3). Adults push their stylets 
through the dense flowers; during feeding 
the body is elevated and the abdomen held 
nearly perpendicular to the surface, as noted 
by Cobben (1953). The bugs also use lepi- 
dopteran tunnels to burrow into the new 
fruiting spikes, which are composed of nu- 
merous, dense minute flowers. Thus, larvae 
of the Holarctic moth L. phragmitella are 
important to C typhae for permitting access 
to the interior of hard newly formed heads, 
and also in spinning silk to hold the downy 
material together so that seeds are retained; 
uninfested heads lose their seeds during 
winter and spring (Claassen, 1921). Other 
Lepidoptera are known to develop in heads 



Figs. 2, 3. Pistillate spikes of Typha latifolia. 2, Head of previous season fluffed out by the tunneling and 
feeding of larvae of the cosmopterygid Limnaecia phragmitella. 3, Current-season head with white, fluffly patches 
indicating new larval tunnels of L. phragmitella. 

of T. latifolia in North America, but L. 
phragmitella "is the most common and the 
most abundant of the insects infesting the 
cat-tail" (Claassen, 1921). One even won- 
ders whether C typhae could have become 
established in North America in the absence 
of the cosmopterygid. 

Chilacis typhae oviposits on the seed or 
pappus (Southwood and Leston, 1959). In 
early August we collected large numbers of 
nymphs in pistillate heads, mainly early in- 
stars of a presumed second generation. The 
heads contained adults, some of them tener- 
al, or adults and fourth and fifth instars in 
late October and early November; adults 
only were present at one site sampled in 
early December. 

Because we did not sample a cattail col- 

ony at regular intervals throughout a season, 
our phenological profile of C. typhae in east- 
em North America is tentative. In England 
adults are known to overwinter, oviposition 
begins in late May or early June, and nymphs 
become adult by mid-July. There is a single 
annual generation, although in Central Eu- 
rope a partial second generation may be 
produced in autumn, the nymphs overwin- 
tering (Southwood and Leston, 1959). The 
adults collected during early June in Penn- 
sylvania may have been a combination of 
overwintered individuals and those of the 
season's first brood since fifth instars were 
present. Collection of all nymphal stages, 
especially early instars, from new pistillate 
spikes in late July-early August suggests the 
beginning of a second generation; and the 



^^^^^^^^^^^^H^^^^^' ' ^'^3^^^^^^^^^^^^^^^^^ 


Fig. 4. Adult habitus of Chilacis typhae, dorsal aspect. Scale line = 1 .0 mm. 

presence of teneral adults and late instars in 
late October, the completion of a second 

This presumed bivoltinism, with adults 
overwintering and occurring throughout the 
summer, contrasts with the life cycle Jordan 
(1935) outlined for C. typhae in Germany. 
He emphasized that because this lygaeid de- 
pends on fruit-bearing stands of cattail, its 

development takes place during the cold 
season. Copulation and some development 
occur in winter, and all nymphs become 
adult the following May. Only adults are 
found during summer when they are said to 
be inactive beneath cattails or rest on other 
marsh plants. In the Netherlands, however, 
Cobben (1953) reported that adults are 
abundant and mate during July and August. 



Additional study is needed to determine 
whether North American populations are 
bivoltine and whether only adults survive 
the winter. 

Recognition features.— The adult of C. 
typhae (Fig. 4) is 3.0 to 4.8 mm long, gla- 
brous, shining yellowish brown or ochreous 
and strongly punctured above, with the head 
and thorax fuscous beneath. It is easily rec- 
ognized by the juga that nearly attain the 
apex of the tylus, the latter extending be- 
yond the apex of antennal segment I; the 
prominent fuscous grooves separating tylus 
from juga; the nearly trapezoidal pronotum 
with sublaminately produced edges; and the 
dark scutellum with a pale V-shaped mark. 

Nymphs are a pale yellowish brown with 
orangeish or reddish markings (or some- 
times transverse reddish bands) on the ab- 
domen and have dorsal abdominal scent 
glands that open between tergites III-IV, 
IV-V, and V-VI; the anterior gland is re- 
duced (Usinger, 1938). Peneau (1909) and 
Collett (1927) illustrated the fifth instar of 
C typhae, and Jordan (1935) described four 
of the five nymphal stages; his (and Col- 
lett's) instar IV is actually V (Puchkov, 
1958). For references to additional descrip- 
tions and illustrations of the immature stages 
of C typhae the reader is referred to Slater's 
( 1 964) catalog. The nymph of A', resedae can 
be distinguished from that of C. typhae by 
the brown or almost fuscous head and 
pronotum and dark red or reddish-brown 
abdomen, and by the well-developed an- 
terior scent gland. 


We are grateful to J. A. Slater (University 
of Connecticut) for confirming the deter- 
mination of C. typhae; R. J. Gallagher, L. 
L. Garrett, R. J. Henry, F. G. Steams, J. F. 
Stimmel, and K. Valley (PDA, BPI) and E. 
R. Hoebeke (Cornell University) for helping 

collect cattail heads; and Slater, Valley, and 
Hoebeke for commenting on the manu- 

Literature Cited 

Claassen, P. W. 1921. Typha insects: Their ecological 

relationships. Cornell Univ. Agric. Exp. Stn. Mem. 

47. pp. 459-531. 
Cobben, R. H. 1953. Bemerkungen zur Lebensweise 

einiger hollandischen Wanzen (Hemiptera-Het- 

eroptera). Tijdschr. Entomol. 96: 169-198. 
Collett, H. R. P. 1927. The earlier stages of Chilacis 

typhae Perr. Entomol. Mon. Mag. 63: 155-157. 
Everett, T. H. 1 982. The New York Botanical Garden 

illustrated encyclopedia of horticulture. Vol. 10, 

Ste-Zy. Garland Pubhshing. New York. pp. 3225- 

Gleason, H. A. and A. Cronquist. 1963. Manual of 

Vascular Plants of Northeastern United States and 

Adjacent Canada. D. Van Nostrand, Princeton, 

N.J. 810 pp. 
Jordan, K. H. C. 1935. Beitrag zur Lebensweise der 

Wanzen auf feuchten Boden. (Heteropt.). Stett. 

Entomol. Ztg. 96: 1-26. 
Peneau, J. 1909. Notules hemipterologiques (3). I.— 

Sur la capture de Chilacis Typhae (Penis). Bull. 

Soc. Sci. Nat. Quest France (2) 9: 511-514. 
Puchkov. v. G. 1958. Larvae of Hemiptera-Heter- 

optera. I. Lygaeidae. Entomol. Rev. 37(2): 332- 

Slater, J. A. 1964. A Catalogue of the Lygaeidae of 

the World. 2 vols. Univ. of Connecticut. Storrs. 

1668 pp. 
Slater, J. A. and H. Brailovsky. 1986. The first oc- 
currence of the subfamily Artheneinae in the 

Western Hemisphere with the description of a new 

tribe (Hemiptera: Lygaeidae). J. N.Y. Entomol. 

Soc. 94:409-415. 
Southwood, T. R. E. and D. Leston. 1959. Land and 

Water Bugs of the British Isles. Frederick Wame 

&. Co., London. 436 pp. 
Sweet, M. H. 1960. The seed bugs: A contribution 

to the feeding habits of the Lygaeidae (Hemiptera: 

Heteroptera). Ann. Entomol. Soc. Am. 53: 317- 

Usinger, R. L. 1938. Dorsal abdominal scent glands 

in nymphs of Lygaeidae. Pan-Pac. Entomol. 14: 

Wheeler, A. G., Jr. 1976. Life history of Kleidocerys 

resedae on European white birch and ericaceous 

shrubs. Ann. Entomol. Soc. Am. 69: 459-463. 

89(2), 1987, pp. 250-263 


Donald W. Webb 

Section of Faunistic Surveys and Insect Identification, Illinois Natural History Survey, 
Champaign, Illinois 61820. 

Abstract.— Tht genus Arthroceras Williston contains seven species and is Holarctic in 
distribution. This paper revises the four species recorded by Nagatomi (1966) for the 
Nearctic Region. Of these four species, one species and two subspecies are relegated to 
synonymy. The immature stages of this genus are unknown and little is known of its 
biology. Keys to species, with descriptions and distributions of each Nearctic species, are 

The genus Arthroceras was erected by 
Williston (1886) and currently contains sev- 
en species (Nagatomi, 1966). It is Holarctic 
in its distribution, although it does not oc- 
cur in Europe. This paper revises the four 
species recorded by Nagatomi ( 1 966) for the 
Nearctic Region. Of these four species, one 
species and two subspecies are relegated to 
synonymy. The immature stages of the ge- 
nus are unknown and little is known of its 

The terminology used here follows 
Mc Alpine (1981) and for the male termi- 
nalia, Stuckenberg (1973). The range for each 
measurement or ratio is followed by the av- 

Arthroceras Williston 

Arthroceras Williston (1886: 107): Coquil- 
lett (19 10: 5 10); James (1965: 298); James 
and Turner (1981: 486); Leonard (1930: 
52); Nagatomi (1966: 44, 1970:293, 1982: 
142, 1984: 142); Nagatomi and Iwata 
(1976: 20). Type-species: A. pollinosum 
Williston by original description. 

Pseudocoenomyia Ouchi (1943: 493): Na- 
gatomi (1955: 57, 1966:44). Type-species: 
P. sinensis Ouchi. 

Ussuriella Paramonov (1929: 181): Naga- 
tomi (1970: 293). Type-species: U. gadi 

Head in lateral view hemispherical. Ver- 
tex truncate, not emarginate lateral to ocel- 
lar tubercle. Ocelli dark red to black; ocellar 
tubercle subtriangular, distinctly raised 
above vertex in male, not raised above ver- 
tex in female. Eyes holoptic in male, dich- 
optic in female; facets in male smaller on 
ventral third, equal in female; setae absent 
or very short, scattered; median margin sin- 
uate, ventral half divergent; in lateral view 
eyes hemispherical to subtriangular. Frons 
reduced in male, broad in female, divergent 
dorsally; setae generally absent in male, 
scattered in female. Antenna (Figs. 1-6) 
shorter than length of head; scape in both 
sexes separated by distance greater than 
width of median ocellus, short, length less 
than width, shorter than pedicel, setae ab- 
sent; pedicel short, globose, shorter than 
wide, setae shorter than length of segment; 
flagellum subulate, longer than combined 
length of scape and pedicel, annulate, setae 
scattered. Gena broad; parafacial setae ab- 
sent. Clypeus broad, anterior surface dis- 



tinctly convex, forming deep lateral grooves; 
setae scattered. Maxillary palpus 2 seg- 
mented; basal segment cylindrical, longer 
than wide; apical segment cylindrical, curved 
ventrally, apex rounded, longer than wide, 
longer than basal segment. 

Thorax with dorsum rounded; vitta gen- 
erally indistinct; setae elongate, scattered, 
in no distinctive pattern. Postpronotal lobe 
concolorous with thorax. Postmetaspiracu- 
lar scale and suprametacoxal pit lacking. 
Scutellum with caudal margin broadly 
rounded. Laterotergite of postnotum with 
elongate setae. 

Wing (Fig. 14) longer than wide; mem- 
brane opaque; pterostigma variable; micro- 
setae minute, over entire membrane; setu- 
late dorsal on length of R,; thyridium absent. 
Costa circumambient, broader along ante- 
rior margin, setae fuscous, appressed. Hu- 
meral crossvein distinct. Subcosta ends dis- 
tad to middle of wing. Subcostal and 
marginal cells elongate, open. Radial sector 
originates from basal fourth of first basal 
cell, r-m situated above basal third of discal 
cell. R, ends distad to fork of R4+5 and apex 
of discal cell. Fork of R4+5 originates basad 
to apex of R|, distad to apex of discal cell, 
angle variable. Cell r4 elongate, enclosing 
apex of wing. R4 ends anterior to apex of 
wing. R5 ends posterior to apex of wing. M, 
and M2 petiolate, contiguous or separate 
from discal cell. Cell m, open. M, and CuA, 
parallel. Posterior cells 5. Discal cell elon- 
gate, apical margin truncate. Posterior cubi- 
tal cell open. First basal cell elongate, apex 
truncate, ends distad to apex of second basal 
cell. Second basal cell emits 4 veins from 
apex. Anal lobe broadly rounded, right-an- 
gled. Alula rounded. Squama large; margin- 
al setae elongate, entire. 

Legs with tibial spurs 0-2-1. Hind coxa 
with distinct anterior tubercle. Empodium 
pulvilliform. Apical claws on tarsomere 5 
fuscous, paired, simple. Hind legs not rap- 

Abdomen with tergite 1 subrectangular, 
anterior margin broadly emarginate. Male 

terminalia with tergite 8 broad, rectangular, 
length 1.5 times width, caudal margin trun- 
cate. Epandrium (Figs. 7, 15) with lateral 
and caudal margins rounded, anterior mar- 
gin broadly emarginate, length 1.5 times 
width. Tergite 10 absent. Cerci simple, flat- 
tened dorsoventrally. Ventral plate of proc- 
tiger subtriangular. Gonocoxite in ventral 
view (Figs. 8, 16) broad, lateral margins 
rounded, fused anteriorly with oval scler- 
otized median plate; in dorsal view (Figs. 
9, 17) gonocoxite with narrow caudal arch 
joining inner margins, aedeagal apodeme 
elongate, extending anteriorly to anterior 
margin of gonocoxite. Gonostylus narrow, 
tapered apically, reflexed. Aedeagus com- 
posed of aedeagal sheath, endophallus, and 
endophallic hilts. Aedeagal sheath broad 
basally, fused to inner margin of gonocoxite, 
tapered caudally to form narrow endophal- 
lic guide. Endophallus elongate anteriorly 
forming endophallic apodeme, caudal third 
oval. Endophallic hilts separated medially, 
thick, heavily sclerotized, tapered anterior- 
ly. Penis valves and endophallic tines lack- 
ing. Female terminalia with tergite 9 (Fig. 
1 9) quadrate, as long as wide, caudal margin 
truncate. Tergite 1 reduced, caudal margin 
truncate to broadly rounded. Cerci 2 seg- 
mented; basal segment broad, equal in length 
to apical segment; apical segment clavate 
with apical depression. Stemite 8 (Fig. 20) 
broad, widened caudally, 1.3 times longer 
than wide, caudal margin rounded with 
deep, median emargination. Sternite 9 
greatly modified, invaginated beneath ster- 
nite 8 to form internal furca. Stemite 10 
broad, membranous, caudal margin sinuate 
with median groove. Internal reproductive 
organs (Figs. 11,21) with furca "Y" shaped, 
anterior apodeme short, not attached lat- 
erally to tergite 9. Common spermathecal 
duct short, trifurcating anteriorly to form 3 
spermathecal ducts. Each spermathecal duct 
membranous, elongate, ending anteriorly in 
dark brown, spherical spermatheca (Figs. 1 1 , 
Immature stages unknown. 



Key TO THE Nearctic 
Species of Arthroceras 

1. Length of apical flagellomere greater than 1.5 
times width of flagellum (Fig. 6); setae on hind 
femur pale yellow, elongate, suberect, length 
greater than 0.5 times greatest width of femur 
pollinosum Williston 

- Length of apical flagellomere less than 1.5 times 
width of flagellum (Figs. 1-5); setae on hind 
femur predominately fuscous, short, ap- 
pressed, length much less than 0.5 times great- 
est width of femur 2 

2. In males, maxillary palpus pale to dark yellow, 
setae predominately stramineous, apical seg- 
ment at least 1.5 times length of basal segment. 
In females, head, antenna, maxillary palpus, 
thorax, femora and abdomen yellow pollinose 

to dark yellow fuhicorne Nagatomi 

- In males, maxillary palpus brown, setae black, 
apical segment about 1.2 times length of basal 
segment. In females, head, antenna, maxillary 
palpus, thorax, femora, and abdomen brown 

to dark brown leptis (Osten Sacken) 

Arthroceras fulvicorne Nagatomi 

Arthroceras fuhicorne ^di%3.Xom.\{\9 66: 46). 

Arthroceras fulvicorne nigricapite Nagatomi 
(1966: 48). New Synonymy. 

Arthroceras fulvicorne subsolanum Naga- 
tomi (1966: 49). New Synonymy. 

Arthroceras subaquilum Nagatomi (1966: 
59). New Synonymy. 

Arthroceras fulvicorne is separated from 
A. pollinosum by having the length of the 
apical flagellomere less than 1.5 times as 
wide as the flagellum and by having the se- 
tae on the hind femur predominately fus- 
cous, short, and appressed. Arthroceras ful- 
vicorne and A. leptis can be separated by the 
characters given in the key. 

Nagatomi (1 966) separated the females of 
A. fulvicorne 2Lnd. A. subaquilum on the basis 
of the coloration of the antennal flagellum 
and the abdomen. At that time he examined 
four specimens of A. subaquilum and 3 1 
females of ^. fulvicorne. In the 57 females 
of ^. fulvicorne from northern Utah that I 
examined, the antennal flagellum varied 
from brown to black; the abdominal tergites 
were generally dark yellow and concolorous, 
but variation ranged from tergites 2-4 being 

dark brown with the caudal margins dark 
yellowish brown to the entire abdomen being 
dark brown. On the basis of this variation, 
I have synonymized A. subaquilum with A. 
fulvicorne. Nagatomi (1966) separated A. 
fulvicorne fulvicorne and A. fulvicorne ni- 
gricapite on the coloration of the female 
head. In female specimens from California 
and Utah, the ground coloration of the head 
varied from fuscous to black. Because of this 
variation, I have synonyrmztd A. fulvicorne 
nigricapite with A. fulvicorne. Nagatomi 
(1966) separated A. fulvicorne subsolanum 
and A. fulvicorne fulvicorne on the basis of 
the antennal flagellum being 7 or 8 seg- 
mented in A. fulvicorne fulvicorne and 5 or 
6 segmented in A. fulvicorne subsolanum. 
Table 1 shows the variation in the number 
of completely separated flagellomeres (fla- 
gellum cleared in 10% KOH). Because of 
the variation in flagellum segmentation in 
specimens examined from Nova Scotia to 
British Columbia and California, I have 
synonymized A. fulvicorne subsolanum with 
A. fulvicorne. 

Male.-Length 6.2-8.2, 7.2 mm. Head 
with ocellar tubercle fuscous to black, prui- 
nosity light yellow grey; setae fuscous to 

Table 1 . Segmentation and fusion in the female fla- 
gellum oi Arthroceras fulvicorne. Flagella were cleared 
in 10% KOH. 


Fusion Pattern of 



Nova Scotia 

1+2,3 + 4 + 5,6,7 




1+2,3 + 4,5,6,7 










1,2,3,4,5,6,7 + 8 




British Columbia 

1+2,3 + 4,5,6,7 


1,2,3,4,5,6,7 + 8 






1,2,3,4,5,6,7 + 8 


1+2,3 + 4.5,6,7 




black, elongate, abundant. Eyes dark brown 
to black. Frons fuscous in ground color with 
dense silver pile. Antenna length 0.6-0.8, 
0.7 times length of head; scape fuscous to 
dark brown, pruinosity light yellow, length 
0.4-0.6, 0.5 times width, 0.5-1.0, 0.8 times 
length of pedicel; pedicel fuscous to dark 
brown, pruinosity light yellow, length 0.4- 
0.8, 0.6 times width, setae fuscous, scat- 
tered; flagellum fuscous to brown, pruinos- 
ity light yellow, basal flagellomere (1, 2, or 
1 + 2) yellowish orange to dark yellow, length 
3.0-4.1, 3.5 times width, 2.7-3.9, 3.3 times 
combined length of scape and pedicel, setae 
silver and fuscous, variable; annuli 7, apical 
flagellomere length 0.9-1.3, 1.1 times width 
of flagellum. Gena pruinosity dense silver 
grey; facial setae white to pale yellow, elon- 
gate, abundant. Clypeus fuscous in ground 
color, pruinosity dense silver grey; setae 
white to pale yellow, elongate, scattered on 
dorsal third. Maxillary palpus with setae pale 
yellow, elongate, abundant, occasionally 
with scattered black setae; basal segment 
generally pale to dark yellow, occasionally 
brown, length 1.8-2.2, 2.0 times width; api- 
cal segment pale to dark yellow, length 2.7- 
3.3, 3.0 times width, 1.3-1.8, 1.5 times 
length of basal segment. Labellum fuscous, 
pruinosity light yellow; setae fuscous, mod- 
erately long, scattered. Postocular setae 
white to pale yellow, elongate, abundant, 
becoming shorter dorsally. 

Thorax fuscous to black in ground color, 
pruinosity light grey to yellowish grey; setae 
yellow, elongate, scattered; vitta indistinct, 
occasionally pale fuscous. Postpronotal lobe 
with setae yellow, elongate, abundant. Pleu- 
ron fuscous in ground color, pruinosity vari- 
able in density; setae pale yellow, elongate, 
abundant on propleuron, scattered on dor- 
sal two-thirds to half of anepistemum, scat- 
tered ventrally and dorsally on katepister- 
num, absent on anepimeron, scattered on 
ventral third and in caudolateral area of 
meron, scattered along caudal margin of 
metapleuron. Halter stalk dark yellow to 
dark yellowish brown, capitulum fuscous to 

dark brown, pruinosity light grey; setae fus- 
cous, short, scattered. Scutellum fuscous, 
pruinosity yellowish grey; setae pale yellow, 
elongate, abundant. Postnotum fuscous, 
pruinosity grey; laterotergite fuscous to dark 
brown, pruinosity light yellowish grey; setae 
pale yellow, elongate, abundant. 

Wing as in A. poUinosum (Fig. 1 4). Length 
6.5-7.5, 7.0 mm, width 2.3-2.8, 2.6 mm, 
length 2.7-3.0, 2.8 times width. Membrane 
pale brown; veins yellowish brown; ptero- 
stigma slightly darker than membrane. Fork 
of R4+5 almost right angled. M,, M., M3 orig- 
inate separately from discal cell. 

Coxae fuscous, pruinosity grey, setae on 
forecoxa pale yellow, elongate, abundant; 
femora pale to dark yellow, glossy; tibiae 
variable from fuscous to dark yellowish 
brown, becoming fuscous apically; tarsi fus- 
cous. Empodia and pulvilli fuscous to dark 

Abdomen fuscous in ground color, prui- 
nosity yellowish grey to brownish grey; setae 
fuscous, moderately long, scattered medi- 
ally, with white to pale yellow setae laterally 
on tergites 1 and 2. Terminalia (Figs. 7-10). 
Gonocoxite in ventral view with mediolat- 
eral enlargement on caudolateral projec- 

Female. — Length 7.5-1 1.4, 9.4 mm. Head 
with ocellar tubercle dark yellow to black 
in ground color, pruinosity grey to yellowish 
grey; setae pale to dark yellow, elongate, 
scattered. Eyes dark red to fuscous. Frons 
variable, pruinosity greenish grey pollinose; 
setae white to dark yellow, short to elongate, 
scattered. Antenna (Figs. 1-4), length 0.7- 
0.9, 0.8 times length of head; scape dark 
yellow, pruinosity light yellow grey, length 
0.7-0.9, 0.8 times length of head; scape dark 
yellow, pruinosity light yellow grey, length 
0.5-0.7, 0.6 times width, length 0.7-1 .0, 0.9 
times length of pedicel; pedicel dark yellow, 
pruinosity light yellow grey, length 0.5-0.8, 
0.6 times width, setae pale to fuscous, mod- 
erately long; flagellum brown to black, basal 
half of flagellomere 1 or 1+2 dark yellow, 
length 3.2-4.0, 3.7 times width, 2.5-3.8, 3. 1 



Sea Ped 



Figs. 1-6. Female antenna, lateral view. 1-4, Arthroceras fulvicome. 1, New Hampshire. 2, Utah. 3, Utah. 
4, California. 5, Arthroceras leptis. 6, Arthroceras pollinosum. Abbreviations: (Flag) Flagellum, flagellomeres 1- 
8. (Ped) Pedicel. (Sea) Scape. 




Figs. 7-11. Arthroceras fuhicome. 7, Male epandrium, cerci, ventral plate of proctiger. 8, Male gonocoxite, 
ventral view. 9, Male gonocoxite, dorsal view. 10, Male endophallus, lateral view. 11, Female furca and sper- 
matheca. Abbreviations: (AA) Aedeagal apodeme. (C) Cerci. (E) Endophallus. (EH) Endophallic hilt. (Epa) 
Epandrium. (Gs) Gonostylus. (Gx) Gonocoxite. (VPP) Ventral plate of proctiger. 



Fig. 12. Arthroceras fulvicome, distribution. 

times combined length of scape and pedicel, 
setae on flagellomere 1 or 1+2 fuscous, 
short, appressed, on apical flagellomeres se- 
tae silver to pale yellow, very short, ap- 
pressed; apical flagellomere length 1.0-1.2, 
1.1 times width of flagellum. Gena polli- 
nose, occasionally pruinosity dense grey; fa- 
cial setae white to pale yellow, elongate, 
abundant. Clypeus pollinose, occasionally 
pruinosity dense grey; setae white to pale 
yellow, elongate, scattered. Maxillary pal- 
pus dark yellow, pollinose, setae pale yel- 
low; basal segment length 1 . 1-2.2, 1 .8 times 
width; apical segment length 3.1-4.2, 3.6 
times width, 1.7-2.8, 2.1 times length of 
basal segment. Postocular setae pale yellow 
to golden, elongate, abundant, becoming 
shorter dorsally but reaching vertex. 

Thorax variable, pollinose, yellowish grey, 
or greenish yellow-grey; vitta variable, in- 
distinct, to faintly dark brown, to grey 
brown. Pleuron pale to dark yellow, prui- 
nosity light yellow grey; setae as in male. 
Halter pale to dark yellow brown. Scutellum 
dark yellow, pruinosity grey. Postnotum 
variable, dark yellow, occasionally with 
ventral fourth fuscous; laterotergite dark 
yellow, pruinosity grey. 

Wing length 8.0-10.5, 9.1 mm, width 2.5- 
3.8, 3. 1 mm, length 2.8-3.4, 3.0 times width. 
Membrane pale yellow to pale yellow brown. 
Cell r4 more elongate than in male. 

Coxae yellow to dark yellow, pruinosity 
light grey; femora dark yellow to brown; 
tarsi fuscous to black. 

Color pattern of abdomen highly vari- 
able, generally dark yellow, subshiny but in 
many specimens tergites 2-4 are dark brown, 
pruinosity grey, caudal margin dark yellow- 
ish brown, tergite 5 dark yellow, and oc- 
casionally the entire abdomen is dark brown; 
setae as in male. Terminalia as in A. polli- 
nosum (Figs. 1 9-20). Internal reproductive 
organs (Fig. 11). Furca length 0.40 mm. 
Spermatheca length 0.10 mm., 1.0 times 

Type material.— The holotype female of 
Arthroceras fulvicome {CNQ, no. 9100) was 
collected at Robson, British Columbia on 
21 June, 1948 by H. R. Foxlee. The holo- 
type female o{ Arthroceras fulvicome nigri- 
capite (USNM, no. 68171) was collected at 
Yosemite, California on 11 June, 1935 by 
A. L. Melander. The holotype female o{ Ar- 
throceras fulvicome subsolanum (USNM, 
no. 68172) was collected at Mt. Washing- 



ton. New Hampshire, by A. T. Slosson. The 
holotype female of Art hwceras subaquilum 
(USNM, no. 68173) was collected at Banff, 
Alberta on 27 August, 1935 by A. L. Me- 

Seasonal activity.— Adults of Art hwceras 
fulvicorne have been collected with aerial 
nets, in Malaise traps baited with dry ice 
(carbon dioxide), and at lights. In the col- 
lections examined adults were taken from 
20 May until 20 August, with the majority 
of specimens collected in July. Females ( 1 29) 
were collected 2.7 times more often than 
males (47) with no evidence of protandry. 

Distribution (Fig. 12).— Arthroceras ful- 
vicorne is a widespread species found from 
Maine to western Ontario in the east and 
from Colorado to California north to Al- 
berta and British Columbia in the west. 

Specimens examined (176). — UNITED 
STATES: CALIFORNIA: Phillips Station 
(Placer County); Snowline Camp (Eldorado 
County); Blodgett Forest, 1 3 mi E George- 
town; Bumble Bee; Giant Forest Camp- 
ground, Sequoia National Park; Yosemite 
Carson Pass (Alpine County). COLO 
RADO: West slope of Loveland Pass, 9850' 
Great Sand Dunes National Monument 
7600'; St. Louis Creek Campground, 3 mi 
SW Eraser, 8800'. IDAHO: Moscow Mt. 
Long Valley, Alpha; Moscow; Chatcolet 
Cub River Canyon, Thomas Spring; Bear 
Valley (Valley County). MAINE: Allagast 
Point, St. Francis. MICHIGAN: Pequam- 
ing; shore of Lake Superior, Marquette. 
MONTANA: Glacier National Park, Med- 
icine Lake; Glacier National Park, Lake 
MacDonald. NEW HAMPSHIRE: Mt. 
Washington; Mt. Alpine. NEW YORK: Mt. 
Slide (Ulster County), 3500'. OREGON: 
Crater Lake; Chief Joseph Mt., Joseph; 
Wallowa Lake; Velvet Creek, 28 mi SE 
Union; Ladd Canyon, 14 mi S La Grande, 
4280'; Butte Lookout Station, 25 air mi E 
Medford; Lake of the Woods; Jordan Creek, 
28 mi SSW La Grande, 4840'. UTAH: Lo- 
gan Canyon; Spring Hollow; Tony Grove 
Junction; Blacksmith Fork Canyon; Willard 

Basin, 9300'; Monte Cristo; Mendon Cold 
Spring; Green Canyon; Smithfield Canyon; 
Logan; Millcreek Canyon; Kaller Hollow 
Camp, 22 mi NNW Vemel, 8900'. VER- 
MONT: Camels Hump, Bolton, 4100'. 
WASHINGTON: Nooksack River, Mt. 
Baker; Summerland Trail, Mt. Ranier; Gla- 
cier Peak Wilderness, 27 mi ESE Darring- 
ton; Moscow Mts.; Field Spring S. P., 4 mi 
S Anatone, 3500-4000'; Bald Knob Camp- 
ground, Mt. Spokane S.P., 4800-5200'; Mt. 
Ranier National Park. WYOMING: Elk 
Mountain; Old Faithful area, Yellowstone 
National Park; Trout Lake, 1 mi SW Round 
Prairie, Yellowstone National Park, 6900'; 
Grand Teton National Park; Bottle Creek 
Camp, 7 mi SW Encampment, 8800'; Low- 
er Green River Lake, Wind River Range; 
Old Faithful area, Yellowstone National 
Park, 7000-7500'; South Brush Creek 
Campground, Medicine Bow National For- 
est, 8000-8500'. CANADA: ALBERTA: 
Island Lake, Coleman, 4500'; Banff; Wa- 
terton; Island Lake, Coleman, 4500'; Mtn. 
Hope Mts.; Terrace Mtn. NOVA SCOTIA: 
Mile 15, Highland Road. ONTARIO: Sud- 
bury; Macdiarmid; Burke Falls. 

Arthroceras leptis (Osten Sacken) 

Arthropeas leptis Osten Sacken (1878: 223). 
Arthroceras leptis (Osten Sacken) (1878: 

223); Aldrich (1905: 213); James (1965: 

298); Leonard (1930: 52). 

Arthroceras leptis is easily separated from 
A. pollinosum by the length of the apical 
flagellomere being less than 1.5 times as wide 
as the flagellum and the setae on the hind 
femur being predominately fuscous, short, 
and appressed. Arthroceras leptis and A. ful- 
vicorne can be easily separated by the char- 
acters in the key. 

Male. — Length 5.8-6.5, 6.2 mm. Head 
with ocellar tubercle fuscous to black, prui- 
nosity light grey; setae pale yellow to pale 
fuscous, elongate, abundant. Eyes brown to 
fuscous. Frons fuscous, pruinosity grey; se- 



tae pale yellow to pale fuscous, elongate, 
scattered dorsally. Antenna length 0.46- 
0.92, 0.67 times length of head; scape dark 
brown, length 0.3-0.6, 0.4 times width, 0.5- 
0.8, 0.6 times length of pedicel, setae absent; 
pedicel length 0.5-0.8, 0.6 times width, se- 
tae pale yellow to pale brown, moderately 
long, subapical; flagellum length 1.9-3.9, 3.3 
times width, 2.1-4.3, 3.0 times combined 
length of scape and pedicel, setae pale brown, 
scattered subapically; 7 annuli, apical flag- 
ellomere length 0.9-1 .4, 1 .2 times flagellum 
width. Gena with dense grey pruinosity; fa- 
cial setae pale yellow, elongate, abundant. 
Clypeus with dense grey pruinosity; setae 
pale yellow to gold, elongate, abundant. 
Maxillary palpus fuscous, pruinosity grey, 
setae black, elongate, abundant; basal seg- 
ment length 2.4-3.0, 2.5 times width; apical 
segment length 3 . 2-3 .3,3.2 times width, 1.1- 
1.4, 1.2 times length of basal segment. La- 
bellum fuscous, pruinosity grey; setae fus- 
cous, elongate, scattered. Postocular setae 
pale yellow to gold, elongate, abundant ven- 
trally, becoming shorter dorsally, not reach- 
ing vertex. 

Thorax dark brown, pruinosity yellowish 
brown to grey; setae gold, elongate, abun- 
dant; vitta generally indistinct, occasionally 
dark brown, faint. Postpronotal lobe with 
setae brown, elongate, on anterior half. 
Pleura fuscous in ground color, pruinosity 
grey; setae dark yellow to gold, elongate, 
abundant on propleuron, on dorsal half of 
anepistemum, scattered dorsally and ven- 
trally on katepistemum, absent on anepi- 
meron, in caudodorsal patch on meron, 
scattered on dorsal half of metapleuron. 
Halter stalk dark yellowish brown, capitu- 
lum fuscous, pruinosity light grey. Scutellum 
fuscous, with grey pruinosity; setae brown, 
elongate, scattered. Postnotum fuscous, with 
dense grey pruinosity; setae dark yellow to 
brown, elongate, abundant. 

Wing as in A. pollinosum (Fig. 1 4). Length 
5.7-6.3, 6.1 mm, width 1.8-3.0, 2.2 mm, 
length 2.2-3.2, 2.9 times length. Membrane 
pale brown; veins brown; pterostigma 

slightly darker than membrane. M, and M2 
generally petiolate, occasionally originate 
contiguously from discal cell. Squama pale 
brown; setae pale yellow, elongate, entire. 

Coxae fuscous, pruinosity grey, setae gold, 
elongate, abundant; femora fuscous to dark 
brown, glossy or with pruinosity light grey; 
tibiae and tarsi pale brown to fuscous. Em- 
podia and pulvilla dark brown. 

Abdomen dark brown, pruinosity light 
grey; setae pale yellow, elongate, abundant. 
Terminalia as in A. pollinosum (Figs. 7-9). 

Female. -Length 7.7-8.9, 8.2 mm. Head 
with ocellar tubercle dark brown; setae gold- 
en, shorter than in male. Eyes brown. Frons 
fuscous, pruinosity pale brown, pruinosity 
on lateral and ventral margins silver to grey; 
setae gold, moderately long, scattered sub- 
apically. Antenna (Fig. 5), length 0.78-0.86, 
0.82 times length of head; scapal length 0.3- 
0.6, 0.5 times width, 0.4-1.0, 0.7 times 
length of pedicel; pedicel length 0.5-0.8, 0.6 
times width; flagellum length 3.1-3.9, 3.5 
times width, 2.8-4.4, 3.3 times combined 
length of scape and pedicel; apical flagello- 
mere length 0.8-1.3, 1.1 times flagellum 
width. Clypeal setae not as abundant as in 
male. Basal segment of maxillary palpus 
length 1.9-3.0, 2.4 times width; apical seg- 
ment length 2.7-3.1, 2.9 times width, 1.5- 
2.7, 1.8 times length of basal segment. Post- 
ocular setae reach vertex. 

Thoracic setae shorter than in male. Vitta 
dark brown, separated by yellowish grey 
pruinosity. Metapleural setae scattered along 
dorsal margin. 

Wing length 7.0-8.0, 7.6 mm, width 2.7- 
3.0, 2.9 mm, length 1.2-1.6, 1.3 times width. 
Veins Mi, M2, M, originate separately from 
discal cell. 

Terminalia and internal reproductive or- 
gans as in A. pollinosum (Figs. 19-21). 

Type material.— The lectotype female, 
here designated, of Arthropeas leptis (MCZ) 
was collected in the White Mountains, New 
Hampshire, by E. P. Austin. 

Seasonal activity. — In the collections ex- 
amined Arthroceras leptis was taken from 7 



Fig. 13. Arthroceras leptis, dislrihution. 

July until 1 4 August. Females ( 1 9) were col- 
lected 1 .4 times more often than males ( 1 4), 
with no evidence of protandry. 

Distribution (Fig. \3).—Arthroceras lep- 
tis is found from New Hampshire to Mich- 
igan in the east and from Oregon to Wash- 
ington in the west. 

Specimens examined (33). — UNITED 
HAMPSHIRE: Mt. Washington; White Mts. 
NEW YORK: Whiteface Mts., Adiron- 
dacks; Lake Tear (Essex County). ORE- 
GON: Mt. Hood. VERMONT: Jay Peak, 
3400-3800'. WASHINGTON: Mt. Ranier; 
Sunrise, Mt. Ranier; Deer Lake near Chew- 
elah; 42 mi SE Randle; Bald Knob Camp- 
ground, Mt. Spokane S. P., 4800-5200'. 
CANADA: ALBERTA: Johnston Canyon, 
Park, Valle View. QUEBEC: Mt. Oxford. 

Arthroceras poUinosum Williston 

Arthroceras poUinosum Williston (1886: 
108); James (1965: 298); Leonard (1930: 
53, 1931: 322); Nagatomi (1966: 54). 

Leptis pruinosa Bigot (1887: 115); Leonard 
(1931: 322); Nagatomi (1966: 54). 

Arthroceras poUinosum is easily separat- 
ed from both A. fulvicorne and A. leptis by 
the length of the apical flagellomere being 
greater than 1.5 times the width of the fla- 
gellum and the setae on the hind femur being 
pale yellow, elongate, and suberect. 

Male. -Length 5.2-5.8, 5.6 mm. Head 
with ocellar tubercle fuscous to black, pru- 
inosity grey; setae stramineous to brown, 
elongate. Eyes fuscous. Frons fuscous with 
dense silver grey pile. Antenna fuscous, 
pruinosity light grey, length 0.84-0.95, 0.89 
times length of head; scapal length 0.6-0.8, 
0.7 times width, 0.8-1.0, 0.9 times length 
of pedicel, setae pale yellow, short; pedicel 
length 0.6-0.7, 0.7 times width, setae pale 
yellow, short; flagellum length 4.7-6.0, 5.3 
times width, 3.5-4.3, 4.0 times combined 
length of scape and pedicel, annuli 7, apical 
flagellomere length 1.6-2.2, 1.9 times width 
of flagellum, setae fuscous, short, apical. 
Gena fuscous, pruinosity dense silver grey; 
facial setae white to pale yellow, elongate, 
abundant. Clypeus pruinosity dense silver 
grey; setae pale yellow, elongate, on lateral 
margins. Maxillary palpus fuscous, pru- 
inosity light grey, setae white to pale yellow, 
elongate, abundant; basal segment length 
1.4-2.2, 2.0 times width; apical segment 
length 2.4-4.3, 3.3 times width, 1 . 1-2. 1 , 1 .5 
times length of basal segment. Labellum 
dark brown to fuscous, pruinosity grey; se- 
tae a mixture of short fuscous setae and 
elongate, pale yellow setae. Postocular setae 
pale yellow, elongate, abundant ventrally, 
absent dorsally. 

Thorax fuscous in ground color, pruinos- 
ity dense grey yellow; vitta indistinct; setae 
pale yellow, elongate, scattered. Postprono- 



Figs. 14-2L Arthroceras pollinosum. 14, Wing. 15, Male epandrium, cerci, ventral plate of proctiger. 16, 
Male gonocoxite, ventral view. 17, Male gonocoxite, dorsal view. 18, Male endophallus, lateral view. 19, Female 
terminalia, dorsal view. 20, Female terminalia, ventral view. 21, Female furca, spermatheca. Abbreviations: (C) 
Cerci. (F) Furca. (Sp) Spermatheca. (S8) Stemite 8. (SIO) Stemite 10. (T9) Tergite 9. (TIO) Tergite 10. 



tal lobe with setae pale yellow, elongate on 
anterior half. Pleura fuscous in ground col- 
or, pruinosity dense silver grey; setae pale 
yellow on propleuron, dorsal and caudal 
margin of anepistemum, scattered along 
dorsal margin of katepistemum, along dor- 
socaudal margin of meron, scattered dor- 
socaudally on metapleuron, absent on ane- 
pimeron. Halter stalk yellow to dark brown, 
capitulum fuscous. Scutellum with pru- 
inosity dense grey yellow; setae pale yellow, 
elongate, abundant. Postnotum with pru- 
inosity grey, laterotergite with pruinosity 
grey; setae pale yellow, elongate, abundant. 

Wing (Fig. 14) length 4.8-5.7, 5.2 mm, 
width 1.7-2.0, 1.9 mm, length 2.7-2.9, 2.8 
times width. Membrane pale yellow; veins 
pale brown; pterostigma absent to pale 
brown. Fork of R4+5 originates above or dis- 
tad to apex of discal cell. Squama and mar- 
ginal setae pale yellow. 

Coxa fuscous, pruinosity dense grey; fem- 
ora fuscous, pruinosity grey, setae pale yel- 
low, elongate; tibiae pale brown; tarsi fus- 
cous. Empodia and pulvilli fuscous. 

Abdomen fuscous in ground color, pru- 
inosity dense grey; setae pale yellow, elon- 
gate, abundant. Epandrium, cerci, and ven- 
tral plate of proctiger (Fig. 15). Gonocoxite 
in ventral view (Fig. 1 6); in dorsal view (Fig. 
1 7). Endophallus (Fig. 1 7) flattened laterally 
in dorsal view, length 0.36 mm, elongate; 
in lateral view endophallus (Fig. 18) with 
anterior third expanded. Endophallic hilts 
(Fig. 1 7) thick, heavily sclerotized, tapered 
anteriorly, separated medially. 

Female. — Length 5.5-7.5, 6.7 mm. Head 
with ocellar tubercle pollinose; setae pale to 
dark yellow. Frons pollinose; setae pale to 
dark yellow, elongate. Antenna (Fig. 6) fus- 
cous, apical third of pedicel and basal half 
of flagellomere 1 + 2 yellow, length 1 .0- 
1.1, 1.1 times length of head; scapal length 
0.6-0.8, 0.7 times width, 1.3 times length 
of pedicel; pedicel length 0.4-0.6, 0.5 times 
width; flagellum length 5.3-6.0, 5.6 times 
width, 4.1-6.0, 4.9 times combined length 
of scape and pedicel, apical flagellomere 

length 1 .9-2.5, 2.2 times width of flagellum. 
Gena pollinose. Clypeus pollinose, setae pale 
to dark yellow, elongate, lateral. Maxillary 
palpus pollinose to fuscous, setae pale to 
dark yellow, elongate, abundant; basal seg- 
ment length 1 .8-2.4; 2. 1 times width; apical 
segment length 2.3-2.8, 2.5 times width, 1.5- 
1.7, 1.6 times length of basal segment. Post- 
ocular setae pale to golden yellow, elongate, 
abundant, extending to vertex. 

Thorax pollinose, setae fuscous, elongate, 
with small fuscous spot at base of each. 
Postpronotal lobe with setae entire. Pleura 
with pruinosity dark yellow to grey. Halter 
dark yellow, pruinosity light grey. Scutellum 
dark yellow, glossy or pruinosity light grey. 
Postnotum dark yellow, pruinosity of me- 
dian area light grey. 

Wing length 6.0-7.0, 6.5 mm; width 2.2- 
2.5, 2.4 mm; length 2.6-3.0, 2.8 times width. 

Coxa dark yellow, pruinosity grey; fem- 
ora dark yellow, glossy; tibiae and tarsi pale 
brown to fuscous, pruinosity light grey. 

Abdomen fuscous in ground color, pru- 
inosity dense yellowish grey. Terminalia in 
dorsal view (Fig. 1 9), in ventral view (Fig. 
20). Internal reproductive organs (Fig. 21). 
Furca " Y"-shaped, anterior apodeme short. 
Spermatheca length 0.09 mm, width 0.10 

Type material.— The lectotype male, here 
designated, o{ Arthroceras poUinosum (SEM, 
no. 5846) has no locality data on the spec- 
imen. The lectotype male, here designated, 
of Leptis pruinosa (BMNH) was collected 
at Mount Hood, Oregon. 

Seasonal activity.— Adults of Arthroceras 
pollinosum have been collected at eleva- 
tions as high as 9800 feet, from Phacelia, 
with an aerial net, or in Malaise traps, and 
at lights. In the collections examined, adults 
were taken from 6 June until 27 July. Males 
(45) were collected 1.1 times more often 
than females (40), with no evidence of pro- 

Distribution (Fig. 22).— Arthrocer as pol- 
linosum is a western species found from 
California and New Mexico north to Wy- 



Fig. 22. Arthroceras pollinosum. distribution. 

oming and Washington. A single female was 
collected in Wisconsin. 

Specimens examined (85). -UNITED 
Rowdy Creek; Prairie Creek; Weott; moun- 
tains near Orick; Dry Lagoon Beach State 
Park; Blacksburg Road; Areata. COLO- 
RADO: St. Louis Creek Camp, 3 mi SW 
Fraser, 8800'; Aspen; Dolittle Ranch, Mt. 
Evans, 9800'; Chicago Creek, 8800'; Happy 
Hollow; Little Beaver; Clear Creek; 7 mi N 

Ward; Lake City, 9000'; Science Lodge; west 
end of Grand Mesa. NEW MEXICO: Ther- 
ma; Tajique; Hyde State Park, 8 mi NE San- 
ta Fe, 8700'; Santa Fe Campground near 
Santa Fe. OREGON: Robinson Butte Look 
Out, 25 air mi E Medford; Oregon Creek; 
Portland; Corvallis; Forest Grove; 14 mi S 
Ranier; 20 mi E Waldport; Black Rock, 10 
mi SW Dallas; Beverly Beach. UTAH: Kal- 
er Hollow Camp, 22 mi NNW Vernal. 
WASHINGTON: Canyon Creek; Seattle; 



Vancouver; Electron; 7 mi E Randle, Rt. 
Creek Camp, 7 mi SW Encampment, 8800'; 
Lower Green River Lake, Wind River 
Range, 8000'. 


I thank L. M. Page, G. L. Godfrey, and 
W. E. LaBerge for reviewing this manu- 
script and E. Steger for her editorial com- 
ments. I thank the curators of the following 
institutions and collections for the loan of 
material relevant to this study: American 
Museum of Natural History, P. Wygodzin- 
sky; British Museum of Natural History 
(BMNH), J. Chainey; California Academy 
of Sciences, P. H. Amaud, Jr.; Canadian 
National Collection (CNC), H. J. Teskey; 
Cornell University, L. L. Pechuman; Flor- 
ida State Collection of Arthropods, H. V. 
Weems, Jr.; Kansas State University, H. D. 
Blocker; Museum of Comparative Zoology, 
Harvard University; Ohio State University, 
C. A. Triplehom; Oregon State University, 
J. D. Lattin; Royal Ontario Museum, G. 
Wiggins; W. J. Turner Collection; United 
States National Museum (USNM), W. 
Mathis; University of British Columbia, S. 
G. Cannings; University of California, 
Berkeley (California Insect Survey), E. I. 
Schlinger; University of California, Davis, 
R. O. Schuster; University of Colorado, U. 
N. Lanham; University of Georgia, C. L. 
Smith; University of Idaho, J. B. Johnson; 
University of Kansas, Snow Entomological 
Museum (SEM), G. W. Byers; University 
of Michigan, T. E. Moore; University of 
New Hampshire, D. S. Chandler; Univer- 
sity of Vermont, R. T. Bell; Utah State Uni- 
versity, W. J. Hanson; Washington State 
University, W. J. Turner. This paper is a 
contribution of the Illinois Natural History 
Survey supported in part by a grant from 
the University of Illinois Research Board 
and an Ernst Mayr Grant from the Museum 
of Comparative Zoology, Harvard Univer- 

Literature Cited 

Aldrich, J. M. 1905. ACatalogueof North American 

Diptera (or Two-Winged Flies). Smithson. Misc. 

Coll. 46(1444): 1-680. 
Bigot, J. M. F. 1887 Dipteres nouveaux ou peu con- 

nus. Leptidi, Muscidi. Bull. Soc. Zool. France 12: 

Coquillett, D. W. 1910. The type-species of the North 

American genera of Diptera. Proc. U.S. Natl. Mus. 

27: 499-647. 
James, M. T. 1965. Family Xylophagidae, pp. 296- 

298. In Stone, A. et al., eds., A Catalog of the 

Diptera of America North of Mexico. U.S. Dep. 

Agric. Agric. Handb. 276: 1-1696. 
James, M. T. and W. J. Turner. 1981. Family Rhagi- 

onidae. 33, pp. 483-488. In McAlpine, J. R. et 

al.. eds.. Manual of Nearctic Diptera. Res. Br., 

Agric. Canada Monogr. 27(1): 1-674. 
Leonard, M. D. 1930. A revision of the dipterous 

family Rhagionidae (Leptidae) in the United States 

and Canada. Mem. Am. Entomol. Soc. 7: 1-181. 
. 1931. Some notes on my revision of the 

Rhagionidae (Diptera). Trans. Am. Entomol. Soc. 

57: 321-323. 
McAlpine, J. F. 1981. Morphology and terminolo- 
gy—Adults. 2, pp. 9-63. In McAlpine, J. F. et al., 

eds.. Manual of Nearctic Diptera. Res. Br., Agric. 

Canada Monogr. 27(1): 1-674. 
Nagatomi, A. 1955. A new genus and species of the 

dipterous family Coenomyiidae from Japan. Mu- 

shi 29: 57-60. 
. 1 966. The Arthrocems of the world (Diptera, 

Rhagionidae). Pac. Insects 8: 43-60. 
. 1970. i^^55//r/>//a Paramonov, a new synonym 

of .4r//7rac£'ra5 (Diptera, Rhagionidae). Mem. Fac. 

Agr. Kagoshima Univ. 7: 293. 
. 1982. Geographical distribution of the lower 

Brachycera (Diptera). Pac. Insects 24: 139-150. 
. 1984. Male genitalia of the lower Brachycera 

(Diptera). Beitr. Entomol. 34(1): 99-157. 
Nagatomi, A. and K. Iwata. 1976. Female terminalia 

of lower Brachycera. I. (Diptera). Beitr. Entomol. 

Berlin 26: 5-46. 
Osten Sacken, C. R. 1 878. Catalogue of the Described 

Diptera of North America (2nd ed.). Smithson. 

Misc. Coll. 16(270): 1-276. 
Ouchi, Y. 1 943. Diptera Sinica. Coenomyiidae 1 . On 

a new genus belonging to the family Coenomyiidae 

from east China. Shanghai Sizenkagaku Kenkyu- 

sholho 13: 493-495. 
Paramonov, S. J. 1929. Dipterologische Fragmente. 

Trav. Mus. Zool. Kieff. 7: 181-182. 
Stuckenberg, B. R. 1973. The Athericidae, a new fam- 
ily in the lower Brachycera (Diptera). Ann. Natal 

Mus. 21(3): 649-673. 
Williston, S. W. 1886. On two interesting new genera 

of Leptidae. Entomol. Am. 2: 105-108. 


89(2), 1987, pp. 264-268 





E. E. Grissell and L. De Santis 

(EEG) Systematic Entomology Laboratory, BBII, Agricultural Research Service, 
U.S.D.A., % U.S. National Museum NHB 168, Washington, D.C. 20560; (LDS) Director, 
Museo de La Plata, Paseo del Bosque, 1 900 La Plata, Republica Argentina. 

Abstract.— Erixestus pachy neuron Grissell and De Santis, n. sp., is described from Ar- 
gentina. This species was reared from eggs of Calligrapha polyspila (Germar) (Chryso- 
melidae) collected on Sida rhombifolia L. (Malvaceae). The genus Erixestus heretofore 
has been known only from the type species, E. winnemana Crawford, from the north- 
eastern Nearctic, reared from eggs of Calligrapha spp. The two species of Erixestus are 
compared and the geographic range of £". winnemana is expanded westward to Idaho and 
New Mexico. 

Since its description in 1910, the genus 
Erixestus has been represented by a single 
species distributed in the northeastern 
Nearctic (Burks, 1979). The genus was de- 
scribed for the species winnemana (Craw- 
ford, 1910), an internal parasitoid of eggs 
of the chrysomelid genus Calligrapha. 
Among New World pteromalids, Erixestus 
is unique as an internal egg parasite. Recent 
discovery of an undescribed species of Erix- 
estus in Argentina, also an internal egg par- 
asite of Calligrapha, suggests that the genus 
is both widespread and perhaps quite spe- 
cialized in its biology. The large number of 
Calligrapha species (ca. 100) and their ex- 
tensive geographic distribution throughout 
the New World (Blackwelder, 1 944; Wilcox, 
1972) suggest that Erixestus may be more 
widespread than present records indicate. 

The new species of Erixestus described 
below was collected by H. A. Cordo (U.S. 
Department of Agriculture, Biological Con- 
trol of Weeds Laboratory, Hurlingham, Ar- 
gentina) in connection with studies of Cal- 

ligrapha polyspila (Germar) on Sida 
rhombifolia L. in Argentina. The chryso- 
melid is being studied as a possible biocon- 
trol agent for Sida spinosa L. (Malvaceae), 
an introduced weed of southeastern United 

Erixestus pachyneuron Grissell and 
De Santis, New Species 

Figs. 1-4 

Female. — Body length 0.9 to 1.2 mm. 
Black with faint blue or green reflections on 
the head. Abdomen brownish black. Scape, 
antenna, mouth parts, tegula, and legs (in- 
cluding coxae) pale yellowish white. Wings 
hyaline with yellowish veins. Body polished 
except occiput, anterior part of mesoscu- 
tum, scutellum, and axillae faintly aluta- 
ceous. Propodeum polished except nucha 
reticulate, medially with 3 complete longi- 
tudinal carinae intersected by transverse ca- 
rina, partial longitudinal carinae present lat- 

Head and thorax with sparse, but obvious 



Figs. 1, 2. Erixestus pachyneuron (female). 1, Forewing. 2, Antenna. 

setae (some 2 x or more as long as diameter 
of ocellus). Eyes glabrous. Midlobe of meso- 
scutum with irregularly placed long setae 
(ca. 8 to 16), sidelobes with 6 or 7, axillae 
each with 5 or 6, scutellum with 3 pairs. 
Hindcoxae and lateral regions of propo- 
deum covered with long white setae. 

Head as wide as thorax. Frontovertex 
width greater than eye height. Clypeus with 
anterior edge bilobed, malar distance about 
as long as longitudinal diameter of an eye, 

without genal suture. Ocelli in obtuse tri- 
angle, posterior ocellus at least 2 x own di- 
ameter from inner margin of eye. Dorsal 
edge of occiput rounded. Antennae inserted 
midway between median ocellus and free 
edge of clypeus; antennal proportions as 
shown in Fig. 2. Collar of pronotum with 
anterior edge vaguely carinate. Spiracle el- 
liptic, anterior border obscured by meta- 
notum which overhangs it. Wing with dis- 
tribution of setae as in Fig. 1. Ratio of 



Figs. 3, 4. Erixestus pachyneiiron (male). 1, Forewing. 2. Antenna. 

postmarginal : marginal veins 0.57 ± SD = 
0.05 (range 0.53-0.64, n = 10), of stigmal : 
marginal veins 0.48 ± 0.03 (range 0.45- 
0.50, n = 10). Tibial spur ca. half length of 
basitarsus. Petiole of abdomen short 
(scarcely visible), much wider than long. 
Gaster oval, nearly as long as head and tho- 
rax together; first three terga subequal in 
length, remainder together shorter than Tl . 
Male. — Body length 0.7 to 1.1 mm. Sim- 

ilar to female, except body with intense green 
to bronze reflections. Sculpture of head and 
thorax more strongly developed than that 
of female. Antennal club less thickened (Fig. 
4). Forewing marginal vein much thickened 
(Fig. 3). Ratio of postmarginal : marginal 
veins 0.60 ± 0.05 (range 0.53-0.67, n = 10), 
for stigmal : marginal veins 0.44 ± 0.04 
(range 0.38-0.55, n = 10). 
Type material. — Holotype $, allotype 3, 



32 6 and 23 $ paratypes from ARGEN- 
TINA, Buenos Aires Province, Dique Lu- 
jan, 1 February and 24 and 27 March 1976, 
H. A. Cordo, reared from egg mass of Cal- 
ligrapha polyspila on Sida rhombifolia; 40 
6, 50 2 paratypes from ARGENTINA, Entre 
Rios Province, 40 km S. Gualenguaychu, 4 
January 1976, H. Cordo, reared from eggs 
of Calligrapha sp. The holotype, allotype, 
and 36 paratypes are deposited in the Mu- 
seo de La Plata, La Plata, Argentina. One 
hundred and nine additional paratypes are 
deposited in the U.S. National Museum, 
Washington, D.C., the British Museum 
(Natural History), and the Canadian Na- 
tional Collection. 

Variation. — Specimens of Erixestus 
pachyneuron vary little in size or coloration 
except that males are slightly smaller than 
females and are metallic blue or green rather 
than black. The abdomens of specimens 
preserved in alcohol tend to have a brown 
or yellowish cast. 

^'\o\o%y . — Erixestus pachyneuron has 
been reared from Calligrapha polyspila eggs 
on Sida rhombifolia. This species of Sida is 
widespread throughout the tropics (M. H. 
Sachet, Department of Botany, Smithsoni- 
an Institution, personal communication) and 
is associated with disturbed areas. 

Discussion.— Erixestus pachyneuron may 
be distinguished from E. winnemana by the 
following characters: in both sexes of pachy- 
neuron the postmarginal vein averages less 
than two-thirds the length of the marginal 
vein (0.59 ± 0.05, n -= 20), whereas in win- 
nemana these veins (Figs. 5, 6) are nearly 
equal (0.97 ± 0.04, n = 20); in both sexes 
of pachyneuron the stigmal vein averages 
less than one-half the marginal vein (0.45 ± 
0.04, n = 20), whereas in winnemana the 
stigmal vein averages two-thirds the mar- 
ginal (0.66 ± 0.4, n = 20); male pachyneu- 
ron have the marginal vein noticeably ex- 
panded throughout (cf. Fig. 3, pachyneuron 
with Fig. 6, winnemana); in both sexes of 
pachyneuron the antennae are white (dark 
brown to black in winnemana); in male 

Figs. 5, 6. Erixestus winnemana, forewing vena- 
tion. 5, Female. 6, Male. 

pachyneuron the flagellomeres are quadrate 
to transverse with short setae (Fig. 4), 
whereas in winnemana each flagellomere is 
longer than wide with long setae (Fig. 7). 

Erixestus winnemana Crawford, 1910 

Figs. 5-7 

Discussion.— This species was described 
from 10 specimens reared from eggs of Cal- 
ligrapha bigsbyana (Kirby) (now C multi- 
punctata bigsbyana) and 1 2 specimens from 
C scalaris (LeConte). All material was col- 
lected on Plummers Island, Maryland. At 
present there are a total of 1 6 syntypic spec- 
imens in the U.S. National Museum. As 
Crawford did not select a holotype at the 
time of description we herein designate and 
label a female as LECTOTYPE. The re- 
maining 15 specimens are paralectotypes. 
Distinguishing characters of this species are 
discussed under E. pachyneuron. 

Distribution.— This species is currently 
reported from Quebec, Maryland, and Vir- 
ginia (Burks, 1979). We have examined 
specimens from the following states which 
are all new records: New York, Illinois, 
Pennsylvania (ex Calligrapha spiraceae 
(Say)), Michigan, North Dakota (ex Cal- 
ligrapha scalaris), Kansas (ex Calligrapha 
scalaris), Wyoming (ex Calligrapha sp.), 
Idaho, and New Mexico. 

Erixestus winnemana might be expected 
to occur wherever its hosts do: Calligrapha 
scalaris is reported from throughout eastern 



Fig. 7. Erixestus winnemana (male). Antenna. 

United States and Canada from Quebec in 
the north to Georgia and Texas in the south; 
C. multipunctata bigsbyana occurs from 
Nova Scotia and British Columbia in the 
north, south to Oregon and Georgia; C. spi- 
raceae occurs from Maine and Michigan 
south to Pennsylvania (Wilcox, 1972). 

Biology. —This species is associated in the 
mid and eastern Nearctic with Calligrapha 
found on plants of northern hardwood for- 
ests: Calligrapha scalaris on Ulmus; C. m. 
bigsbyana on Salix, Tilia, and rarely Pop- 
ulus\ and C. spiraceae on Physocarpus (Wil- 
cox, 1972). Western specimens do not pro- 
vide enough data from which to draw 
conclusions about host relations. 


We thank H. A. Cordo for specimens used 
in this study, Richard White for informa- 

tion on chrysomelid distribution and hosts, 
M. H. Sachet for information on the distri- 
bution of Sida, and Steve Heydon for his 
critical review of the manuscript. 

Literature Cited 

Blackwelder, R. E. 1944. Checklist of the coleopter- 
ous insects of Mexico, Central America, the West 
Indies, and South America. Part 4. Bull. U.S. Natl. 
Mus. 185: 551-763. 

Burks, B. D. 1979. Pteromalidae, pp. 768-835. In 
Krombein, K. V. 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. 

Crawford, J. C. 1910. Three new genera and species 
of parasitic Hymenoptera. Proc. U.S. Natl. Mus. 
38: 87-90. 

Wilcox, J. A. 1972. A review of the North American 
chrysomeline leaf beetles. Bull. N.Y. State Univ. 
421: 1-37. 

89(2), 1987, pp. 269-274 


Richard D. Goeden 

Department of Entomology, University of California, Riverside, California 92521. 

Abstract.— l^ev/ (some initial) host-plant records are reported for Urophora caurina 
(Doane), U. fonnosa (Coquillett), U. rufipes (Curran), U. stenoparia Steyskal, and U. 
timberlakei Blanc and Foote. Urophora rufipes is initially reported from California. The 
host plants of these tephritids include one or more species of Acamptopappus, Chryso- 
thamnus, Grindelia, Gutierrezia, and Haplopappus, all noted to belong to the Subtribe 
Solidagininae of the Tribe Astereae of the Asteraceae. Synphagy among Urophora, Neaspi- 
lota, Procecidochares, Tephritis, and Trupanea species is documented and discussed. 

This is a companion paper to one to be 
written by G. C. Steyskal (retired). System- 
atic Entomology Laboratory, Agricultural 
Research Service, USDA, % National Mu- 
seum of Natural History, Washington, D.C., 
who will treat the taxonomy of some of the 
specimens reported herein. I report here new 
information on the host-plant relations of 
several native Urophora spp. (Diptera: Te- 
phritidae) resulting from the past six years 
of field studies on nonfrugivorous fruit flies 
in southern California, that section of the 
state defined and treated botanically by 

Materials and Methods 

The materials and methods used in the 
sampling of mature flower heads of Aster- 
aceae and the rearing of Tephritidae from 
samples were described by Goeden (1985). 
Sweep net collections of adults limitedly 
supplemented these rearings. Identifica- 
tions of most flies mentioned in this report 
were confirmed or made by G. C. Steyskal. 
All host-plant identifications were con- 
firmed or made by Andrew C. Saunders, 
Curator of the Herbarium of the University 
of California, Riverside. The plant nomen- 

clature used is that of Munz and Keck (1959) 
and Munz (1974); the insect nomenclature, 
that of Steyskal (1979). 

Results and Discussion 

Steyskal (1979) reviewed what little was 
known about the host plants of native North 
American species of Urophora. Except for 
the atypical species, U. acuticornis Steyskal 
reared from Lycium berlandieri Dunal (So- 
lanaceae), all known host plants of Neo- 
tropical Urophora are Asteraceae. Unlike the 
Palearctic Urophora, no North American 
Urophora has been reported from astera- 
ceous thistles. At least four Palearctic species 
have been introduced from Europe to Can- 
ada and the United States for the biological 
control of accidentally introduced, weedy 
thistles in the genera Carduus, Centaurea, 
and Cirsium (Steyskal, 1979; Julien, 1982). 
So far, no European species of Urophora has 
successfully been established in California 
for thistle control (Julien, 1982). Moreover, 
recent surveys of insects infesting the flower 
heads of native Cirsium thistles in northern 
California (Pemberton et al., 1985) and the 
above-ground shoots (including heads) in 
southern California (Goeden and Ricker, 



1986b, 1987a, b) yielded no rearing records 
of native Urophora. It may be as Steyskal 
(1979, p. 25) has suggested, that ". . . the 
American species of Urophora, at least in 
part, may eventually be referred to other 
genera inasmuch as none of them seem to 
be very closely related to any Palearctic 
species, ..." I offer the following informa- 
tion on host-plant relations of native Uro- 
phora in southern California as a contri- 
bution to a better understanding of the 
biology and ecology of the genus. The fol- 
lowing treatment of flies is alphabetical by 

Urophora caurina (Doane). — Steyskal 
(1979) hsted the host of U. caurina as the 
genus Grindelia. I never reared this species 
from a total of 1 3 samples of flower heads 
of five species o{ Grindelia that I have sam- 
pled since 1980. However, 2 <5 of t/. caurina 
were reared from a quantity of flower heads 
of Gutierrezia sarothrae (Pursh) Britton and 
Rusby collected in the Chihauhua Valley, 
NE San Diego Co., Il.xi.l982. Also, 8 <5 
and 8 9 of t/. caurina were reared from a 
quantity of flower heads of Haplopappus er- 
icoides (Lessing) Hooker and Amott ssp. 
blakei C. B. Wolf collected near Orcutt, San- 
ta Barbara Co., 12.xi.l980. Both records 
represent new host-plant genera for this te- 

Urophora formosa (Coquillett). — Was- 
bauer (1972) listed Chrysothamnus viscidi- 
florus (Hooker) Nuttall, Grindelia campo- 
rum Greene, G. nana Nuttall, Grindelia sp., 
Haplopappus squarrosus Hooker and Ar- 
nott ssp. grindelioides (deCandoUe) Keck, 
and H. venetus (Humboldt) Blake as hosts 
of U. formosa. In reviewing these host gen- 
era, Steyskal ( 1979) noted that some of these 
records may properly refer to U. caurina. I 
have reared the following specimens of U. 
formosa: 9 3 and 1 1 $ from flower heads of 
H. squarrosus spp. grindelioides collected 
above upper end of Kitchen Creek, Cleve- 
land Nat. Forest, San Diego Co., 9.ix.l980; 
7 6 and 7 9 from flower heads of//, venetus 
prob. spp. furfuraceus (Greene) Hall col- 

lected in Proctor Valley, SW San Diego Co., 
28.x. 1 98 1 ; 1 5 3 and 1 1 9 reared from flower 
heads of//, venetus spp. vernonioides (Nut- 
tall) Hall collected NW of Temecula, SW 
Riverside Co., 1 5.x. 1 980; 1 6 and 5 9 reared 
from flower heads of H. venetus prob. spp. 
vernonioides collected at Cardiff-by-the-Sea, 
San Diego Co., 15.x. 1980; 12 3 and 16 9 
reared from a quantity of flower heads of 
H. venetus ssp. vernonioides collected near 
U.S. Navy facility on Santa Cruz Island, 
Santa Barbara Co., 8.x. 1985. My rearing 
records definitely confirm Haplopappus as 
hosts of U. formosa; however, as with U. 
caurina, I have not reared U. formosa from 
any sample of Grindelia to date (including 
a sample of G. camporum). As the host rec- 
ords for U. formosa from Grindelia in Was- 
bauer (1972) are based on two or three sep- 
arate plant species and three independent 
sources and F. L. Blanc (in litt. 1986) has 
reconfirmed the record for G. camporum in 
Foote and Blanc (1963), this suggests a def- 
inite relationship under as yet undefined 
conditions with this fly. This host-plant re- 
lationship warrants additional study. The 
published "unpublished" host record for C. 
viscidiflorus in Wasbauer (1972) remains 
unconfirmed. It may properly refer to a 
sweep record or, perhaps, to U. timberlakei 
Blanc and Foote (see discussion below). 

Urophora n. sp. Steyskal. —To complicate 
matters further, the only tephritid that I have 
reared from Grindelia to date in southern 
California apparently is undescribed (Stey- 
skal, in litt. 1986). The sole host plant found 
to date for the robust individuals of this 
species appears to be G. hallii Steyermark, 
a plant species confined to dry flats in the 
Cuyamaca Mountains, San Diego Co. 
(Munz, 1974), and especially common 
around Lake Cuyamaca, where mature 
flower heads sampled in quantity on 
9.ix.l980 and 17.vii.l985 yielded only 2 S 
and 4 9 and 2 S and 1 9, respectively. Flower 
heads sampled in bulk on 2.vii, 31.vii, and 
14.viii.l985 from the same area yielded no 
flies. Factors involved in oviposition site 



Table 1 . Synphagy among Tephritidae reared with Urophora from samples of mature heads of Asteraceae 
from southern CaHfomia 1980-1985. 

Tephntid Genera (No. Species) 

Total No. (%) Adults 

Host Plant^ 

Sample Date" 



Acamptopappus shockleyi 

27 V 1982 



Trupanea (2) 

71(58), 21(17) 



A. sphaerocephalus 

21 V 1982 




20 (71), 2 (7) 



Chrysothamnus nauseosus 

3 IX 1982 







Trupanea (2) 

4 (24), 2 (12) 



C. viscidiflorus 

3 IX 1981 





Grindelia hallii 

9 IX 1981 



17 VIII 1985 



Gutierrezia microcephala 

9 IX 1981 



Gutierrezia sarothrae 

18 IX 1980 



11 X 1982 



Haplopappus acradenius 

8 XII 1983 





28 XI 1984 





11 XII 1984 





H. cuneatus 

16 X 1980 


33 (72) 



Urophora (2) 

8(17), 1(2) 

H. laricifolius 

21 X 1982 


25 (63) 



H. squarrosus 

9 IX 1980 




30 (48) 



H. venetus 

15 X 1980 




26 (90) 

15 X 1980 



Trupanea (2) 

8 (47), 2 (12) 



28 X 1981 

Trupanea (2) 

8 (32), 3 (12) 



8X 1985 




28 (93) 

See text for complete rearing record for Urophora. 

selection will be studied along with other 
aspects of the life history of this tephritid 
beginning in 1987. This species is especially 
suitable for a life history study among Neo- 
tropical Urophora because apparently no 
other species of Tephritidae infests the flow- 

er heads of G. hallii (Table 1 ); whereas, as 
discussed below, the other species of Uro- 
phora commonly are synphagous with other 
genera of Tephritidae. 

Urophora rufipes (Curran).— The single 2 
of U. rufipes reared from a small quantity 



of flower heads of Haplopappus acradenius 
(Curran) prob. spp. eremophilus (Greene) 
Hall collected at Mountain Springs, SW Im- 
perial Co., 8.xii.l983, represented the first 
host-plant record for this species (Was- 
bauer, 1972; Steyskal, 1979). Since then I 
have reared 4 3 and 2 9 and 4 $ and 1 9 from 
quantities of flower heads of H. acradenius 
collected along the south shore of Clark Dry 
Lake, SE San Diego Co., Il.xii.l984, and 
along the Coachella Canal above the ther- 
mal springs area. Riverside Co., 28. xi. 1 984, 
respectively. These also are the first Cali- 
fornia records of U. rufipes, known previ- 
ously only from Arizona (Steyskal, 1979). 

Urophora stenoparia Steyskal. — Similar- 
ly, 1 S and 1 9 of (7. stenoparia reared from 
a quantity of flower heads of Gutierrezia 
sarothrae collected at Pine Valley, Cleve- 
land Nat. Forest, San Diego Co., 18.ix.l980, 
represented the first host-plant record for 
this species (Steyskal, 1979). I additionally 
have reared 3 6 and 1 9 from flower heads 
of Gutierrezia microcephala (deCandolle) 
Gray and 1 9 from the sample of flower 
heads of//, cuneatus Gray along with the 8 
specimens of U. timberlakei Blanc and Foote 
reported below. I also swept 3 3 and 1 9 of 
U. stenoparia from Hymenoclea salsola 
Torrey and Gray during extensive field 
studies of the insect fauna of this common 
desert shrub (Goeden and Ricker, 1986a), 
which is not a host plant of this tephritid. 
Nineteen additional species of Tephritidae 
were swept from H. salsola, only one species 
of which infests the flower heads or other- 
wise reproduces on this common desert 
shrub (Goeden and Ricker, 1986, unpub. 

Urophora timberlakei.— Wasbaucr (1972) 
listed Gutierrezia microcephala (de- 
Candolle) Gray as a host plant of U. tim- 
berlakei, and Steyskal (1979) cited this ge- 
nus and Chrysothamnus as hosts. I have 
reared 1 3 and 7 9 of t/. timberlakei from 
flower heads of Acamptopappus shockleyi 
Gray, collected at the SE end of Kingston 

Mountains, NE San Bernardino Co., 
27.V.1982; 2 $ and 1 9 from flower heads of 
A. sphaerocephalus (Harvey and Gray) Gray 
collected at Snow Creek, Riverside Co., 
21.V.1982; 1 9 from flower heads of C ««m- 
seosus (Pallas) Britton collected in Landers 
Meadow, Sequoia Nat. Forest, Kern Co., 
3.ix. 1981; 1 6 and 2 9 from flower heads of 
C. teretifolius (Durand and Hilgard) Hall 
collected in Westgard Pass, Inyo Nat. For- 
est, Inyo Co., 9.ix.l986; 31 3 and 42 9 and 

3 <5 and 3 9 from flower heads of C. visci- 
dijlorus also collected in Landers Meadow 
on 3.ix.l981 and at Antelope Spring, NE 
Inyo Co., 15.ix.l982, respectively; 4 $ and 

4 9 from flower heads of Haplopappus cu- 
neatus collected in Lark Canyon, San Diego 
Co., 1 6.x. 1 980; and 8 <5 and 7 9 from flower 
heads of H. laricifolius Gray, 1 km NW of 
Kessler Peak, S end of Ivanpah Mountains, 
NE San Bernardino Co., 21.x. 1982. All my 
rearing records are for new host species. Two 
new host genera also are represented. The 
record for G. microcephala in Wasbauer 
(1972), originating from a host listing of R. 
H. Foote, but noted only as a sweep record 
by Blanc and FooiC (1961) and Foote and 
Blanc (1963), still lacks confirmation and is 

My field observations and rearing data 
suggest that all Urophora species native to 
southern California encountered to date are 
flower head-infesting, seed-feeding species, 
with the exception of U. acuticornis, which 
doubtfully belongs in the genus, judging from 
its distinctive host-plant affinities and at- 
tendant mode of development. Steyskal 
(1979) suggested that U. acuticornis likely 
will be referred to a distinct genus when 
more is known about American Myopitin- 
ae. Allen L. Norrbom (in litt. 1986) states 
that U. acuticornis not only is not a Uro- 
phora or myopitine, but rather is a trype- 

I have detected no sign of galls on the 
excised compound inflorescences that 
mainly constituted the flower head samples 



of Chrysothamnus, Gutierrezia, and Hap- 
lopappus reported above. Nor have I noted 
any enlarged flower heads in these and other 
host-plant genera mentioned in this report 
that are symptomatic of some other gallico- 
lous Tephritidae, e.g. Procecidochares in 
Chrysothamnus (Table 1 ). 

Of evolutionary and taxonomic signifi- 
cance is my observation that all of the con- 
firmed host plants of the southern Califor- 
nia Urophora mentioned in this report 
belong to the Subtribe Solidagininae of the 
Tribe Astereae (Munz and Keck, 1959). 
Thus, like so many Eurasian Urophora con- 
fined to hosts in the Tribe Cynareae, Sub- 
tribes Carduinae and Centaurinae, these 
southern California Urophora show a com- 
mon affinity for a definable group of host 
plants in the Asteraceae (Zwolfer, 1965; 
Steyskal, 1979). 

No life history of any Nearctic Urophora 
has been published. One difficulty involved 
is illustrated in Table 1. In southern Cali- 
fornia, at least, flower heads of Urophora 
host-plant species at most locations were 
commonly infested with other Tephritidae 
(synphagy). Genera commonly associated 
with Urophora are Neaspilota, Tephritis, and 
Trupanea. Rarely was more than one species 
of Urophora reared from a single sample. 
Associated genera differed qualitatively and 
quantitatively among some host-plant pop- 
ulations sampled, e.g. Haplopappus acra- 
denius and H. venetus (Table 1). Urophora 
commonly were reared in small numbers 
and usually have been poorly represented 
in my sweep collections, reflecting, perhaps, 
their low population densities in nature. In 
some flower head samples, however, Uro- 
phora outnumbered at least one associated 
tephritid species; in Chrysothamnus visci- 
diflorus, overwhelmingly so (Table 1). Sam- 
ples from two different species of Acamp- 
topappus appeared similarly composed. The 
undescribed Urophora apparently lacks te- 
phritid competitors for heads of Grindelia 
hallii\ whereas, Gutierrezia heads yield 

another genus of Tephritidae, i.e. Trupanea 
(Goeden, 1 985). Much remains to be learned 
about the host-plant relations of these and 
other Nearctic Urophora. 


My thanks to George Steyskal for his 
taxonomic help noted in the text, to Louis 
Blanc, Dick Foote and Al Norrbom for their 
reviews of early drafts of this paper, and to 
Don Ricker for his patience during stops to 
allow my many collecting forays. 

Literature Cited 

Blanc, F. L. and R. H. Foote. 1961. A new genus and 
five new species of California Tephritidae. Pan- 
Pac. Entomol. 37: 73-83. 

Foote, R. H. and F. L. Blanc. 1963. The fruit flies or 
Tephritidae of California. Bull. Calif Insect Surv. 
7. 115 pp. 

Goeden, R. D. 1985. Host-plant relations of Tru- 
panea spp. (Diptera: Tephritidae) in southern Cal- 
ifornia. Proc. Entomol. Soc. Wash. 87: 564-571. 

Goeden. R. D. and D. W. Ricker. 1986a. Phytoph- 
agous insect fauna of the desert shrub Hymenoclea 
salsola in southern California. Ann. Entomol. Soc. 
Am. 79: 39-47. 

. 1986b. Phytophagous insect faunas of the 

two most common, native Cirsium thistles, C. cal- 
ifornicum and C. proteanum, in southern Califor- 
nia. Ann. Entomol. Soc. Am. 79. (In press.) 

. 1987a. Phytophagous insect faunas of the 

native thistles, Cirsium brevistylum. C. congdonii, 
C. occidentale, and C. tioganum, in southern Cal- 
ifornia. Ann. Entomol. Soc. Am. 80. (In press.) 

. 1987b. Phytophagous insect faunas of native 

Cirsium thistles, C. mohavense, C neomexican- 
um, and C nidulum, in the Mojave Desert of 
southern California. Ann. Entomol. Soc. Am. 80. 
(In press). 

Julien, M. H. (ed.) 1982. Biological control of weeds: 
A world catalogue of agents and their target weeds. 
Commonw. Agric. Bur., Commonw. Inst. Biol. 
Control, Famham Royal, Slough. 108 pp. 

Munz, P. A. 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 and Los An- 
geles. 1681 pp. 

Pemberton, R. W., C. E. Turner, and S. S. Rosenthal. 
1985. New host records for tephritid flies (Dip- 
tera) from Cirsium and Saussurea thistles (Aster- 


aceae) in California. Proc. Entomol. Soc. Wash. tera: Tephritidae). Calif. Dep. Agric. Bur. Ento- 

87: 790-794. "^ol. Occas. Pap. 19. 172 pp. 

Steyskal, G. C. 1 979. Taxonomic studies on fruit flies Zwolfer, H. 1965. Preliminary list of phytophagous 

of the genus Urophora (Diptera: Tephritidae). insects attacking wild Cynareae (Compositae) in 

Entomol. Soc. Wash. Misc. Pub. 6 1 pp. Europe. Commonw. Inst. Biol. Control Tech. Bull. 

Wasbauer, M. W. 1972. An annotated host catalog 6:81-153. 
of the fruit flies of America north of Mexico (Dip- 


89(2), 1987, pp. 275-283 




Donald R. Davis 

Department of Entomology, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560. 

Abstract. — Examination of a small collection of Acrolophus in the collections of the 
Instituto de Zoologia in Havana, Cuba, has revealed the presence of three new species 
{A. fuscisignatus, A. guttatus, and A. basistriatus) and the first specimens collected of .4. 
niveipunctatus Walsingham since 1891. All species are fully described and illustrated. 

In 1980, the Academia de Ciencias de 
Cuba and Smithsonian Institution signed an 
agreement designed to promote cooperative 
research in the natural sciences. A vital part 
of this agreement encourages an exchange 
of scholars between our two countries. In 
February 1981, I was privileged to be one 
of the first entomologists to participate in 
this program. 

Although February was certainly not the 
most opportune season for Lepidoptera, a 
respectable sample of specimens, including 
28 species of Tineidae, was collected at five 
sites in western Cuba over an 1 1 day period. 
The purpose of this brief account is not to 
report upon what is hoped to be only the 
first of several trips to Cuba, but instead 
upon a small but interesting series oi Ac- 
rolophus found in the collections of the In- 
stituto de Zoologia in Havana. Most of the 
specimens were collected by Pastor Alayo 
and associates; all but one species were pre- 
viously undescribed; and none was encoun- 
tered during my brief excursion. 

Only five species of Acrolophus have been 
described from Cuba (Davis, 1984). Be- 
cause the unique holotypes of four of these 
{A. dimidiella Wlsm., A. niveipunctatus 
Wlsm., A. noctuina Wlsm., A. vitellus Poey) 
are now lost, their identities have been un- 

clear. Fortunately, one of the four species 
found in the collections of the Instituto de 
Zoologia has been determined as conspe- 
cific with A. niveipunctatus. At present, these 
represent the only examples of this species 
collected in nearly a century, which merely 
indicates again how poorly known the Cu- 
ban microlepidoptera are. It is further in- 
teresting to note that all of the species treat- 
ed herein, with the exception of A. 
fuscisignatus, new species, are members of 
the North American plumifrontellus group. 
This group, which was previously known to 
contain only A. plumifrontellus Clemens 
(Hasbrouck, 1964), is characterized pri- 
marily by the bifid apex of the male valva. 
Institutional acronyms referred to in this 
paper are: IZAC for Instituto de Zoologia, 
Academia de Ciencias de Cuba, Havana; 
and USNM for National Museum of Nat- 
ural History (formerly United States Na- 
tional Museum), Smithsonian Institution, 
Washington, D.C, USA. 

Acrolophus fuscisignatus Davis, 

New Species 
Figs. 1-3, 7-10, 23 

Adult (Figs. 1-3). — Length of forewing: S, 
10.0-14.5 mm; 9 15-17 mm. A relatively 
large species with strongly recurved labial 



Figs. 1-6. Adults. 1, Acrolophus fuscisignatus, paratype male, Pico Turquino, length of forewing 17.7 mm. 
2, A. fuscisignatus, paratype male, Moa, length of forewing 10 mm. 3, A. fuscisignatus, paratype male, Pico 
Turquino, length of forewing 15.6 mm. 4, A. guttatus. holotype male, Finca La Ciega, length of forewing 12 
mm. 5, A. niveipunctatus, male, Hongolosongo, length of forewing 13.9 mm. 6, A. basistriatus, holotype male, 
Cuabal de Gulindo, length of forewing 12.5 mm. 



palpi in the male. The forewing is white, 
variously marked with fuscous. 

Head: Vestiture mostly white with a fringe 
of fuscous hairs around eyes. Eye smooth 
with long dark lashes from posterior rim. 
Antenna approximately 0.4-0.5 the length 
of forewing, 61-64 segmented; scape en- 
tirely white; flagellum subserrate with gray- 
ish-white scales dorsally and often indis- 
tinctly banded with light brown; naked 
ventrally except for dense, very short whit- 
ish sensory setae. Labial palpus 3 segment- 
ed, extremely long in male and recurved 
over thorax to metanotum; relatively short 
in female, length about 2 x diameter of eye 
and porrect; vestiture mostly white with lat- 
eral surfaces of basal segment dark fuscous. 

Thorax: Pronotum mostly white, some- 
times with heavy suffusion of fuscous an- 
teriorly and medially. Venter white. Fore- 
wing white, variously marked by fuscous as 
shown in Figs. 1-3; most specimens with a 
prominent sinuate fuscous band extending 
longitudinally for most of the length of the 
wing along the medial vein; cilia white with 
scattered patches of fuscous. Hindwing uni- 
formly pale to dark gray; cilia sometimes 
paler and with whitish apices. Foreleg fus- 
cous to buff dorsally, white ventrally. Mid- 
leg slightly paler; femur brown dorsally; tib- 
ia mostly white usually with two dorsal 
brown spots; tarsi brown with white band- 
ing. Hindleg pale brown to nearly white dor- 
sally, white ventrally. 

Abdomen: Pale gray to buff dorsally, white 

Male genitalia: As shown in Figs. 7-10. 
Uncus relatively deeply bifid. Gnathos en- 
tire, consisting of a broadly rounded lobe. 
Valva simple, straight with apex evenly 
rounded. Aedoeagus moderately short, 
about two-thirds the length of valva, with 
an elongate cluster of 30 or more minute 
spines extending from middle to apex. 

Female genitalia: As shown in Fig. 23. 
Seventh sternum evenly rounded posterior- 
ly. Bursa copulatrix short, approximately 

1.5 X the length of posterior apophyses; 
ductus bursae very short, with heavily fur- 
rowed walls, corpus bursa simple, without 
signum or spicules. 

Immature stages unknown. 

Holotype.— <5, Trinidad Mts., San Bias, 
Cienfuego Province, Cuba; 5 May 1932, S. 
C. Bruner and A. Otero, E.E.A. Cuba Ento. 
No. 9967, USNM Type No. 100676 

Paratypes.— CUBA: Specific locality un- 
known: 1 3, W. Schaus Coll. (USNM). Ca- 
maguey Prov. [?]: Las Animas, Sierra Ran- 
gel, 1500 ft. [457 m]; 1 3, Aug. 1922, H. 
Roberto (USNM). Cienfuego Prov.: Same 
data as holotype; 2 $ (USNM). La Habana 
Prov.: Arroyo Nararrjo; 1 3, 13 May 1934, 
L. C. Scaramuzza (USNM). Santiago de las 
Vegas; 1 2, 15 May 1934, A. R. Otero 
(USNM); 1 9, 24 May 1932, A. R. Otero, 
E.E.A. Cuba Ento. No. 9967 (USNM). Hol- 
guin Prov.: Moa, El Johnson; 1 $, June 1954, 
Zayas & Alayo (IZAC). Santiago de Cuba 
Prov.; Pico Turquino; 3 3, 1 9, June 1963; 
Alayo and Garcia (IZAC, USNM); 1 3, June 
1964, Garcia (IZAC); 2 3, 10-29 June 1936, 
J. Aoulla (USNM). Sierra Maestra, 1000 ft. 
[305 m]; 2 3, 7-21 June 1930, O. Querci 

Host. — Unknown. 

Flight period.— May to August. Most rec- 
ords indicate a May-June emergence, al- 
though this may be only a collecting artifact. 

Distribution.— This species appears to be 
widely distributed over Cuba, occurring at 
rather low elevations to as high as 450 m. 
It may also occur in the Bahamas, but the 
single male examined in the USNM from 
Mangrove Cay, Andros Island, is in too poor 
condition (rubbed and without genitalia) to 

Etymology.— The specific name is de- 
rived from \.2Li\n fuscus (dusky, dark) and 
signatus (mark, stamp) in reference to the 
forewing markings. 

Discussion.— This species, with its white 
forewings heavily streaked with fuscous, ap- 



pears distinct from all other West Indian 
Acrolophus. Consequently, its nearest affin- 
ities remain uncertain, although the male 
genitalia closely resemble a few currently 
unnamed species from the Virgin Islands. 
The forewing pattern can vary consider- 
ably from the most common expression (Fig. 
1) in which a heavy, sinuate streak of dark 
fuscous extends from the basal fourth of the 
costa through the discal cell to the termen. 
The smallest male examined (Fig. 2) from 
Holguin Province possesses the most re- 
duced markings but exhibited genitalia in- 
separable from the typical form (Fig. 1). 

Acrolophus guttatus Davis, 
New Species 
Figs. 4, 11-14 

Adult (Fig. 4). — Length of forewing: 6, 
12.0-12.5 mm. A moderately large species 
with strongly recurved labial palpi in the 
male. The forewing is reddish brown marked 
by 4-5 large, reticulated whitish spots. 

Head: Vestiture densely hairy, brown 
where exposed; frons and vertex largely hid- 
den by labial palpi. Eye hairy; lashes not 
evident. Antenna approximately 0.5 x the 
length of forewing, 56-59 segmented; scape 
brown dorsally, pale buff to white ventrally; 
flagellum laminate, with pale buff to white 
scales dorsally; naked ventrally except for 
dense, very pale, short sensory setae. Labial 
palpus 3 segmented, extremely long in male 
and recurved over thorax to metanotum; 
vestiture brown with lateral portions of dis- 
tal half mostly suffused with pale buff to 

Thorax: Pronotum brown with scattered, 
indistinctly white tipped scales. Tegula dark 
brown. Venter pale buff to white, very hairy. 
Forewing predominantly orange brown with 
4-5 large variegated whitish spots arranged 
as in Fig. 4; each spot mostly cream colored 
with small scattered patches of silvery- white 
scales and a reticulate pattern of smaller or- 
ange scales; fringe brown. Hindwing pale 
gray with fringe only slightly darker. Foreleg 
and midleg brown dorsally, pale buff to 

nearly white ventrally. Hindleg paler, most- 
ly whitish buff with spurs and tarsi brown. 

Abdomen: Pale whitish gray dorsally, 
slightly darker, more buff ventrally. 

Male genitalia: As shown in Figs. 1 1-14. 
Uncus shortly bifid. Gnathos divided into 
a pair of elongate, rounded lobes. Valva di- 
vided less than a third its length at apex into 
a pair of similar, nearly straight lobes. Ae- 
doeagus moderately stout, about two-thirds 
the length of valva, and without comuti. 

Female and immature stages unknown. 

Holotype.— (5. Finca La Ciega, Camaguey 
Province, Cuba; 18 June 1955, P. Alayo, at 
light (IZAC). 

Paratypes.— Cuba: Pinar del Rio Prov- 
ince: Guanahacabibes Peninsula: Cabo Cor- 
rientes thickets: 1 3, 15 May 1956, P. Alayo, 
at light (USNM). 

Host.— Unknown. 

Flight period. — May to June. 

Distribution. — Known only from two 
disjunct localities, at the western tip of Cuba 
and from east-central Cuba. 

Etymology.— The specific name is de- 
rived from the Latin guttatus (spotted, 
speckled) in reference to the conspicuous 
spotted pattern on the forewing. 

Discussion. —The presence of 4 to 5 large, 
reticulated whitish spots on the forewing of 
this species readily distinguishes it from all 
other Acrolophus. The divided valva in the 
male genitalia amply demonstrates its affin- 
ities to the plumifrontellus group. The rel- 
atively straight, unswoUen cucullar lobe and 
the absence of discernible comuti is diag- 
nostic of the species. 

Acrolophus niveipunctatus 


Figs. 5, 15-18 

Acrolophus niveipunctata Walsingham, 
1892: 513; Davis, 1984: 20, no. 113. 

Acrolophus niveipunctatus Walsingham, 
1897: 174. 

Adult (Fig. 5). — Length of forewing, <5, 13- 
14 mm. A moderately large species with 



Figs. 7-14. Male genitalia. 7, Acrolophus fuscisignatus, ventral view. 8, Lateral view. 9, Lateral view of 
valva. 10, Aedoeagus. \\,A. guttatus. ventral view. 12, Lateral view. 13, Lateral view of valva. 14, Aedoeagus. 
All scales = 0.5 mm. 



Figs. 15-22. Male genitalia. 15, Acrolophus niveipunctatus, ventral view. 16, Lateral view. 17, Lateral view 
of valva. 18, Aedoeagus. 19, A. basistriatus, ventral view. 20, Lateral view. 21, Lateral view of valva. 22, 
Aedoeagus. All scales = 0.5 mm. 



Strongly recurved labial palpi in the male. 
The forewing is pale reddish brown with a 
whitish-buff colored anal area possessing a 
bicrenulate anterior margin bordered by an 
interrupted line of broad, white scales. 

Head: Vestiture densely hairy, reddish 
brown where exposed; frons and vertex of 
male largely covered by labial palpi. Eye 
hairy with long lashes. Antenna approxi- 
mately 0.5 X the length of forewing, 60-62 
segmented; scape reddish brown dorsally, 
buff to white ventrally; flagellum laminate, 
with pale buff to white scales dorsally, naked 
ventrally except for dense, very pale, short 
sensory setae. Labial palpus 3 segmented, 
extremely long in male and recurved over 
thorax to metanotum. Vestiture dark red- 
dish brown over basal two-thirds and apex; 
subapical region with white-tipped, pale buff 
scales laterally. 

Thorax: Pronotum generally pale brown 
with a mixture of piliform, dark brown- 
tipped scales and paler, broader, white- 
tipped scales. Tegula reddish brown. Venter 
mostly reddish brown, very hairy. Forewing 
anteriad of CuA dark reddish brown, grad- 
ually fading to pale buff along subterminal 
margin; anal area posteriad of CuA con- 
trastingly pale gray to buff except for dark 
reddish-brown suffusion across base of anal 
area and a semicircular protrusion below 
middle of wing from Cu; margin between 
the two areas continues obliquely to tomus 
along CuA2; also situated disjunctly along 
this margin is a thin line of broad, silvery- 
white scales along base of CuA and more 
distally along CuA2 to tomus but inter- 
rupted medially by the semicircular lobe of 
dark scales; cilia dark reddish brown. 
Hindwing uniformly pale grayish brown ex- 
cept for slightly darker cilia. Foreleg and 
midleg reddish brown dorsally, buff colored 
ventrally. Hindleg much paler, pale buff with 
mixture of indistinctly white-tipped scales; 
spurs and tarsi more brown. 

Abdomen: Pale gray dorsally; reddish 
brown ventrally. 

Male genitalia: As shown in Figs. 15-18. 

Uncus shortly bifid. Gnathos divided into 
a pair of elongate rounded lobes. Valva di- 
vided less than a third its length at apex into 
a pair of straight lobes; ventral lobe (cucul- 
lus) with slender base and slightly enlarged 
apex. Aedoeagus moderately broad, about 
0.75 X the length of valva; a cluster of 10- 
1 2 minute spines present near apex. 

Female and immature stages unknown. 

Type. — Holotype, S. "Museum Staudin- 
ger," present deposition unknown. 

Type locality. — Cuba. 

Host. — Unknown. 

Flight period.— May to June. 

Distribution. — Known only from the 
Sierra Maestra of Santiago de Cuba Prov- 

Material examined. — 3 6. CUBA: Santia- 
go de Cuba Prov.: Honglosongo, Cobre, 
Loma de Gato; 1 <5, 20 June 1952 (IZAC). 
Gran Piedra, Caney; 1 <5, June 1954, Zayas 
and Alayo (IZAC). Gran Piedra Mt., Sierra 
Maestra; 1 5, 30 May 1959, P. Alayo, at 
light (USNM). 

Discussion.— The discovery of the above 
specimens in the Instituto de Zoologia de 
Cuba is significant because it establishes the 
identity and relationship of a previously 
named but otherwise unknown species. Most 
of the uncertainty surrounding A. nivei- 
punctatus is caused by the absence of the 
type specimen, present whereabouts of 
which remains unknown. All specimens 
bearing this name in the collections that I 
have examined were found to be misiden- 
tified. Fortunately, this species possesses a 
rather distinctive wing pattern in which an 
interrupted series of broad white scales are 
most diagnostic. This pattern, as carefully 
described by Walsingham, agrees complete- 
ly with the specimens before me. It should 
be noted that even if the male holotype is 
found, it would still be impossible to deter- 
mine the specific group relationship of niv- 
eipunctatus solely on that specimen, assum- 
ing Walsingham's original statement is 
correct: "Abdomen missing, (a female ab- 
domen is stuck on to this specimen)." 



Fig. 23. Acrolophus fuscisignatus, female genitalia, 
ventral view. Scale = 0.5 mm. 

Acrolophus basistriatus Davis, 
New Species 
Figs. 6, 19-22 

Adult (Fig. 6).— Length of forewing: <5, 1 1- 
12 mm. A moderately large species with 
strongly recurved labial palpi in the male. 
The forewing is reddish brown with a prom- 
inent white streak extending along the base 
of the cubital vein. 

Head: Vestiture densely hairy, reddish 
brown with white-tipped piliform scales 
where exposed; frons and vertex of male 
largely hidden by labial palpi. Eye hairy with 
long lashes. Antenna approximately 0.4 x 
the length of forewing, 59-62 segmented; 
scape reddish brown dorsally, mostly white, 
irrorated with reddish-brown scales ven- 
trally; flagellum laminate, with light brown 
scales dorsally, naked ventrally, except for 

dense, very pale, short sensory setae. Labial 
palpus 3 segmented, extremely long in male, 
recurved over thorax to metanotum. Ves- 
titure similar to head, consisting of dense 
piliform scales of reddish brown with mi- 
nute white tips. 

Thorax: Pronotum similar to head in col- 
or and vestiture. Venter paler in color, pale 
reddish brown to buff. Forewing uniformly 
reddish brown except for a prominent white 
streak extending along the base of CuA; dis- 
tal half of streak tends to diffuse into anal 
area. Cilia consisting of very short, broad 
scales with dull white tips. Hindwing uni- 
formly pale brown. Foreleg reddish brown 
with paler, more wooly vestiture ventrally. 
Midleg similar to foreleg in color. Hindleg 
much paler in color, pale buff with indis- 
tinctly white-tipped scales dorsally; vesti- 
ture mostly white ventrally. 

Abdomen: Densely covered with light 
brown wooly scales dorsally and ventrally; 
usually a concentration of darker brown 
scales along mid- venter. 

Male genitalia: As shown in Figs. 19-22. 
Uncus shortly bifid. Gnathos divided into 
a pair of elongate, rounded lobes. Valva di- 
vided about one-third its length at apex into 
a straight, relatively broad, costal lobe and 
a more slender, strongly curved cucullar 
lobe. Aedoeagus moderately slender, nearly 
as long as valva, and with a central apical 
mass of approximately 12-14 short spines. 

Female and immature stages unknown. 

Holotype.— 3. Cuabal de Gulindo, Valle 
del Yumuri, Matanzas, Cuba; June 1970, P. 
Alayo (IZAC). 

Paratypes. — Same data as holotype, 5 S 

Host. — Unknown. 

Flight period.— June. 

Distribution. — Known only from the type 
locality in Matanzas Province. 

Etymology.— The specific name is de- 
rived from the Labin basis (base, bottom) 
and stria (furrow, line, stripe) in reference 
to the single white streak along the base of 
the forewing cubital vein. 



Discussion.— Acrolophus basistriatus is 
the only Cuban member of the genus with 
a whitish streak along the cubital vein. The 
male genitalia are diagnostic in possessing 
the largest comuti and the most curved cu- 
cullar lobe (of the valva) of any member of 
the plumifrontellus group. 


I am grateful to the Instituto de Zoologia, 
Academia de Ciencias de Cuba, for spon- 
soring my trip to Cuba in 1981 and to the 
Smithsonian Institution for a Fluid Re- 
search Grant that allowed me to participate 
in this program. I am especially thankful to 
Pastor Alayo of the Instituto de Zoologia 
for allowing me to study his material. The 
line drawings were executed by my former 
assistant, Biruta Akerbergs Hansen, and the 
photographs are by Victor Kranz of the 

Smithsonian Photographic Laboratory. Sil- 
ver West of our department typed the final 
draft of the manuscript, which was reviewed 
by J. F. Gates Clarke, and an anonymous 

Literature Cited 

Davis, D.R. 1984. Tineidae, pp. 1 9-24. /« Heppner, 
J. R., ed.. Atlas of Neotropical Lepidoptera, vol. 
2, Checklist, pt. 1 Micropterigoidea to Immoidea. 
W. Junk, The Hague. 

Hasbrouck, F. F. 1964. Moths of the family Acro- 
lophidae in America north of Mexico (Microlep- 
idoptera). Proceedings of the United States Na- 
tional Museum 114(3475): 487-706. 

Walsingham, Lord (Thomas de Grey). 1892. On the 
Micro-Lepidoptera of the West Indies, pp. 511- 
517, 544-545. In Proceedings of the Zoological 
Society of London. 

. 1897. Revision of the West-Indian Micro- 
Lepidoptera, with descriptions of new species, pp. 
1 69-1 75. 7/7 Proceedings of the Zoological Society 
of London. 


89(2), 1987, pp. 284-287 




Weldon L. Burrows 

Consulting Aquatic Biologist, 902 Pennsylvania Ave., St. Albans, West Virginia 25177. 

Abstract. — Nymxih^ and reared imagoes ofAmeletus tarteri, new species, are described. 
The new species is presently known from West Virginia, Virginia, and New York. Mor- 
phological characters of the male imago and nymph of the new species are illustrated and 
compared with the nominally described eastern Nearctic species and with three western 
Nearctic species that possess similar male genitalia. Notes on the ecology and habits of 
the nymph of the new species are included. 

The genus Ameletus was established by 
Eaton (1885). Thirty-two species o{ Ame- 
letus have previously been described from 
North America including six species from 
eastern North America (Edmunds et al., 
1976; Carle, 1978). During an ecological 
survey of streams in central West Virginia, 
nymphs of an apparent new species ofAme- 
letus were discovered. Subsequent rearings 
revealed characters of the male imago that 
differed from those of the known North 
American species o^ Ameletus. Since little 
is known of the biogeography oi Ameletus, 
available type specimens of three western 
species with similar male genitalia, as well 
as five of the six nominal eastern species, 
were examined. The male of the eastern 
species A. walleyi Harper has complex gen- 
italia (see Harper, 1970, figs. 1 and 2) quite 
unlike that of the new species and was not 
examined; the nymph of A. walleyi is un- 

Ameletus tarteri Burrows, 

New Species 

Figs. 1-5 

Male imago (in alcohol). — Body length 
7.5-10.0 mm. Forewing length 8.0-9.5 mm. 

Head brown. Scapes and pedicels of anten- 
nae tan; flagella light purple. Compound eyes 
yellow-green dorsally, tan ventrally. Thorax 
mostly brown with lighter and darker areas 
and streaks. Legs tan with forelegs darker. 
Forewings hyaline; costal and subcostal cells 
suffused with amber; bullae darkly mar- 
gined. Hind wings hyaline; costal cells suf- 
fused with amber. Abdominal tergum 1 
brown; terga 2-10 pale, shading to tan at 
posterior and lateral margins and each with 
submedian dark marks; sterna pale with dark 
marks submedian on 2-8 and in ganglionic 
areas; sternum 9 with gray-brown anterior 
and lateral margins. Genitalia (Figs. 1 and 
2) mostly gray-brown; medial area and an- 
terior margin of subgenital plate lighter, with 
median apical emargination, and with apex 
of mesad apical processes attaining apex of 
first forceps segment; dorsal sclerotization 
extending mesad from bases of penis lobes 
and contiguous mesally; dorsal sclerotized 
bar at fused base of penes; pair of straight, 
acuminate, laterad titilators with 1-3 den- 
ticles along ventral margin of each. Caudal 
filaments tan. 

Female imago (in alcohol). — Body length 
8.0-10.5 mm. Forewing length 9.0-11.5 



Figs. 1-5. Ameletus tarteri. 1, Male genitalia, dorsal, ap (mesad apical process), bs (sclerotized bar at fused 
base of penes). Is (mesad extension of sclerotization from bases of penis lobes), pi (penis lobe), / (titilators, 
hidden). 2, Left penis lobe, lateral, d (denticle), pi (penis lobe), / (titilator). 3, Abdomen of mature nymph, dorsal. 
4, Abdomen of mature nymph, ventral. 5, Left mandible of mature nymph, // (inner incisor), Im (lacinia mobilis), 
mh (molar brush), sg (setal gap), sr (setal row). 

mm. Head tan with black submedian lines 
on vertex. Compound eyes olive-brown 
dorsally, olive ventrally. Abdomen (with 
eggs) orange except segment 1 brown; sterna 
with dark marks submedian on 2-7, 9 and 
in ganglionic areas; subanal plate variable 
but often with anterior and lateral margins 
entire. Remainder similar to male imago. 

Mature nymph (in alcohol).— Body length 
7.0-1 1.5 mm; caudal filaments about 0.5 x 
body length. Head gray-brown; lighter areas 
on frons and mouthparts; female with sub- 
median dark lines on vertex extending to 
lateral ocelli. Antennae tan, shading distally 
to purple. Labrum tan, often with median 
"V" outlined darker. Distal seta of inner 

margin of mandibles (Fig. 5) about 1.5 x 
length of proximal seta of row; length of 
setal row about 2.8 x length of setal gap; gap 
between proximal seta of row and molar 
setal brush about 1.7 x length of proximal 
seta. Thorax tan with lighter and darker 
streaks and areas. Legs pale with gray-tan 
shading; each leg from foreleg progressively 
lighter; tarsi with apical black band. Ab- 
dominal terga (Fig. 3) tan with lighter and 
darker areas; dark submedian marks on 2- 
10; spinules on posterior margins of 3 or 4- 
10; lateral margins of 1 or 2-9 posteriorly 
produced, spinelike on 2 or 3-9. Gills hya- 
line with amber tracheae in fresh specimens; 
sclerotized at ventral margin and near dor- 



sal margin. Abdominal sterna (Fig. 4) pale 
with darker anterior margins; dark median, 
submedian, and sublateral markings vari- 
ably on 3-9. Caudal filaments alternately 
banded light-dark-light-dark from base; 
proximal dark band dark gray with all ar- 
ticulations similar; distal dark band purple- 
gray; each light band tan on cerci, white on 
terminal filament; hairlike setae of proximal 
half of each filament purple-brown, white 
in distal half of each. 

Holotype. — (3 imago. West Virginia, 
Greenbrier Co., Hamrick Run at West Vir- 
ginia Route 39/55 near confluence with 
North Fork of Cherry River, 900 m el. 
(38°13'40"N, 80°24'04"W), 15 June 1983, 
W. L. Burrows. 

Paratypes.— 4 $ imagoes, 3 9 imagoes, 
same data as holotype; 7 nymphs, same data 
as holotype but 13 July 1985; 2$ imagoes, 

3 9 imagoes, Greenbrier Co., Carpenter Run 
at WV Route 39/55 near confluence with 
North Fork of Cherry River, 750 m up- 
stream from Hamrick Run, 15 June 1983, 
W. L. Burrows; 2 6 imagoes, same data but 
9 July 1982; 16 nymphs, same data but 13 
July 1985. All material in alcohol; all adults 
reared with exuviae included. The holotype 
will be deposited at the U.S. National Mu- 
seum of Natural History (USNMNH); para- 
types will be deposited at USNMNH, Ca- 
nadian National Collection, West Virginia 
Benthological Survey (Marshall Universi- 
ty), Purdue University, Rorida A&M Uni- 
versity, and the Academy of Natural Sci- 
ences, Philadelphia. 

Other material examined.— Nymph, New 
York, Chemung Co., McCom Creek, 7 April 
1976, Lamb, Cornell University collection; 

4 nymphs, Virginia, Giles Co., Stony Creek, 
above White Rock Branch, 10 March 1985, 
Burrows, author's collection. 

Etymology. — I am pleased to name the 
new species after Donald C. Tarter, Chair- 
man, Dept. of Biology, Marshall Universi- 
ty. His work, as both student and professor, 
has contributed much to the knowledge of 
the regional aquatic fauna. 

Discussion.— The penes (Figs. 1 and 2) of 
the male imago of A. tarteri resemble those 
of the western species A. cooki Mc- 
Dunnough, /i. imbellis Day, andyl. vemalis 
McDunnough in that each has a pair of 
straight, acuminate, laterad titilators. The 
penes of the eastern species A. cryptostim- 
ulus Carle are somewhat similar, but ex- 
amination of the genitalia of the holotype 
reveals that the titilators are not only nearly 
contiguous as in Carle's (1978) figure, but 
that they are apparently flattened through- 
out their lengths. The male imago of A. tar- 
teri can be distinguished from all known 
Ameletus from North America by the pres- 
ence of denticles (Fig. 2) on the ventral mar- 
gins of the acuminate titilators. 

The nymphal tarsi of all species examined 
have a dark band at both the base and apex 
except for A. tarteri and A. cryptostimulus, 
which have only a dark apical band. Al- 
though the maculation of the dorsal abdo- 
men of the nymph of A. cryptostimulus is 
similar to A. tarteri (Fig. 3), the following 
should reliably distinguish the two species: 
on tergum 7 the dark, submedian, curved 
marks are followed by a median spot in A. 
cryptostimulus but by submedian spots or, 
in many cases, by no such posterad spots in 
A. tarteri; on tergum 10 the dark submedian 
marks are expanded anterolaterally in A. 
tarteri, but not in A. cryptostimulus, to form 
a dark margin extending to the lateral mar- 
gin of the tergum. The labra of A. tarteri 
and A. cryptostimulus are mostly tan, often 
with a darker median "V" in the former and 
nearly always with a dark macula reminis- 
cent of the small form of the Greek letter 
omega in the latter. Spinules occur on the 
posterior margins of abdominal terga 3 or 
4-10 in yi. tarteri, on 5 or 6-10 in yl. cryp- 
tostimulus, and on 1-10 or 2-10 in the other 
seven species of Ameletus examined. Sev- 
eral characters of the inner mandibular setal 
row (Fig. 5), including numbers of setae and 
relative lengths of setae, setal row, and setal 
gap, are useful for distinguishing A. tarteri 
from the other species examined. For ex- 



ample the setal row of the species examined 
is less than 1.5 x the setal gap except in A. 
tarteri (2.8 x), A. ludens Needham (4.5 x), 
A. lineatus Traver (4.8 x), and A. tertius 
McDunnough, whose setal row is continu- 
ous to the molar setal brush with no gap. 

Ecology and habits.— Nymphs o{ A. tar- 
teri were found in rocky first and second 
order streams. These streams typically had 
low pH, low alkalinity, and low specific con- 
ductivity. The immature nymphs were often 
found on horizontal substrate; later instars 
were typically found on nearly vertical faces 
of boulders; pre-emergent nymphs, how- 
ever, were often found congregated on boul- 
der faces angled beyond the perpendicular. 
The nymphs were most efficiently collected 
by sweeping those surfaces with a small 
aquarium net. 


I thank the following people for making 
specimens available for study: D. Azuma 
and D. Otte, the Academy of Natural Sci- 
ences, Philadelphia; N. D. Penny and W. J. 
Pulawski, California Academy of Sciences; 
R. Foottit and J. E. H. Martin, Canadian 
National Collection; J. K. Liebherr, Cornell 

University; R. D. Davie, personal collec- 
tion; O. S. Flint and G. F. Hevel, U.S. Na- 
tional Museum of Natural History; B. C. 
Kondratieff" and J. R. Voshell, Jr., Virginia 
Polytechnic Inst, and State University. I 
thank Dean Adkins, Dept. of Biology, Mar- 
shall University and Jan Hacker, Plant Pest 
Control Div., W.V. Dept. of Agriculture, for 
critically reviewing the manuscript. I thank 
Steve Lawton for illustrating the nymphal 
abdomen and Vicki Crager for help with 
typing. I am grateful to several mayfly work- 
ers, including anonymous reviewers, for the 
generous sharing of their expertise, insight, 
and time. 

Literature Cited 

Carle, F.C. 1978. A new species of >4m£'/£'/M5(Ephem- 

eroptera: Siphlonuridae) from western Virginia. 

Ann. Entomol. Soc. Am. 71: 581-584. 
Eaton, A. E. 1885. A revisional monograph of Recent 

Ephemeridae or mayflies. Trans. Linn. Soc. Lond. 

(2)Zool. 3: 210. 
Edmunds, G. P., Jr., S. L. Jensen, and L. Bemer. 1976. 

The Mayflies of North and Central America. Univ. 

Minn. Press, MinneapoUs, MN. 330 pp. 
Harper, F. 1970. A newspeciesof.-lmf'tow^ (Ephem- 

eroptera: Siphlonuridae) from southern Ontario. 

Can. J. Zool. 48: 603-604. 


89(2), 1987, pp. 288-295 




J. E. McPherson, R. J. Packauskas, and P. P. Korch, III 

Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901; RJP, 
present address: Biological Sciences Group, University of Connecticut, Storrs, Connecticut 

Abstract. — The life history of Pelocoris femoratus (Palisot de Beauvois) was studied in 
southern Illinois, and the immature stages were described. The bug was reared from egg 
to adult in the laboratory. Adults of this univoltine species overwintered in mud and 
detritus at the bottom of their aquatic habitat and became active in early March. Eggs 
were found between late April and mid-May and had been glued singly to leaves of 
Cemtophyllum demersum L. beneath the surface of the water. First instars appeared in 
mid-May followed by marked overlapping of the subsequent instars. Active adults were 
last observed in November. This species was reared on Chaoborus americanus (Johannsen) 
larvae under a 16L:8D photoperiod at 26.7 ± 1.5°C. The incubation period averaged 
17.7 days. Durations of the five subsequent stadia averaged 10.5, 9.7, 11.0, 12.6, and 
19.5 days, respectively. 

The naucorid Pelocoris femoratus (Pali- 
sot de Beauvois) ranges in continental 
United States from New England south to 
Florida and west to the Dakotas (Slater and 
Baranowski, 1978), Kansas, Oklahoma, and 
Texas (Sanderson, 1982). It occurs through- 
out Illinois (Lauck, 1959). 

Scattered notes have been published on 
this bug's field life history. It is predaceous, 
feeding on small moUusks, dragonfly naiads, 
and other aquatic animals (Lauck, 1959; 
Uhler, 1884). It inhabits various lentic (e.g. 
lakes, ponds, pools) (Blatchley, 1926; Bobb, 
1974; Ellis, 1952; Froeschner, 1962; Gon- 
soulin, 1973; Hungerford, 1927 [as P. car- 
olinensis Torre-Bueno; see La Rivers, 1948]; 
Lauck, 1959; Polhemus, 1979; Slater and 
Baranowski, 1978; Torre-Bueno, 1923; 
Wilson, 1958) and the sluggish parts of lotic 
habitats (Gonsoulin, 1973; Polhemus, 1979) 
where it is usually well concealed amidst 

thick growths of aquatic plants (e.g. Alter- 
nanthera, Cham, Lemna, Mwiophyllum, 
Nitella) (Bobb, 1974; Ellis, 1952; Gonsou- 
lin, 1973; Hungerford, 1927; Lauck, 1959; 
Polhemus, 1979; Slater and Baranowski, 
1978; Torre-Bueno, 1903, 1905, 1923; Wil- 
son, 1958). 

Adults overwinter at the bottoms of ponds 
and pools in muck and detritus (Blatchley, 
1926; Bobb, 1974; Uhler, 1884) and emerge 
in spring to feed and reproduce. Oviposition 
begins in spring (Bobb, 1974; Torre-Bueno, 
1 903) and continues at least until the middle 
of the summer (Torre-Bueno, 1903). 
Nymphs have been found during the sum- 
mer (Bobb, 1974; Torre-Bueno, 1903). 
There appears to be only one generation per 
year (Sanderson, 1982) although nymphs 
may be collected in several stages at the 
same time (Sanderson, 1982; Torre-Bueno, 
1903, 1923). 



Pelocoris femoratus has been reared in 
the laboratory under uncontrolled condi- 
tions from egg to adult and the immature 
stages have been briefly described (Hunger- 
ford, 1927; Torre-Bueno, 1903). 

For the past three years (i.e. 1983-1985), 
we have studied the life history of a popu- 
lation of P. femoratus occurring in the La 
Rue-Pine Hills Ecological Area. This area, 
located ca. 30 km northeast of Cape Gi- 
rardeau, Missouri, in the northwest comer 
of Union County, Illinois, is part of the 
Shawnee National Forest. It includes both 
heavily forested areas atop limestone bluffs, 
and moist forests at the base of these bluffs 
that surround La Rue Swamp and Winters 
Pond. These aquatic habitats are continu- 
ous and the naucorid occurs throughout the 
area. Much of the study area is blanketed 
with duckweeds (i.e. Lemna, Spirodela, 
Wolffia, and Wolffiella) along the shoreline. 

This paper presents information on the 
life history and laboratory rearing of P. fe- 
moratus and includes descriptions of the 
immature stages. 

Materials and Methods 

Life history.— The study began in March 
1983, before the bugs emerged from over- 
wintering sites that year. Samples of adults 
and/or nymphs were taken with an aquatic 
net at ca. weekly intervals at six sites along 
the edge of the study area into November 
after all nymphs had disappeared and adult 
activity had markedly decreased. Sampling 
during the following two years was con- 
ducted similarly. All samples were pre- 
served in 75% ethanol and examined in the 
laboratory to accurately determine the de- 
velopmental stages present in each sample. 
Eggs were collected by hand-picking and re- 
turned to the laboratory for incubation to 
confirm their identity when the first instars 
emerged. Occasional collections were made 
also during the winter months to determine 
overwintering stage(s) and sites. Data gath- 
ered during the three years of this study were 

combined to gain a better understanding of 
the annual life cycle. 

Laboratory rearing. — Eighteen adults were 
collected during late March 1985, returned 
to the laboratory, and divided between two 
aquaria (8, 10 adults; no attempt was made 
to sex the individuals). Each aquarium (ca. 
30 X 20.5 X 15 cm) was covered on the 
bottom with aquarium gravel and filled with 
ca. 6 cm of dechlorinated water. Cerato- 
phyllum demersum L. (Ceratophyllaceae) 
was provided as an oviposition site. Adults 
were maintained on larvae of Chaoborus 
americanus (Johannsen) (Diptera: Cha- 
oboridae). Aquaria were cleaned as the water 
became fouled, ca. every 20 days. 

The C. demersum was replaced daily; any 
sections with attached eggs were removed 
and placed in petri dishes. Each dish (ca. 9 
cm diam, 4 cm depth) was covered on the 
bottom with filter paper and filled with ca. 
1 cm of distilled water, which was sufficient 
to keep the eggs submerged. Upon hatching, 
the first instars were also placed in petri 
dishes. Each dish was again covered on the 
bottom with filter paper and filled with ca. 
1 cm of distilled water, more than sufficient 
to cover the bugs. Later instars were also 
provided sufficient water to just keep them 
submerged. Five first instars were placed in 
each dish but further separated as they de- 
veloped through subsequent instars. 

One C. americanus larva was provided 
daily as food for first and second instars, 
increased to two larvae for third and fourth 
instars, and to three for fifth instars. Dishes 
were checked daily for exuviae and prey car- 
casses removed. Clean dishes, paper, and 
water were provided at each molt; water was 
changed more frequently if it became fouled. 

Three pairs of newly emerged F, adults 
were placed in finger bowls (one male and 
one female per bowl) to determine if they 
would reproduce. Each finger bowl (10.5 cm 
ID, 3.5 cm depth) was filled with 2.5 cm of 
dechlorinated water, and C. americanus 
larvae and a section of Elodea canadensis 
Michaux (Hydrocharitaceae) were provided 













25 — 







25 — 












25 — 














Figs. 1, 2. Field life cycle of P. femoratus during 1983-1985 combined seasons in Union Co., Illinois. 1, 
Percent of individuals in each stage per sample. 2, Percent in each sample of total individuals of same stage 
collected during season. For both figures, dashed lines between first and last dates for first instar indicate no 
specimens were found. 



as food and an oviposition site, respectively. 
The finger bowls were cleaned as the water 
became fouled (ca. every 20 days). 

The aquaria, petri dishes, and finger bowls 
were kept in incubators maintained at ca. 
26.7 ± 1.5°Canda 16L:8Dphotoperiod(ca. 
260 ft-c). 

Descriptions of immature stages. — First 
to fifth instars were selected from field sam- 
ples, eggs from those deposited in the lab- 
oratory by field-collected adults; all speci- 
mens had been preserved in 75% ethanol. 
The description of each stage is based on 
ten individuals. Drawings were made with 
the aid of a camera lucida, measurements 
with an ocular micrometer. Dimensions are 
expressed in mm as A' ± SE. 

Results and Discussion 

Life history.— This species overwintered 
as adults at the bottom of the swamp in mud 
and detritus and became active in early 
March (Figs. 1, 2). Eggs (10) were found 
between late April and mid-May but un- 
doubtedly were laid well beyond this period, 
based on the late seasonal appearance of 
first instars. All had been glued singly (i.e. 
not in clusters) to leaves of C. demersum 
beneath the surface of the water. 

The first instars were found from mid- 
May to the third week of August, second 
instars from the third week of May to late 
August, third instars from late May to early 
September, fourth instars from early June 
to early September, and fifth instars from 
mid-June to late September. No active adults 
were found after November. 

This species is apparently univoltine. Our 
conclusion is based in part on the fluctua- 
tions in numbers of the adults and nymphs 
during the season. Although there was 
marked overlapping of the various stages 
and, thus, any particular sample could have 
any combination of individuals (Fig. 1), 
weekly plotting of data for each stage showed 
only one peak for the first, fourth, and fifth 
instars, and adults (i.e. a buildup of new 
generation adults); the second and third in- 

Table 1 . Duration (in days) of each immature stage 
of P. /^A/iorarw^ under controlled laboratory conditions. 





.v± SE 

Mean Age 




17.7 ± 0.06 



1st instar 



10.5 ± 0.11 


2nd instar 



9.7 ± 0.12 


3rd instar 



11.0 ± 0.13 


4th instar 



12.6 ± 0.16 


5th instar 



19.5 ± 0.18 



were laid. 

Stars show more fluctuations in seasonal 
numbers but this may be a reflection of smafl 
sample sizes which, by chance, was not ap- 
parent with the first instars (Fig. 2). 

Laboratory rearing.— As in the field, eggs 
were glued singly to the leaves of C. de- 
mersum beneath the surface of the water. 
They were white at oviposition but dark- 
ened to yellowish with brown markings dur- 
ing maturation. The incubation period av- 
eraged 17.7 days (Table 1). 

The first instar emerged through a slit in 
the cephalic end of the egg. It was whitish 
at this time but soon darkened to its normal 
color (see description). It usually fed on its 
first C americanus larva within one day. 

The first through fifth stadia averaged 
10.5, 9.7, 11.0, 12.6, and 19.5 days, re- 
spectively (Table 1). The total develop- 
mental period averaged 81.0 days. 

F, adults maintained in the laboratory for 
six weeks never reproduced. This further 
supports our conclusion that this species is 
univoltine in southern Illinois. 

Descriptions of Immature Stages 

Egg (Fig. 3). -Length, 1.37 ± 0.01; width, 
0.77 ± 0.01. Eggs laid singly and glued to 
aquatic vegetation, each egg white at ovi- 
position, but turning yellowish with brown 
markings during maturation; chorion with 
primarily irregular hexagonal pattern; mi- 
cropylar plug at cephalic end. 



Table 2. Measurements (mm)^ of P. femoratus instars. 


1st Instar 

2nd Inslar 

3rd Instar 

4lh Instar 

5th Instar 

Body length 

2.55 ± 






± 0.05 





± 0.09 

Body width" 

1.58 ± 






± 0.02 





± 0.05 

Width at eyes 

0.90 ± 






± 0.01 





± 0.02 


0.49 ± 






± 0.02 





± 0.01 

Head length^ 

0.60 ± 






± 0.02 





± 0.04 

Pronotal length*^ 

0.25 ± 






± 0.01 





± 0.02 

Mesonotal length"^^ 

0.20 ± 






± 0.01 





± 0.03 

Metanotal length^ 

0.31 ± 






± 0.01 





± 0.01 

Leg lengths: 


0.60 ± 






± 0.01 





± 0.02 


0.34 ± 






± 0.01 





± 0.01 


0.21 ± 






± 0.01 





± 0.01 


0.50 ± 






± 0.01 





± 0.02 


0.41 ± 






± 0.01 





± 0.01 


0.25 ± 






± 0.01 





± 0.01 


0.64 ± 






± 0.01 





± 0.03 


0.71 ± 






± 0.02 





± 0.03 


0.39 ± 






± 0.01 





± 0.01 

X ± SE; SE values rounding off to less than 0.005 listed as 0.00. 
' Measured across metanotum. 
Measured along midline. 

Nymphal instars.— The first instar is de- 
scribed in detail, but only major changes 
that have occurred from previous instars are 
described for subsequent instars. Length is 
measured from tip of tylus to tip of abdo- 
men; width across metanotum for lst-4th 
instars, across mesonotal wing pads at level 
of metanotum for 5th. Additional measure- 
ments are given in Table 2. 

First instar (Fig. 4).-Length, 2.55 ± 0.05; 
width, 1.58 ± 0.02. Body elongate oval, 
general appearance dorsoventrally flat- 
tened, greatest width at metathorax; dor- 
sally convex; ventrally, flattened to slightly 
concave laterally, convex medially; brown- 
ish with yellowish white markings dorsally, 
yellowish white with brown markings ven- 

Head broadly triangular, recessed into 
prothorax to level of eyes; anterior margin 
convex, continuous with lateral margins of 
prothorax; posterior margin arcuate, often 
subtruncate medially. Dorsally, head yel- 

lowish white with posteriorly directed, 
broad, brown, anchor-shaped mark medi- 
ally, and thin yellowish line originating near 
midpoint of inner margin of each eye, both 
lines continuing posteriorly and converging 
medially near back of head to form broad 
V-shaped pattern and giving rise posteriorly 
to short line that is continuous with mid- 
dorsal yellow line of thorax. Eyes red, synth- 
lipsis (taken near anterior margin of eyes) 
ca. 2.5 X width of one eye. Antennae yel- 
lowish, 3-segmented, segment 2 ca. 2x 
length of segment 1 and 0.7 x length of seg- 
ment 3. Beak yellowish, broadly triangular, 
3-segmented, segment 2 ca. 3x length of 
segment 1 and ca. 1.5 x length of segment 3. 
Thoracic nota generally brownish with 
yellowish or whitish markings, more whit- 
ish laterally, these lateral areas usually lack- 
ing brown spots. Anterior margin of prono- 
tum concave; posterior margin nearly 
straight. Mesonotum shortest of thoracic 
nota, ca. 0.8 x length of pronotum along 



midline, posterior margin straight medially, 
arcuate laterally; mesonotal wing pads ev- 
ident. Metanotum longest of the thoracic 
nota, ca. 1 .6 x length of mesonotum along 
midline; posterior margin nearly straight; 
ratio of mesonotal wing pad to metanotum 
along lateral edge ca. 3:4. Metanotum often 
overlapping abdominal segment 1 and much 
of 2. 

Prothoracic leg raptorial, yellowish, fre- 
quently with brownish streak on outer mar- 
gin of femur. Procoxa ca. 2 x length of tro- 
chanter and 0.5 X length of femur. Profemur 
flattened laterally, markedly tapering dis- 
tally, basally ca. 4 x width distally; ventrally 
with 2 contiguous rows of pegs. Protibia and 
tarsus much narrower than femur, together 
equal to length of femur ventrally. Tibia 
with ventral surface forming shallow groove, 
this groove extending onto tarsus; pegs of 
femur approximating edge of groove of tibia 
and tarsus when these segments are ap- 
posed; tarsus 1 -segmented with single mi- 
nute claw. 

Meso- and metathoracic legs yellowish; 
all leg segments on metathorax longer than 
respective segments on mesothorax. Me- 
socoxa elongate, ca. 2 x length of trochanter 
and 0.8 X length of femur. Mesotarsus 
2-segmented, segment 1 ca. 0.2 x length of 
2, the 2 claws of equal length. Metacoxa and 
trochanter resembling those of mesothorac- 
ic leg in shape and relative proportions, 
fringed with long hairs on outer and inner 
margins, respectively. Meso- and metatib- 
iae and tarsi furnished with row of swim- 
ming hairs on outer surface, most developed 
on hindlegs. 

Abdomen, dorsally, brown with yellow- 
ish white markings; ventrally, yellowish 
white, covered by long hairs and greatly 
convex in middle Vy, 7 pairs of spiracles 
evident but minute, 1st pair more medially 
placed. Paired ostioles of scent glands pres- 
ent dorsally between segments 3-4. 

Second instar (Fig. 5). — Length, 3.39 ± 
0.01; width, 2.13 ± 0.03. Body widest at 
metanotum and 3rd abdominal segment. 

Synthlipsis ca. 2 x width of one eye. An- 
tennae with segment 2 ca. 3 x length of seg- 
ment 1 and 0.6 X length of segment 3. 

Thoracic nota medially generally brown- 
ish with yellow markings or yellowish with 
brown markings, brown markings may be 
much reduced; lateral whitish areas of 
pronotum now often with few brown spots. 
Pronotum subequal to metanotum in length 
along midline, both longer than meso- 
notum. Posterior margin of mesonotum 
slightly arcuate medially, clearly arcuate lat- 
erally; ratio of mesonotal wing pad to meta- 
notum along lateral edge ca. 1:1. Double 
row of pegs of profemur reduced to single 
row except at base, area formerly occupied 
by remainder of 2nd peg row now occupied 
by short setae. Row of swimming hairs pres- 
ent on outer surface of mesotibia and tarsus, 
and on both inner and outer surfaces of 
metatibia and tarsus. Spiracles now more 

Third instar (Fig. 6). — Length, 4.74 ± 
0.05; width, 2.97 ± 0.02. Antenna with seg- 
ment 2 ca. 2 X length of 1 and 0.5 x length 
of 3. Thoracic nota medially often with 
brown markings reduced or absent; lateral 
whitish areas of pronotum now with several 
brown spots. Pronotum longest of thoracic 
nota along m idline, slightly exceeding 
metanotum and much exceeding meso- 
notum. Mesonotum clearly arcuate medi- 
ally and laterally; ratio of wing pad to meta- 
notum along lateral edge ca. 4:3. Single row 
of pegs on profemur now surrounded on 
each side by setae, peg row may be incom- 

Fourth instar (Fig. 7). — Length, 6.24 ± 
0.08; width, 3.93 ± 0.06. Lateral area of 
pronotum now yellowish. Pronotum longest 
of thoracic nota, much exceeding either 
meso- or metanotum along midline. Ratio 
of mesonotal wing pad to metanotum along 
lateral edge ca. 2:1. Single row of pegs on 
profemur now incomplete, area covered by 
numerous setae. 

Fifth instar (Fig. 8). -Length, 8.17 ± 0.09; 
width, 5.33 ± 0.05. Wing pads of meso- 



1.0 mm 

Figs. 3-8. Immature stages of P. femoral us. 3, Egg. 4, First instar. 5, Second instar. 6, Third instar. 7, Fourth 
instar. 8, Fifth instar. 



and metanota (latter present but covered by 
those of mesonotum) just extending to ab- 
dominal segment 3 laterally. Single row of 
pegs on profemur now almost absent, nu- 
merous setae present. 


We thank S. L. Keffer and T. E. Vogt, 
Department of Zoology, SIU-C, for their 
help with various aspects of this project, and 
Karen A. Schmitt, Scientific Photography 
and Illustration Facility, SIU-C, for the final 
illustrations and photographs of the various 
figures. We thank also R. H. Mohlenbrock, 
Department of Botany, SIU-C, for identi- 
fying the species oi Ceratophyllum and Elo- 
dea mentioned in this study. 

Literature Cited 

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Eastern North America with Especial Reference 
to the Faunas of Indiana and Florida. Nature Publ. 
Co., Indianapolis. 1116 pp. 

Bobb, M. L. 1974. The insects of Virginia: No. 7. 
The aquatic and semi-aquatic Hemiptera of Vir- 
ginia. Va. Polytech. Inst. State Univ. Res. Div. 
Bull. 87: 1-195. 

Ellis, L. L. 1952. The aquatic Hemiptera of south- 
eastern Louisiana (exclusive of the Corixidae). Am. 
Midi. Nat. 48: 302-329. 

Froeschner, R. C. 1962. Contributions to a synopsis 
of the Hemiptera of Missouri, Part V. Hydro- 
metridae, Gerridae, Veliidae, Saldidae, Ochteri- 
dae, Gelastocoridae, Naucoridae, Belostomatidae, 
Nepidae, Notonectidae, Pleidae, Corixidae. Am. 
Midi. Nat. 67: 208-240. 

Gonsoulin, G. J. 1973. Seven families of aquatic and 

semiaquatic Hemiptera in Louisiana. Entomol. 
News 84: 83-88. 

Hungerford, H. B. 1927. Life history of the creeping 
water bug, Pelocoris carolinensis Bueno (Naucor- 
idae). Bull. Brooklyn Entomol. Soc. 22: 77-83. 

La Rivers, I. 1948. A new species of PWocorw from 
Nevada, with notes on the genus in the United 
States (Hemiptera: Naucoridae). Ann. Entomol. 
Soc. Am. 41: 371-376. 

Lauck, D. R. 1959. The taxonomy and bionomics of 
the aquatic Hemiptera of Illinois. M.S. Thesis, 
University of Illinois, Urbana. 

Polhemus, J. T. 1979. Family Naucoridae/creeping 
water bugs, saucer bugs, pp. 131-138. In Menke, 
A. S., ed.. The Semiaquatic and Aquatic Hemip- 
tera of California (Heteroptera: Hemiptera). Calif 
Insect Surv. Bull. 21: 1-166. 

Sanderson, M. W. 1982. Aquatic and semiaquatic 
Heteroptera, pp. 6.1-6.94. In Brigham, A. R. et 
al., eds.. Aquatic Insects and Oligochaetes of North 
and South Carolina. Midwest Aquatic Enterprises, 
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Slater, J. A. and R. M. Baranowski. 1978. How To 
Know the True Bugs (Hemiptera— Heteroptera). 
Wm. C. Brown Co., Dubuque, Iowa. 256 pp. 

Torre-Bueno, J. R. de la. 1903. Brief notes toward 
the life history of Pelocoris femorata Pal. B. with 
a few remarks on habits. J. N.Y. Entomol. Soc. 
11: 166-173. 

. 1905. Practical and popular entomology- 
No. 4. Notes on collecting, preserving and rearing 
aquatic Hemiptera. Can. Entomol. 37: 137-142. 

. 1923. Family Naucoridae, pp. 402-404. In 

Britton, W. E., ed., Guide to the insects of Con- 
necticut. Part IV. The Hemiptera or sucking in- 
sects of Connecticut. Conn. State Geol. Nat. Hist. 
Surv. Bull. 34: 1-807. 

Uhler, P.R. 1884. Order VI -Hemiptera, pp. 204- 
296. In Kingsley, J. S., ed.. The Standard Natural 
History. Vol. II. Crustacea and Insects. S. E. Cas- 
sino and Company, Boston. 555 pp. 

Wilson, C. A. 1958. Aquatic and semiaquatic He- 
miptera of Mississippi. Tulane Stud. Zool. 6: 115- 


89(2), 1987, pp. 296-302 


Barry C. Poulton and Kenneth W. Stewart 

Department of Biological Sciences, North Texas State University, Denton, Texas 76203. 

Abstract.— Three new species of Plecoptera representing three families are described 
from the Ozark-Ouachita Mountain region: Allocapnia oribata (Capniidae), Alloperla 
caddo (Chloroperlidae), and Isoperla szczytkoi (Perlodidae). Morphological descriptions, 
illustrations, biological notes, and comparisons with other regional species are provided. 
All stages of Isoperla szczytkoi are described. 

Stark et al. (1983) and Ernst et al. (1986) 
have recently described new Alloperla and 
Neoperla species from the Ozark-Ouachita 
Mountain region of central North America. 
During our ongoing study of the regional 
Plecoptera fauna, three additional species 
representing three families were discovered. 
The following descriptions and morpholog- 
ical terms follow those of Ross and Ricker 
(1971), Surdick (1981), and Szczytko and 
Stewart (1976, 1979, 1984). 

Allocapnia oribata Poulton and 

Stewart, New Species 

Figs. 1-3 

Male. — Micropterous. Color dark brown 
in alcohol. Length of body 5.5-6.5 mm; 
wings reaching 6th tergum. Seventh tergum 
without a dorsal process, but with a distinct 
rounded membranous area connected to 
posterior margin (Fig. 2). Eighth tergum with 
anterior margin sloping abruptly upward 
from base forming a sclerotized process, ap- 
pearing as rounded knob in lateral view (Fig. 
1), and wide, shallow, and U-shaped in dor- 
sal view (Fig. 2). Eighth tergum process sub- 
conical in posterior view (Fig. 3). Upper 
limb of epiproct narrow, and slightly wider 
apically in dorsal view (Fig. 2), with sub- 
equal proximal and distal sections, and cla- 

vate apically in lateral view (Fig. 1). Upper 
and lower epiproct limbs subequal in length 
in lateral view, and lower limb spatulate and 
abruptly curved downward at tip (Fig. 1). 

Female. —Unknown. 

Material examined. — Holotype $, Arkan- 
sas, Searcy Co., Middle Fork Little Red 
River, Hwy 65 @ Shirley, 6-1-1985, B. C. 
Poulton; two additional <5 in poor condition, 
same locality and date, B. C. Poulton; 1 
paratype <5, Arkansas, Van Buren Co., Ar- 
chey Creek, Hwy 254, 12.1 km NE Rupert, 
6-1-1985, B. C. Poulton. Holotype depos- 
ited at United States National Museum, 
paratype deposited at North Texas State 
University museum. 

Etymology.— The species name is de- 
rived from the Greek root oribat-, meaning 
"mountain roaming." 

Diagnosis.— The species represents the 
first discovery of a new Allocapnia from the 
Ozarks since A. warreni Ross and Yama- 
moto (1966). It is not clear which Ross and 
Ricker (1971) group this species belongs to, 
since: 1) the epiproct upper limb and the 
shape of the eighth dorsal process in dorsal 
view are typical of the pygmaea group; how- 
ever the process is unnotched, and 2) the 
arcuate elevated posterior ridge of the eighth 
dorsal process is characteristic of the recta 



(^b B 

Figs. 1-3. Allocapnia oribata. 1, Lateral terminalia, <5. 2, Dorsal terminalia, 6. 3, 
process, <3. 4-7, AUoperla caddo. 4, Lateral (A) and anterior (B) view, <5 epiproct tip. 
Ventral, 9 subgenital plate. 7, Lateral, 2 abdominal segments 7-9. 

Posterior view, 8th dorsal 
5, Dorsal terminalia, 6. 6. 

group (Figs. 1, 3). Even though A. oribata 
is similar to A. malverna Ross of the recta 
group, neither it nor A. malverna have the 
thin blade-Uke epiproct upper limb typical 
of A. recta (Claassen) and other members 
of that group. 

Biological notes.— The type males were 
collected on bridges from permanent, 4th 
order streams which have rock-rubble sub- 
strate. Large numbers of .4. mohri Ross and 
Ricker, A. rickeri Prison, and A. granulata 
(Claassen) were also collected at these lo- 

calities. Females of these species exhibit 
considerable variation in subgenital plate 
form; therefore we were unable to discern 
the female of .4. oribata. 

AUoperla caddo Poulton and 

Stewart, New Species 

Figs. 4-7 

Male.— Macropterous. General color am- 
ber to white in alcohol. Forewing length 6- 
7 mm; body length 5-7 mm. Abdominal 
dorsal stripe absent. Epiproct tip ca. 2x 



longer than wide; lateral margins conver- 
gent near base (Fig. 5); anterior margin 
sharply upturned in lateral view and with a 
prominent anterior knob visible in dorsal 
view (Figs. 4A, 5). Epiproct tip with ventral 
groove visible in front view (Fig. 4B). Lat- 
eral and dorsal abdominal setae present, with 
longer brushes of setae on segments 7-9, 
and dorsally on segment 10. Posterior mar- 
gin of abdominal pleura 8 and 9 with 4-8 
brown spinules. Basal cereal segments with 
thick setae ca. 1-2 x segment width. Ae- 
deagus membranous. 

Female. — Macropterous. Color similar to 
male. Forewing length 6-7 mm; body length 
5-7 mm. Subgenital plate little produced, 
with posterior margin rounded in ventral 
view (Fig. 6), and slightly protruding ven- 
trad in lateral view (Fig. 7). Cereal and ab- 
dominal setae similar to male. Fine setae 
on subgenital plate (Figs. 6, 7) ca. '/2 length 
of lateral setae. Vagina membranous. 

Material examined. — Holotype 3, allo- 
type 9, 7 paratype 3, and 4 paratype 2, Ar- 
kansas, Garland Co., Middle Fork Saline 
River, Hwy 7 at Iron Springs Rec. Area, 
6-VI-1984, B. C. Pouhon; 4 paratype <5 and 
1 paratype 9, Arkansas, Perr>' Co., Dry Fork 
Creek, Hwy 7, 14.3 km S. of Mollis,' 6- VI- 
1 984, B. C. Poulton; 4 paratype 5, Arkansas, 
Perry Co., Bear Creek, Hw>' 7, 4.4 km SE 
of Mollis, 12-V-1985, B. C. Poulton. 

Etymology.— This species is named after 
the Caddo mound builders, an American 
Indian tribe that inhabited parts of the 
Ouachita Mountains from 750-1200 AD. 

Diagnosis. — Recent descriptions and an 
illustrated key to Ozark-Ouachita Alloperla 
were provided by Stark et al. (1983), in- 
cluding the new species A. ouachita Stark 
and Stewart, collected in the same region of 
the Ouachita Mountains as A. caddo. The 
unique saddle-shaped epiproct tip of.-l. cad- 
do (Fig. 4) easily separates it from A. leon- 
arda Ricker and A. hamata Surdick. The 
two last species have flat, blade-like epi- 
proct tips, with either lateral serrations or 

points (Surdick, 1981). Alloperla ouachita 
and A. caudal a Frison have broadened epi- 
proct tips, but the lateral horns of .4. ouachi- 
ta and the appressed hairs of A. caudata 
(Mitchcock, 1974; Stark etal., 1983) are ab- 
sent in A. caddo. The membranous basal 
lobes of the epiproct cowl in A. caddo are 
relatively smaller than those of .4. ouachita 
and A. hamata (Stark et al., 1983). A. caddo 
represents the only Ozark-Ouachita Allo- 
perla with an evenly rounded female 
subgenital plate; the other species all possess 
pointed subgenital plates similar to those 
described by Surdick (1981) for .4. hamata 
and A. furcula Surdick. 

Biological notes.— The type localities are 
all in the Ouachita Mountains and comprise 
first or second order, rock-rubble, intermit- 
tent streams. Adults were collected by 
sweeping riparian vegetation. 

Isoperla szczytkoi Poulton and 
Stewart, New Species 

Figs. 8-20 

Male. — Macropterous. Forewing length 
10-11 mm, body length 9-10 mm. Body 
color yellow in life, light brown to amber in 
alcohol, with darker pattern on head, 
pronotum, and thorax. Mead with brown 
circular patch enclosing light spot, con- 
necting median ocellus with anterior frons; 
lateral and median ocelli connected with an 
inverted U-shaped dark band with lighter 
peripheral patches, posterior ones extending 
to near eyes and back to occiput (Fig. 1 5). 
Pronotum with median light stripe; disks 
medium brown with dark brown rugosities. 
Pronotum stripe continues through meso- 
notum; rest of mesonotum and metanotum 
light brown. Cerci and antennae medium 
brown. Legs light brown, tibiae darker than 
femora, both with darker brown bands near 
their articulation. Cereal segments with short 
plumose setae and a single, long, dark brown 
posterioventral seta on each segment. Wings 
light brown with dark brown veins. Eighth 
sternum with vesicle 1.5-2 x wide as long 



,1 ',^SV'i< 

• •••','» • • • . . •/ 

# • A • « 

Figs. 8-10. hoperla szczytkoi egg. 8, 700 x . 9, 200 x . 10, 700 ; 


» » » T ;, 


(Fig. 13), with setae present along lateral 
and anterior margins. Paraprocts medium 
length, curving upward and inward, narrow- 
ing to a sharp point, and overlapping pos- 
terior margin of 1 0th tergum (Fig. 1 1 ). Ninth 
pleura with a single brown spot. Aedeagus 
membranous with bulb-like posteriodorsal 
section 1.5-2 x wider than aedeagal base; 
posterior portion with a row of 6-7 scler- 
otized, digited golden brown teeth sur- 
rounded by an unsclerotized triangular area; 
blunt spinulae present on elevated circular 
region posteriodorsal to sclerotized digited 
teeth; ventral portion of aedeagus with el- 
evated region containing sparse blunt spi- 
nulae, and a round patch with anterior half 
containing stout brown spinulae of medium 
density (Fig. 13). Basal portion of aedeagus 
with circular bands containing stout, blunt 
spinulae and short, rounded spinulae; lat- 
eral regions with patches of sclerotized 
punctures and sharp golden brown spinulae 
(Fig. 12). Posterodorsal tip of aedeagus 
without spinulae; separate membranous 
dorsal lobe between cerci unsclerotized and 
without spinulae (Fig. 12). 

Female. — Macropterous. Forewing length 
11-12 mm, body length 9-11 mm. Body 
coloration and external morphology similar 
to male. Subgenital plate broadly rounded 

and produced posteriorly to anterior '/3 of 
9th sternum (Fig. 14). Vagina membranous. 
Nymph. — Body length 10-13 mm. Gen- 
eral color brown in alcohol. Abdomen with 
2 faint dorsal longitudinal stripes with light 
borders, appearing slightly diagonal on each 
segment, with a median light blotch be- 
tween them; a row of 6-8 dark spots, 2 al- 
ways positioned dorsally inside median light 
blotch, and others dorsolaterally (Fig. 16). 
Head pattern with dark inverted U-shaped 
band connecting ocelli, with light area be- 
tween lateral ocelli, extending to posterior 
portion of head, and light oval area anterior 
to median ocellus (Fig. 1 7). Pronotum vari- 
able with median longitudinal brown band 
surrounded by lighter borders; irregular dark 
markings on discs and light lateral margins 
(Fig. 17). Lacinia with 2 teeth; subapical 
tooth '/3-'/2 length of apical tooth and par- 
tially hidden behind 1 of 2 incomplete rows 
of stout hairs that continue along entire 
length of inner margin (Fig. 20). Labrum 
with median swelling (Fig. 1 7). Right man- 
dible deeply cleft, with 5 short, stout apical 
teeth, the outer 2 with inner serrated ridges. 
Inner margin of mandible with a dense brush 
of long setae (Fig. 1 8). Glossae produced 
and broadly pointed upward at apex; para- 
glossae slender, curved, with length ca. 1.5- 



Figs. 1 1-20. Isoperla szczytkoi. 1 1, Dorsal, S segment 10. 12, Lateral terminalia, 3. 13, Ventral terminalia, 
(5. 14, Ventral terminalia, 9. 15, Head and pronotum, adult S. 16, Abdominal tergites, nymph. 17, Head and 
pronotum, nymph. 18, right mandible, nymph. 19, Nymphal labium. 20, Right lacinia. nymph. 



2 X width of base (Fig. 1 9). Posterior margin 
of abdominal segments with continuous 
fringe of hairs. Cereal segments with a dor- 
sal row of long setae, and whorles of short 
hairs on posterior margins. 

Egg.— Outline oval, cross section circular. 
Length 0.35 mm; width 0.27 mm. Collar 
absent. Chorion sculptured with numerous 
small punctations, and a few raised knobs, 
possibly marking a micropylar ring (Figs. 

Material examined. — Holotype $ and al- 
lotype 9 (reared), Arkansas, Logan Co., Gut- 
ter Rock Creek, 33 km SE of Paris on side 
road near Hwy 309, 20-IV-1985, B. C. 
Poulton; 6 paratype S and 3 paratype 9 
(reared), and 6 nymphs, same locality and 
date, B. C. Poulton; 1 paratype $ (reared), 
and 4 nymphs, same locality, 13-IV-1985, 
B. C. Poulton. Holotype, allotype, and 2 
nymphs deposited at USNM; paratypes de- 
posited at NTSU Museum. 

Etymology.— This species is named in 
honor of Stanley W. Szczytko, who is pres- 
ently working on the revision of eastern 
Nearctic Isoperla and has greatly contrib- 
uted to knowledge of Isoperla in North 

Diagnosis. — During our intensive sam- 
pling in the Ozark and Ouachita Mountain 
region, our collections have revealed a total 
of nine Isoperla species. The most recent 
descriptions of regional Isoperla are pro- 
vided by Szczytko and Stewart (1976) and 
Stark and Stewart (1973). Stanley W. 
Szczytko (personal communication), has in- 
dicated that /. szczytkoi belongs to the /. 
decepta Frison-/. mohri Frison complex. 
Common characters of this complex include 
the following: 1) outer nymphal mandibular 
teeth deeply cleft and serrated, 2) nymphal 
lacinial shelf reduced, 3) nymphal subapical 
tooth of lacinia reduced or absent, 4) 
nymphal head pattern dark, 5) aedeagus with 
sclerotized patch-like process bearing teeth, 
and 6) ovum without a collar. Based on these 
characters, /. szczytkoi most closely resem- 

bles /. decepta, another Ozark-Ouachita 
mountain species. Nymphs of these two 
species have a similar head pattern, but /. 
decepta lacks a median pronotal stripe and 
has a much weaker apical lacinial tooth. 
Adult /. decepta lack a contrasting head pat- 
tern as in /. szczytkoi (Fig. 1 5). 

Biological notes.— This species is known 
only from the type locality, an intermittent 
first-order stream with large rubble substra- 
tum, that flows down the north side of Mag- 
azine Mountain, having the highest eleva- 
tion in the Ozark-Ouachita region. 


We thank S. W. Szczytko for the SEM 
photography and helpful suggestions in di- 
agnosing the new Isoperla species. This study 
was supported in part by National Science 
Foundation Grant BSR 8308422 and the 
Faculty Research Fund of North Texas State 

Literature Cited 

Ernst, M. R., B. C. Poulton, and K. W. Stewart. 1 986. 
Neoperla (Plecoptera: Perlidae) of the southern 
Ozark and Ouachita mountain region, and two 
new species o{ Neoperla. Ann. Entomol. Soc. Am. 
79: 645-661. 

Hitchcock, S. W. 1974. Guide to the insects of Con- 
necticut, Pt. VII. The Plecoptera or stoneflies of 
Connecticut. Bull. State Geol. Nat. Hist. Surv. 
Conn. 107: 1-262. 

Ross, H. H. and W. E. Ricker. 1971. The classifi- 
cation, evolution, and dispersal of the winter 
stonefly genus AUocapnia. 111. Biol. Monogr. 45: 
166 pp. 

Ross, H. H. and T. Yamamoto. 1966. Two new sister 
species of the winter stonefly genus AUocapnia 
(Plecoptera: Capniidae). Entomol. News. 77: 265- 

Stark, B. P. and K. W. Stewart. 1973. New species 
and descriptions of stoneflies (Plecoptera) from 
Oklahoma. Entomol. News. 84: 192-197. 

Stark, B. P., K. W. Stewart, and J. Feminella. 1983. 
New records and descriptions of Alloperla (Ple- 
coptera: Chloroperlidae) from the Ozark-Ouachita 
region. Entomol. News. 94: 55-59. 

Surdick, R. F. 1981. New Nearctic Chloroperlidae 
(Plecoptera). Great Basin Nat. 41: 349-359. 

Szczytko, S. W. and K. W. Stewart. 1976. Three new 



species of nearctic Isoperla (Plecoptera). Great Ba- 
sin Nat. 36:211-220. 

— . 1979. The genus Isoperla (Plecoptera) of 
western North America; holomorphology and sys- 
tematics, and a new stonefly genus Cascadopeiia. 
Mem. Am. Entomol. Soc. 32: 1-120. 

— . 1984. Descriptions of Ca///per/a Banks, ^/cA:- 
era Jewett, and two new western nearctic Isoperla 
species (Plecoptera: Perlodidae). Ann. Entomol. 
Soc. Am. 77: 251-263. 

Book Review 

89(2), 1987, p. 302 

Foundations for a National Biological 
Survey. Edited by K. C. Kim and Lloyd 
Knutson. Association of Systematic Col- 
lections, Washington, D.C. 1986, xii + 
215 pp. $18.00. 

This important book is primarily a result 
of a symposium presented at the annual 
meeting of the Association of Systematic 
Collections held in May of 1 985 in Victoria, 
British Columbia. A total of twenty-four 
contributors examine and comment on vir- 
tually all aspects of the formation, funding, 
and perceived results and effects of a 
National Biological Survey. After a brief 
foreword by E. O. Wilson and a preface by 
the editors, the heart of the book is pre- 
sented in six sections. An introduction by 
K. C. Kim and Lloyd Knutson addresses 
the scientific bases for such a Survey. This 
introduction provides an excellent descrip- 
tion of what a National Biological Survey 
might encompass and what might be ex- 
pected as a direct result of such a Survey. 
The introduction also provides in brief form 
and with well-chosen words what is ex- 
panded upon in the following sections of 
the volume. 

The second section consists of papers 
summarizing the relations of such a Survey 
on ecological and environmental consider- 
ations. Especially cogent are the articles on 
the relation of systematics to long-range 
ecologic research and the role of a National 
Biological Survey on environmental pro- 
tection, food production and plant protec- 
tion. This is followed by a section on bio- 
logical survey information in which the form 
of the data, how it is to be managed and 
how and to whom it is to be disseminated, 
is discussed. A fourth section dealing with 

legislative and historical perspectives for a 
National Biological Survey examines both 
federal and state legislation of the past with 
comments on possible future legislation. A 
fifth section describes the ambitious biolog- 
ical survey programs of Australia and Can- 
ada and the results of these surveys, and 
discusses the formation of these surveys and 
their possible use as models for such a sur- 
vey in this country. 

A final brief section consists of two arti- 
cles. First, a brief article by L. L Nevling, 
Jr. summarizes the conference and provides 
a list of nine statements in which the con- 
tributors, representing highly diverse view- 
points, were in agreement. Second, R. M. 
West and W. D. Duckworth list five rec- 
ommendations distilled from the presen- 
tations of the participants. A final brief epi- 
logue by the editors completes the volume. 

This reviewer found considerable varia- 
tion in quality of the presentations, as is 
usually the case in symposium volumes. 
However, the editors did a remarkable job 
in finding excellent speakers and writers and 
a fine job of editing and introducing the 
results. Some ideas expressed in this book 
are of general interest, as well as of interest 
in relation to a possible National Biological 
Survey. For instance, this reviewer found 
Barry Chemoff s discussion of the three main 
relationships between ecology and system- 
atics to be especially well expressed and 
thought provoking. The volume, in short, 
is well worth examining and should be on 
the shelf of anyone doing systematic, eco- 
logic, or survey studies involving the fauna 
or flora of North America. 

Wallace E. LaBerge, Illinois Natural His- 
tory Survey, Champaign, Illinois 61821. 


89(2), 1987, pp. 303-321 




James S. Miller 

Postdoctoral Fellow, Department of Entomology, Smithsonian Institution, Washington, 
D.C. 20560. 

Abstract.— T\it genus Phryganidia Packard (Lepidoptera: Dioptidae) is revised and a 
new species, Phryganidia chihuahua, is described. Following a brief review of the Diop- 
tidae, the three species of Phryganidia and the placement of the genus are discussed. Six 
autapomorphies for Phryganidia are identified. Relevant morphological characters are 
illustrated and distributional data are presented. 

When I began studying the systematics of 
the Dioptidae, John Bums presented me 
with a copy of a poem he had written en- 
titled "In Temperate Spring" (1986). The 
last two lines read, "Like the California oak- 
moth, I'm a mindless Phryganidia.''' As the 
ironies of life would have it, I now find my- 
self revising the genus. 

To most North American entomologists, 
Phryganidia californica, the California Oak 
Moth, is the only member of the obscure 
family Dioptidae with which they are fa- 
miliar. However, the family contains nearly 
400 species (Bryk, 1930), all of which, with 
the exception of P. californica, occur from 
Mexico south to Uruguay and Argentina. 
Most dioptids are relatively small, light- 
bodied, brightly colored moths that appear 
to mimic members of groups as disparate 
as the Pericopinae and Lithosiinae in the 
Arctiidae, and the Ithomiinae in the Nym- 
phalidae. The 39 other dioptid genera will 
be the subject of future publications; this 
paper concerns the species of Phryganidia. 
Because the Dioptidae are so poorly under- 
stood, it is necessary to review briefly the 
taxonomic history of the family and de- 

scribe some of the morphological characters 
that previous authors have discussed. 

Ever since the family was erected by 
Walker in 1865, its phylogenetic position 
has been controversial. After examining the 
larva and pupa of Phryganidia californica, 
it was Packard's (1895) opinion that the 
Dioptidae were closely related to the Geo- 
metridae. Dyar (1896) suggested that the 
group was associated with the Pericopinae. 
Forbes (1916) argued that dioptids belonged 
to the Noctuid/Arctiid lineage within the 
Noctuoidea, but subsequently (1922) fol- 
lowed Fracker (1915) and Mosher ( 1 9 1 6) in 
moving them to a position close to the No- 
todontidae, a relationship with which all re- 
searchers have since agreed. Not until Minet 
(1983) was a significant change in the phy- 
logenetic position of the Dioptidae pro- 
posed. Minet argued that the group should 
be recognized as a tribe, the Dioptini, within 
the Notodontinae, but he was uncertain 
about the tribe's affinities within that 
subfamily. Minet based his proposal on a 
single character complex, the structure of 
the last pair of larval prolegs. 

Several character complexes have figured 



prominently in theories concerning dioptid 
relationships. The structure of the dioptid 
tympanum indicates a close relationship 
with the Notodontidae. The Noctuoidea 
{sensii Hodges et al., 1983; Watson et al., 
1980) can be defined by the presence of a 
thoracic tympanum (Richards, 1933; Brock, 
1971; Minet, 1983), and only in the Diop- 
tidae and Notodontidae is the tympanal 
membrane directed ventrally (Richards, 
1933; Sick, 1940; Kiriakoff, 1950a, b, c, 
1963; Minet, 1983). Variation in tympanal 
structure within the Dioptidae has also re- 
ceived considerable attention (Sick, 1940; 
Kiriakoff, 1950a). 

Since Fracker (1915), many authors have 
recognized a similarity between the larvae 
of Phryganidia and those of notodontids. 
Minet (1983) described the anal prolegs of 
dioptids as being 'reduced or absent' and 
claimed that this apomorphic character state 
is shared with the Notodontinae. I have ex- 
amined larval specimens representing the 
dioptid genera Phaeochlaena, Phryganidia, 
Zunacetha, and Josia. The anal prolegs are 
sometimes elongate (e.g., Phaeochlaena 
gyon, figured in Bastelberger, 1 908), but they 
are not "reduced or absent" as Minet sug- 
gested. His hypothesis is therefore in doubt. 

Wing venation has provided the majority 
of characters previous authors have used to 
define dioptid genera (Prout, 1918; Hering, 
1925; Bryk, 1930), but offers little infor- 
mation concerning higher-level relation- 
ships. All species are "trifid" in the fore and 
hind wings (M, being closer to Mi than to 
M3), a plesiomorphic character state shared 
with the Notodontidae and many lepidop- 
teran families (see Forbes, 1948). Two char- 
acters that are frequently observed in the 
Notodontidae, the presence of an "acces- 
sory cell" in the fore wing (Forbes, 1942, 
1 948), and Sc in the hind wing being parallel 
with R for two-thirds the length of the discal 
cell (Franclemont, 1970), do not occur in 

As this discussion indicates, the phylo- 
genetic position of the Dioptidae remains 

unresolved. Available evidence strongly 
supports a relationship with the Notodon- 
tidae but is inadequate to allow a more pre- 
cise hypothesis. Until more becomes known 
it seems better to retain family-level status 
for the group. 

No autapomorphies for the Dioptidae are 
known, but a character of the larvae appears 
promising. The larval skin of all species I 
have examined is covered with microscopic 
cuticular projections, described by Fracker 
(1915) for Phryganidia californica. This 
condition has also been termed "minutely 
rugose" (Forbes, 1942) or "shagreened" 
(Peterson, 1965). The presence of these cu- 
ticular projections may ultimately prove to 
be a synapomorphy for the Dioptidae, but 
it is important that the immatures of very 
few species have been collected. 

Phryganidia has historically included only 
two species. Packard (1864) described the 
genus in the subfamily Psychinae (Bom- 
bycidae), with P. californica as the type 
species. He chose the name to note conver- 
gent similarities between these moths and 
trichopteran species in the Phryganidae. 
Phryganidia was subsequently referred to 
the Zygaenidae (Stretch, 1872), and finally 
to the Dioptidae (Kirby, 1892). Like P. cal- 
ifornica, P. naxa was originally placed with 
the psychids. Druce (1885) described P. 
naxa in the psychid genus Typhonia Bruand. 
Prout (1918) united both species in the 
Dioptidae as members of Phryganidia but 
expressed doubt as to the correct affinity of 
P. naxa. The only other taxon that has been 
named, fasciata, was described by Hering 
(1928) as a form of P. naxa. This paper 
treats these three taxa and describes a new 
species, P. chihuahua. 


Morphological terminology of the head 
and antennae follows Forbes (1923) and 
Hodges (1971). Names for the structures of 
the genitalia follow Klots ( 1 970) and Hodges 
(1971). However, the homology of lepidop- 
teran genitalic structures needs careful study. 



For example, it is uncertain whether the 
terms socii, uncus, and transtilla used here 
to describe dioptids are homologous with 
structures in other groups to which the same 
names have been applied. I will use the terms 
while recognizing that future research is es- 
sential. The color guide of Smithe (1975) 
was followed as closely as possible in de- 
scriptions of vestiture and wings. 

Genitalia and adomens were prepared by 
soaking the entire abdomen in 10% KOH 
for 12-15 hours. Specimens were placed in 
20% ethanol and the abdomen was cut lon- 
gitudinally along the pleural membrane on 
the right side. The terminalia were then sep- 
arated from the rest of the abdomen and 
scales and soft tissues were removed with a 
brush. The aedoeagus was removed, and the 
vesica was inflated using fluid pressure from 
a syringe inserted in the anterior (proximal) 
end of the aedoeagus. The bursa was simi- 
larly inflated using a syringe inserted in the 
ostium. Preparations were then stained for 
10 to 20 seconds in Chlorazol Black (ICN 
Pharmaceuticals), followed by one hour in 
Safranin O (Wards's Natural Science Estab- 
lishment), and mounted on slides. Line 
drawings were made using a camera lucida 
attached to a Wild M5 dissecting micro- 
scope. Lateral views of genitalia were drawn 
from specimens in ethanol. Drawings of pal- 
pi, legs, anal views of male genitalia, ae- 
doeagi, and tergites/stemites were made 
from specimens mounted on slides in Can- 
ada Balsam. 

Scanning electron micrographs were tak- 
en with a Cambridge Scan 1 00 microscope. 
Specimens were mounted on stubs and 
sputter-coated with gold/palladium. Before 
being mounted, antennae were cleaned by 
rinsing them for several minutes in 95% eth- 
anol. The thorax of dried specimens was 
carefully brushed free of scales for micro- 
graphs of the tympanum. Preserved pupae 
were removed from ethanol and air dried 
before being mounted and sputter-coated. 

Acronyms for museums are as follows: 
American Museum of Natural History, New 

York, (AMNH); British Museum (Natural 
History), London, (BMNH); Canadian Na- 
tional Collections, Ottawa, (CNC); Cornell 
University Insect Collections, Ithaca, (CU); 
Carnegie Museum of Natural History, Pitts- 
burgh, (CMNH); Essig Museum of Ento- 
mology, Berkeley (UCB); Natural History 
Museum of Los Angeles County, Los An- 
geles, (LACM); Museum of Comparative 
Zoology, Cambridge, (MCZ); National Mu- 
seum of Natural History, Washington, 

Genus Phryganidia Packard 

Phryganidia Packard, 1864: 348 [type 
species: Phryganidia californica Packard 
by monotypy]. 

Diagnosis. — Small to medium-sized 
moths, fore wing length 1 2 to 22 mm. Most- 
ly light brown to gray-brown. 

kdwW.— Head: Labial palpi upturned to 
just above clypeus, segment 2 slightly longer 
than segment 1 (Fig. 1). Antennae of male 
widely bipectinate (Fig. 4), those of female 
narrowly bipectinate (Fig. 5). 

Thorax: Pleural region and tegulae cov- 
ered with long, hair-liked scales. Epiphysis 
long and slender (Fig. 2), reaching to distal 
end of tibia or beyond. Fore wing (Fig. 3) 
with R, arising from discal cell, other radials 
in the pattern R2 + [(R3 + R4) + R5]; M3 and 
CuA, stalked; posterior angle of discal cell 
oblique. Hind wing with M3 and CuA, 
stalked, posterior angle of discal cell oblique. 
Tympanal membrane not enclosed by met- 
epimeron (Figs. 6 and 7). 

Male abdomen (Figs. 10, 15, 18,20):Ter- 
gite VIII arched dorsally, anterior margin 
slightly emarginate; posterior margin weak- 
ened medially, with numerous membra- 
nous folds and 2 lateral excavations. Pos- 
terior margin of stemite VIII excavated 
medially, with a pair of sclerotized lateral 
pockets; a single broad, blunt apodeme on 
anterior margin. Genitalia with uncus broad 
and hood-like, ventral surface covered with 
a mat of short setae; socii elbowed, with 



Figs. 1,2. 1, Right labial palpus of Phryganidia californica male in lateral view (California, AMNH slide 
no. JSM 1 10; scale line = 0.5 mm). 2, Left fore leg of Phryganidia chihuahua male in lateral view (Mexico, 
CNC slide no. JSM 115, scale line = 1.0 mm). 

short. Spine-like setae; anal tube kinked dis- 
tally; valvae with a small lateral patch of 
androconial scales, a short process at dorsal 
margin of apex; sacculus small; arms of 
transtilla narrow, meeting above aedoeagus 
anteriorly. Aedoeagus short, with a ventral 
tooth and small dorsal sclerite distally; vesi- 
ca with 15 to 25 spine-like comuti. 

Female abdomen (Figs. 11, 19): Genitalia 
with ostium simple; ductus bursae short; 
corpus bursae with small spicules at base 
and a pair of small signa; posterior margin 
of tergite VIII with a pair of long, slender 
projections. Posterior margin of stetnite VII 
excavated medially. 

Immatures. — Larvae with skin sha- 
greened; anal prolegs not elongate, crochets 
present. Pupae with hook-shaped setae on 
dorsum of abdominal segments 7-10 and 
on cremaster (Figs. 8 and 9). 

Natural history.— Within the genus only 
the immatures of P. californica are known. 
The larvae feed on many oak species (Tietz, 
1972), often causing severe defoliation 

(Brown and Eads, 1965). Its primary host 
is Quercus agrifolia (Volney et al., 1983; 
Puttick, 1986). Largely as a result of its eco- 
nomic importance, P. californica has been 
the subject of ecological (Harville, 1955), 
physiological (Volney et al., 1983; Puttick, 
1 986), and bio-control (Milstead et al., 1 980) 
research. Hochberg and Volney ( 1 984) found 
evidence of a female produced pheromone. 
The parasites of P. californica have been 
studied by Young (1982, and references 

Apomorphies. — Of the morphological 
characters described above, the following 
appear to be autapomorphies for Phrygan- 
idia: 1) Epiphysis long and slender; 2) Pos- 
terior margin of male tergite VIII with nu- 
merous folds and two lateral excavations; 
3) Posterior margin of male stemite VIII 
with a pair of sclerotized, lateral pockets; 4) 
Uncus broad and hood-like; 5) anal tube 
with a distal kink; and 6) Posterior margin 
of female tergite VIII with a pair of long, 
slender projections. 



Fig. 3. Fore and hind wing venation of Phryganidia chihuahua (Mexico, CNC slide no. JSM 1 14). A, anal 
vein; C, costal vein; CuA, cubito-anal vein; M, medial vein; R, radial vein; Rs, radial sector; Sc, subcostal vein. 

Key to the Species of 
Phryganidia Packard 

Ground color of fore wings gray-brown, hind 

wings charcoal gray P. naxa Druce 

Ground color of fore and hind wings concol- 
orous 2 

Ground color light brown; tegulae buff- yellow; 

fore wing length 1 2 to 18 mm 

P. californica Packard 

Ground color gray-brown; tegulae gray-brown; 

fore wing length 19 to 22 mm 

P. chihuahua, new species 



Phryganidia califomica Packard 
Figs. 1, 4-14 

Phryganidia californica Packard, 1864: 348. 

Diagnosis. — Fore wing length 12 to 18 
mm. Ground color lighter than other species 
in the genus. 

Head: Scales on first two segments of la- 
bial palpus buff-yellow, third segment light 
gray-brown; front tan to light gray-brown. 
Antennae black ventrally, light gray-brown 
scales dorsally. 

Thorax: Legs, pleural region, and dorsum 
gray-brown; patagia buff to light brown; teg- 
ulae buff-yellow. Wings translucent, light 
brown; dorsal surface of male fore wing with 
a cream colored fascia beyond discal cell, 
fascia fainter below. 

Abdomen: Light brown. Male terminalia 
as in Fig. 10; stemite VIII with a large, 
membranous "window"; a small patch of 
androconial scales on lateral surface of val- 
vae. Female terminalia as in Fig. 1 1 ; stemite 
VII with short, lateral processes on anal 
margin, largely membranous medially; ter- 
gite VIII with a membranous region me- 
dially; corpus bursae with two small patches 
of spicules at base and two indistinct, spi- 
culate signa. 

Type material. — The male lectotype, 
herein designated, bears three labels: "San 
Mateo, California, [no date], A. Agassiz"; a 
red label stating "MCZ Type 23388"; and 
an "A.S. Packard Type" label. It is in fair 
condition and is in the MCZ. A second male 
with the same label data is here designated 
as a paralectotype. 

Other specimens examined. — 64 6, 32 9 
(USNM); 82 3, 21 9 (AMNH); 32 6, 4 9 
(CMNH); 9 pupae, 25 larvae (USNM). Dis- 
sected 5 3, 5 9. 

Distribution.— Oregon. Two localities in 
Lane County (Wickman and Kline, 1985). 
California. Happy Camp, Siskiyou County 
(CMNH); Overwintering from southern 
Mendocino County south to San Diego, in- 
land only in the Riverside area (Fig. 14). 
Also recorded from the Channel Islands 

(Anacapa, Santa Cruz, and Santa Rosa Is- 
lands), and Catalina Island. A single spec- 
imen in the USNM labeled "Cn[Canyon] 
de las Cruces, Baja California, Mexico." Two 
specimens (one from the AMNH and one 
from the LACM) are recorded as being from 

Phryganidia chihuahua J. Miller, 

New Species 
Figs. 2, 3, 15-17 

Diagnosis. — Fore wing length 19 to 22 
mm. This is the largest of the three Phry- 
ganidia species. It is darker than most spec- 
imens of P. califomica, the species with 
which it is superficially most similar. Fe- 
male specimens are not known. 

Head: Scales on first segment of labial 
palpus light yellow, second and third seg- 
ments gray-brown; front light yellow, gray- 
brown at outer margins; antennae black 
ventrally, dark gray-brown scales dorsally. 

Thorax: Legs, pleural region, dorsum, and 
tegulae gray-brown; patagia light yellow. 
Wings gray-brown, translucent, without 
markings either above or below; (a single 
specimen with a faint fascia beyond discal 

Abdomen: Gray-brown. Male terminalia 
as in Fig. 15; anterior apodeme of stemite 
VIII broad; valvae narrow, a small patch of 
androconial scales on lateral surface. 

Type material.— The holotype male (Fig. 
1 6) bears two labels. The first has the data: 
"Mexico. Chi[huahua], Mesa del Huracan, 
108°15'30°4', 7400', VIII-21-25-1964, J. E. 
H. Martin." The second is a "holotype" la- 
bel with red lettering. The specimen is in 
excellent condition and is deposited in the 
Canadian National Collections, Ottawa. 
Paratypes: 9 6, all with the same label data 
as the holotype. Dissected 3 $. 

Other specimens examined. — Mexico. 
Durango, El Salto, 28 mi E, 8000', VII- 
22-64, J. Powell (1 3, UCB). 

Distribution (Fig. 1 7).— Known only from 
two localities; the type-locality, and El Sal- 
to, Durango. 



Figs. 4-9. Scanning electron micrographs of Phryganidia californica. 4, Male antenna (ventral view, scale 
line = 100 txm). 5, Female antenna (ventral view, scale line = 100 tixn). 6, Meso- and metathoracic segments 
plus first two abdominal segments (anterior at left, scale line = 0.5 mm). A 1, abdominal tergite 1; A2, abdominal 
tergite 2; C, metacoxa; E, metepimeron; M2, mesomeron; M3, metameron; S, metepistemum; SL, metascutellum; 
ST2, mesoscutum, T, tympanum. 7, Metathoracic and first abdominal segments (anterior at left, scale line = 
250 jum). E, metepimeron; TM, tympanal membrane. 8, Terminal segments of pupa (dorsal view, scale line = 
0.5 mm). C, cremaster; H, hook-shaped setae on segment 10. 9, Hook-shaped setae on segment 10 of pupa 
(scale line = 100 nm). 



Fig. 10. Male terminalia of Phryganidia californica (California, AMNH slide no. JSM 1 10; scale line = 1.0 
mm). 10a, Genitalia in anal view with aedoeagus removed. A, anal tube; P, apical process; S, socius; Sc, sacculus; 
T, transtilla; U, uncus. 10b, Aedoeagus in lateral view (anterior at left). 10c, Tergite VIII in dorsal view. lOd, 
Stemite VIII in ventral view. 





Fig. 1 1 . Female terminalia of Phryganidia californica (California, USNM slide no. 29171; scale line = 1 .0 
mm). 1 la. Genitalia in lateral view (anterior at left). 1 lb, Tergite VIII in dorsal view. 1 Ic, Stemite VII in ventral 

Etymology.— A noun in apposition. The 
name comes from the type locality. 

KQxmLr\i.s.— Phryganidia chihuahua and 
P. californica are more similar in habitus 
than either is to P. naxa, but without fe- 
males of P. chihuahua there is insufficient 
character information to resolve relation- 
ships within the genus. 

Phrvganidia naxa Druce 
Figs. 18-26 

Typhonia naxa Druce, 1885: 229. 
Phryganidia naxa: Prout, 1918: 412. 

Diagnosis. — Fore wing length 12 to 18 
mm. The fore wings are narrower and the 

hind wings less full than in the other Phry- 
ganidia species. In addition, the wings are 
not concolorous and are only slightly trans- 
lucent (not at all in males). 

Head: Scales on labial palpus and front 
uniformly gray. Antennae black ventrally, 
dark gray scales dorsally. 

Thorax: Legs, pleural region, patagia, 
dorsum, and tegulae gray. Males (Figs. 21, 
23-26) with fore wing ground color gray- 
brown, ventral surface with a yellow fascia 
beyond discal cell; some color forms (fas- 
ciata of Hering, Fig. 21) with an equivalent 
fascia on dorsal surface as well. Males with 
hind wings charcoal gray above and below. 



Figs. 12, 13. Phryganidia californica (dorsal view). 12, Male; 13, Female. 

Females with fore wings gray-brown, hind 
wings charcoal gray; slightly translucent; 
without markings either above or below. A 
single male specimen from Guatemala (Figs. 
20, 25, 26) with a light yellow longitudinal 
stripe on ventral surface of fore wing from 
base stopping short of fascia; a longitudinal 
orange-yellow stripe on both surfaces of hind 
wing from base stopping short of outer mar- 
gin; fringe scales on anal margin yellow. 

Abdomen: Gray, lighter ventrally. Male 
genitalia as in Figs. 1 8 and 20; costa of valve 
expanded; uncus bulbous. Female genitalia 
as in Fig. 19; corpus bursae with a sclero- 
tized region and two patches of spicules at 
base; two more or less distinct, spiculate 

Type material.— The female lectotype 
(Fig. 22), herein designated, is labeled 
"Guatemala City, 5000 feet, [no date], 
Champion." It bears two additional labels: 
"B.C. A. Lep. Het. Typhonia naxa," and 
"Godman-Salvin Coll. 98.-40." A third 
handwritten label by D. Goodger reads 
"probable syntype of naxa Druce." It is in 
good condition and is in the British Mu- 
seum (Natural History). A female with the 
data: "S. Geronimo, [no date]. Champion," 
and bearing a blue, circular "syntype" label 
is here designated as a paralectotype. It has 
no abdomen and is in rather poor condition. 

Other specimens examined. — Guatema- 
la. [18]87-88, Conradt, det. Martin Hering 

(1 9, BMNH); [no date], B. Bruckner S.G., 
(form "fasciata") det. Martin Hering (1 6, 
BMNH); Guatemala City, Sept. & Nov. [no 
year], Schaus and Barnes (2 3, USNM); Vol- 
can de Santa Maria, Nov. [no year], Schaus 
and Barnes (1 9, USNM). Mexico. Nuevo 
Leon, Chipinque Mesa, 4300', IX- 19-20- 
1975 (at light), J. Powell, J. Chemsak, and 
T. Friedlander ( 1 4 <5, 4 9, UCB); Nuevo Leon, 
4 mi W Iturbide, 5500', IX-25-1975 (black- 
light trap), J. Powell, J. Chemsak, T. Fried- 
lander (1 (5, UCB); Nuevo Leon, 6.4 km W 
Iturbide, 24°44'N-99°56'W, 1800 M, VII- 
16-79, D.C. Darling (1 <5, CMNH). Dis- 
sected 6 3, 3 9. 

Distribution (Fig. 17). — Known from 
Nuevo Leon, Mexico and Guatemala. I have 
been unable to verify a locality reported by 
Prout (1918) for Panama. Collecting in 
higher elevation sites between the Mexican 
and Guatemalan localities will likely yield 
additional specimens. 

Remarks.— The wing pattern of P. naxa 
is extremely variable. The specimen in Fig. 
2 1 is indistinguishable from one figured by 
Hering (1928) as ""^ovm fasciata.''' I found 
no genitalic differences between specimens 
with either this color pattern, or that of the 
specimen in Figs. 23 and 24, in which there 
is only a ventral fascia. The specimen most 
distinct from others, placed in the USNM 
collection as ''form fasciata,'' is shown in 
Figs. 20, 25 and 26. It is essentially identical 




Department of Entomology and Parasitology 

Fig. 14. Overwintering range of Phryganidia californica. 

in male genitalic morphology to other spec- 
imens of P. naxa dissected (compare Figs. 
1 8 and 20). The color forms are sympatric 
at both localities but never differ morpho- 
Although the number of available speci- 

mens was small, morphological differences 
between Mexican and Guatemalan speci- 
mens were observed: 1) The vestiture is sil- 
ver-gray in Mexican specimens and brown- 
gray in those from Guatemala. 2) The male 
socii are somewhat more sharply elbowed 



Fig. 15. Male terminalia of Phryganidia chihuahua (Mexico, CNC slide no. JSM 1 12; scale line = 1.0 mm). 
15a, Genitalia in anal view with aedoeagus removed. 15b, Aedoeagus in lateral view (anterior at left). 15c, 
Tergite VIII in dorsal view. 1 5d, Stemite VIII in ventral view. 

in Mexican specimens than in Guatemalan 
specimens. 3) In females from Guatemala 
M, is short- stalked with R2_5, whereas it 
arises from the discal cell in examples from 

Mexico. Rather than recognize these as dis- 
tinct species, I have chosen to wait and see 
whether collecting in localities between 
Nuevo Leon and Guatemala yields additional 



Fig. 16. Phryganidia chihuahua holotype 
(Mexico, dorsal view). 


specimens. Their discovery may indicate that 
morphological differences between the 
northern and southern populations form 
continuous clines. 


Most of the character states described for 
Phryganidia are shared with species in one 
or more additional dioptid genera. Some are 
widespread in the family. The combination 
of wing venational characters found in 
Phryganidia, including the arrangement of 
the radial veins and stalking of M3 and CuA, 
in the fore and hind wings, is characteristic 
of over 270 dioptid species belonging to 14 
genera (approximately 68% of the family). 
This venational pattern does not occur else- 
where in the Noctuoidea and provides per- 
haps the most convenient means for im- 
mediately recognizing the majority of species 
in the Dioptidae. The only large genus with 
different venation is Myonia (57 species), in 

Fig. 17. Geographical distributions of P. chihuahua (A) and P. naxa (■). 




Fig. 18. Male terminalia q{ Phryganidia naxa (Guatemala, USNM slide no. 29174; scale line = 1.0 mm). 
18a, Genitalia in anal view with aedoeagus removed. 18b, Aedoeagus in lateral view (anterior at left). 18c, 
Genitalia in lateral view (anterior at left). 18d, Tergite VIII in dorsal view. 18e, Stemite VIII in ventral view. 



Fig. 19. Female terminalia of Phryganidia naxa (Guatemala, USNM slide no. 29176; scale line = 1.0 mm). 
19a, Genitalia in lateral view (anterior at left). 19b, Tergite VIII in dorsal view. 19c, Stemite VII in ventral 

which R. and M3 of the fore wing arise from 
the discal cell. 

Sick (1940) and Kiriakoff (1950a) argued 
that the tympanum of Phryganidia is ple- 
siomorphic within the Dioptidae. In diop- 
tids with a derived tympanal configuration 
such as Josia and related genera, the meta- 
thoracic epimeron forms a "pocket" that 
almost completely encloses the tympanal 
membrane (figures in Sick and Kiriakoff). 

The hook-shaped setae on the abdomen 
of Phryganidia pupae were first noted by 
Mosher (1916: plate XXVII, fig. 116). I ob- 
served identical structures on the abdomen 

of pupae in other dioptids, including Zu- 
nacetha annulata and Tithraustes demades. 
The complete taxonomic distribution of this 
and other pupal traits will provide invalu- 
able phylogenetic information. 

The labial palpi in many dioptid genera 
are upturned to the dorsal portion of the 
front (sometimes far beyond), and segment 
2 in these cases is extremely long. The pres- 
ence of relatively short palpi in Phryganidia 
is likely a plesiomorphic character state. 

In many Neotropical notodontid genera 
the sacculus of the male valve is greatly ex- 
panded, consisting of a series of "pleats" 



Fig. 20. Male terminalia of Phryganidia naxa '"form fasciata" (Guatemala, USNM slide no. 29175; scale 
line = 1.0 mm). 20a, Genitalia in anal view with aedoeagus removed. 20b, Aedoeagus in lateral view (anterior 
at left). 20c, Tergite VIII in dorsal view. 20d, Stemite VIII in ventral view. 

which are associated with long androconial 
scales (Forbes, 1942; Barth, 1955). Similar 
valvae occur throughout the Dioptidae. The 
absence of a pleated sacculus in Phryganidia 
may be a derived state within the family. 
The transtilla does not join medially in the 
notodontids I have examined whereas it does 
in virtually all dioptids. It is not known 
whether the character state in dioptids is 
plesiomorphic or apomorphic relative to the 

Notodontidae. There is variation in the 
structure of the transtilla within the Diop- 
tidae as well. 

Prout ( 1 9 1 8) and Hering (1925) suggested 
that Phryganidia is structurally most similar 
to species in the genus Polypoetes. I ob- 
served female genitalic similarities between 
members of these two genera. However, my 
research suggests that Polypoetes is not 
monophyletic. A complete understanding 










Figs. 21-26. Phryganidia naxa. 21, P. «axa male similar to form fasciata of Hering (Mexico, UCB; dorsal 
view). 22, P. naxa female lectotype (Guatemala, BMNH; dorsal view). 23, P. naxa male (Mexico, UCB; dorsal 
view). 24, Same specimen (ventral view). 25, P. naxa ""tormfasciata" male (Guatemala, USNM; dorsal view; 
Genitalia, Fig. 20). 26, Same specimen (ventral view). 

of Phryganidia's relationship to the other 
genera will require cladistic analyses, in- 
cluding a thorough investigation of char- 
acter state distributions throughout the 
Dioptidae and Notodontidae. 


I am extremely grateful to J. Donahue 
(LACM), who first noticed the specimens of 
P. chihuahua in the collection at the CNC. 
He also provided valuable distributional 

data for P. califomica. J. Powell was par- 
ticularly helpful in providing the map of 
California, as well as distributional data for 
P. califomica and several references on its 
biology. J. Donahue, B. Poole, S. Weller, J. 
Franclemont, J. Rawlins, and R. Gagne 
kindly offered comments on the manu- 
script. The photographs were taken by V. 
Krantz (USNM). I thank the following in- 
dividuals and their institutions for the loan 
of material: F. Rindge (AMNH), A. Watson 



(BMNH), D. Lafontaine (CNC), J. Rawlins 
(CMNH), D. Bowers (MCZ), J. Powell 
(UCB), J. Liebherr (CU), B. Poole (SEL- 
USDA at USNM). This research was sup- 
ported by a postdoctoral fellowship from 
the Smithsonian Institution. 

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Brock, J. P. 197 1 . A contribution towards an under- 
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Brown, L. R. and C. O. Eads. 1965. California Oak 
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Bryk F. 1930. Dioptidae. In Strand, E., ed., Lepi- 
dopterorum Catalogus (42). Junk, Berlin. 65 pp. 

Bums, J. M. 1986. In temperate spring. BioScience 
36: 294. 

Druce, H. 1 885. Insecta, Lepidoptera-Heterocera, Vol. 
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Dyar, H. G. 1896. On the probable origin of the 
Pericopidae: Composia fidelissima. J. N.Y. Ento- 
mol. Soc. 4: 68-70. 

Forbes, W. T. M. 1916. On the tympanum of certain 
Lepidoptera. Psyche 23: 183-192. 

. 1922. The position of the Dioptidae (Lepi- 
doptera). J. N.Y. Entomol. Soc. 30: 71. 

. 1923. Lepidoptera of New York and neigh- 
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. 1942. Family Dioptidae, pp. 318-322. The 

Lepidoptera of Barro Colorado Island, Panama. 
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. 1948. Family Notodontidae, pp. 203-237. 

Lepidoptera of New York and Neighboring States, 
Part 2. Cornell Agric. Exp. Memoir 274. 263 pp. 

Fracker, S. B. 1915. The classification of lepidopter- 
ous larvae. 111. Biol. Monogr. 2: 1-161. 

Franclemont, J. G. 1970. Dioptidae, pp. 9-11. In 
Dominick, R. B. et al., eds., Perspectus of the Moths 
of America North of Mexico. E. W. Classey Lim- 
ited and the Wedge Entomological Research 
Foundation, London. 

Harville, J. P. 1955. Ecology and population dynam- 
ics of the California Oak Moth Phryganidia cali- 
fornica Packard (Lepidoptera: Dioptidae). Mi- 
croentomology 20: 83-166. 

Hering,E. M. 1925. Dioptidae, pp. 501-534. /^Seitz, 
A., ed.. Die Gross-schmetterlinge der Erde, Vol. 
6. Alfred Kemen, Stuttgart. 

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Zoologischen Staatsmuseum Berlin. Dtsch. Ento- 
mol. Z. "Iris" (Dresden). 42: 268-282. 

Hochberg, M. E. and W. J. A. Volney. 1984. A sex 
pheromone in the California Oakworm Phrygan- 
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Hodges, R. W. 1971. The Moths of America North 
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Bombyces. Gumey and Jackson, London. 95 1 pp. 

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Lepidoptera and taxonomy of the order. J. Lepid. 
Soc. 17: 1-6. 

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89(2), 1987, pp. 322-324 


Alexander D. Huryn 

Department of Entomology, University of Georgia, Athens, Georgia 30602. 

Abstract— Notiphila (Notiphila) kentensis n. sp. is described from a freshwater marsh 
near Kent, Ohio. Notiphila kentensis is a member of the adjusta species group and is most 
closely related to Notiphila mathisi. 

While examining a series of Notiphila spp. 
collected from a marsh near Kent State Uni- 
versity, Portage County, Ohio (Todd, 1985), 
I encountered an undescribed species of the 
subgenus Notiphila, Notiphila (N.) kentensis 
n. sp. Specimens were collected using de- 
tergent pan traps placed in an area domi- 
nated by the emergent macrophyte, Niiphar 
luteum (L.) Sibthorp & Smith (Todd, 1 985). 
Other members oi Notiphila, e.g. Notiphila 
(N.) bella Loew, Notiphila (N.) mathisi Hu- 
ryn, and Notiphila (N.) theonae Huryn, have 
also been collected almost exclusively in as- 
sociation with foliage and flowers of Nuphar 
(personal observation; Huryn, 1984; Todd, 
1985). Although use of Nuphar for ovipo- 
sition and resting sites has been reported in 
the literature, association of the larvae of 
Notiphila with the roots of these plants has 
not been reported (cf Mathis, 1979; Van 
Der Velde and Brock, 1980). At the type 
locality of A^. kentensis, larvae of the sub- 
genus Notiphila were collected in associa- 
tion with the root systems of Nuphar (B. A. 
Foote, personal communication). Although 
specific identification is not possible at pres- 
ent, the association of the immature stages 
of Notiphila (Notiphila) with the yellow 
pond-lily deserves further study. 

In the description below, numerical char- 
acters follow Mathis (1979) and are based 
on male specimens. Unless otherwise des- 

ignated, other character states utilized are 
based upon examination of both male and 
female specimens. 

Notiphila (Notiphila) kentensis Huryn, 
New Species 

Description. — Shore flies of medium size 
[males 3.5-4,0 mm (n = 6); females 4.6-4.9 
mm (n = 3)]. Ground color blue-grey; ex- 
treme dorsolateral margins of mesonotum 
bordered by pair of distinct dark-brown 

//^«^.- Eye ratio 1:0.7 1-1 :0.78(n = 6); eye 
to cheek ratio 1 :0. 1 5-1:0.22; postfrons ratio 
1:1.25-1:1 .43; prefrons ratio 1 :0.62-l :0.75. 
Frons generally concolorous throughout, 
blue-grey. Paravertical bristles medium in 
size, noticeably more robust than postocel- 
lar setae. Single fine proclinate fronto orbital 
seta present. First and second antennal seg- 
ments brown, third segment variable, either: 
( 1 ) light brown proximally becoming darker 
distally, or (2) entirely dark brown; arista 
with 10-12 dorsal branches, usually 1 1 . Face 
microtomentose, variable: ( 1 ) generally yel- 
low near antennal bases becoming silver to- 
ward oral margin, or (2) yellow throughout. 
Facial setae fine; gena blue-grey; genal bris- 
tle similar in dimensions to paraverticle 
bristle; maxillary palps orange. 

Thorax: Mesonotum and pleural regions 
concolorous, blue-grey; lateral margins with 



Figs. 1-8. 1, 2, 4-8, Notiphila kentensis. 3, Notiphila mathisi. 1, Epandrium, ventral. 2, Epandrium, lateral. 
3, Basiphallus, dorsal. 4, Same. 5, Aedeagal apodeme, lateral. 6, Basiphallus, lateral. 7, Hypandrial process, 
lateral. 8, Hypandrial receptacle, lateral, a, Scale for Figs. 5-8. b. Scale for Figs. 1-4. 

distinctive brown stripes extending poster- 
iad from area anterior of presutural bristle, 
across the extreme dorsal region of noto- 
pleuron, terminating near base of supra-alar 

bristle. Anepisteraum with variable dark 
brown region consisting of either: ( 1 ) dark- 
ened region about prothoracic spiracle, (2) 
two darkened regions, one about spiracle 



and one located posterodorsally on pleurite, 
or (3) an elongate, rectangular darkened re- 
gion extending posteriorly from spiracle to 
posterior margin of pleurite. Lateral mar- 
gins of scutellum nearly black with pigmen- 
tation extending anteriad onto mesonotum 
to form short stripes terminating near bases 
of intra-alar bristles. Femora light-grey, yel- 
low apically; tibia and tarsi yellow; setal fas- 
cicle of hind basitarsus yellow. 

Abdomen: Abdominsd ratio 1:0.60-1:0.66 
(n = 5); tergum V/IV ratio 1:0.64-0.79; ter- 
gum V ratio 1 :0. 4 1-0.66. Ground color blue- 
grey with dark-brown geminate fascia on 
segments III-IV (e.g., fig. 2, Huryn, 1984). 
Male genitalia: epandrium generally rect- 
angular in shape (Fig. 1) with extreme an- 
terior tapered and bilobed, produced into 
anteriorly directed projection extending 
ventrad of epandrial processes (Fig. 2); 
epandrial processes narrow, parallel, form- 
ing lateral boundary of narrow emargina- 
tion (Fig. 1). Aedeagal apodeme as in Fig. 
3. Basiphallus (Fig. 6) strongly sclerotized 
with apical Vs strongly recurved; in dorsal 
view, parallel sided proximally with lateral 
margins converging distally (Fig. 8). Hy- 
pandrial process (Fig. 4) considerably longer 
than wide (width : length ratio ca. 1:7), par- 
allel with no indication of club on apical 
portion; apical third sparsely invested with 
fine spinules; hypandrial receptacle reduced 
to 2 elongate sclerotized strips (Fig. 5). 

Specimens examined.— Holotype 6. Ohio, 
Portage County, 1.3 km E of Kent State 
University. 13 September 1984. Julie L. 
Todd, deposited in National Museum of 
Natural History, Smithsonian Institution. 
Paratypes: 5 <5, 3 5, same data as holotype 
except 19 July 1984; 9 S, same data as ho- 
lotype except 31 August 1984; 1 6, same 
data as holotype. Deposited in USNM (5 6, 
1 9), KSU (5 6, 1 9), and University of Geor- 
gia (5 $, 1 9). 

Etymology. — Through the efforts of Ben- 
jamin A. Foote and his colleagues, the wet- 
land areas surrounding the Kent State Uni- 
versity campus have been the site of 
numerous studies of the Ephydridae. It is 
in recognition of these accomplishments and 
the type locality that I name the new species 
A^. kentensis. 


Notiphila kentensis is a member of the 
adjusta species group as defined by Mathis 
(1979) and is apparently most closely relat- 
ed to Notiphila mathisi. These species are 
readily distinguished by characters of the 
basiphallus. The lateral margins of the bas- 
iphallus of A^. kentensis converge distally to 
form an acutely angled structure (Fig. 4), 
whereas those of TV. mathisi diverge to form 
a spoon-shaped structure (Fig. 3). A^. math- 
isi is known only from the Okefenokee 
Swamp, Georgia (Huryn, 1984). 


All specimens utilized in this study were 
obtained through the efforts of J. L. Todd 
and B. A. Foote. W. N. Mathis examined 
and offered opinion on specimens of A^. ken- 

Literature Cited 

Huryn, A. D. 1984. New Notiphila (Diptera: Ephy- 
dridae) from the Okefenokee Swamp, Georgia. 
Proc. Entomol. Soc. Wash. 86: 942-945. 

Mathis, W. N. 1979. Studies of the Notophilinae 
(Diptera: Ephydridae), I: Revision of the Nearctic 
species oi Notiphila Fallen, excluding the caudata 
group. Smithson. Contrib. Zool. 287: l-1 1 1 + iv. 

Todd, J. L. 1985. The community organization of 
acalypterate Diptera in a freshwater marsh. M.S. 
thesis. Kent State University, Kent, Ohio. 

Van Der Velde, G. and Brock, Th. C. M. 1980. The 
life history and habits of Notiphila bnmnipes Ro- 
bineau-Desvoidy (Diptera: Ephydridae), an aut- 
ecological study on a fly associated with nym- 
phaeid vegetations. Tijdschr. Entomol. 123: 105- 


89(2), 1987, pp. 325-328 


W. R. M. Mason 

Biosystematics Research Centre, Agriculture Canada, Research Branch, Ottawa, Ontario 
KIA 0C6, Canada. 

Abstract. — Vadum volatum, a new genus and species of Diospilini (Braconidae), is de- 
scribed from southern Missouri and eastern Texas. It is distinguished from its nearest 
relatives, Diospilus and Taphaeus, by the long, narrow first metasomal tergum, the trun- 
cately toothed clypeus and the four-segmented labial palpus. 

Diospilini are a small but widely distrib- 
uted tribe of Braconidae placed in Helcon- 
inae and believed to be parasitic on Co- 
leoptera larvae with few or no exceptions. 
There are about 10 described genera, the 
majority of which contain only one or two 
tropical species of uncertain status. Generic 
names representing tropical groups that I 
believe should be placed in Diospilini are: 
Austrodolops Blanchard, Eudiospilus Sze- 
pligeti, Parabaeacis Granger, Repetiodio- 
spilus Shenefelt, and Westwoodiella Szepli- 
geti. There is an obvious need for 
considerably more taxonomic work, espe- 
cially on the rich tropical forest faunas, and 
one should not lightly add new genera. 
However, the Holarctic fauna is much bet- 
ter known, containing only a few genera: 
many species oi Diospilus Haliday, a few of 
the doubtfully different Taphaeus Wesmael, 
and one or two each placed in Baeacis 
Foerster and Aspigonus Wesmael. The new 
genus here described differs radically from 
all Diospilini known to me by its extremely 
long and untapered first metasomal tergite 
and by the median elevated bilobate pro- 
jection of the clypeus that appears like a 
truncate tooth (most Diospilini have the 
clypeal margin flat and evenly curved, 
sometimes, e.g. Aspigonus and Baeacis, 

bearing a small single median point). In ad- 
dition, the new genus has four-segmented 
labial palpi and the second flagellomere 
about '/3 longer than the first, features ap- 
parently unique within the tribe, where the 
labial palpi are usually three-segmented and 
the first flagellomere is normally the longest. 

Vadum Mason, New Genus 

Type species. — Vadum volatum Mason, 
new species. 

Head transverse and large (Fig. 1), about 
50% wider than mesoscutum between teg- 
ulae; clypeus (Fig. 3) rather flat and wide, 
about 3 X as wide as long, central quarter 
elevated and bilobately protruding, forming 
a truncate tooth-like elevation; labrum about 
half as wide as clypeus, its margin arcuate 
but flattened medially; anterior tentorial pits 
very large, with a deep groove between them; 
laterally clypeus separated from face by a 
distinct fine groove; mandibles not twisted, 
teeth subequal; maxillary palpi with 6 ar- 
ticles, length about equal to height of head; 
labial palpi short, with 4 articles; hyposto- 
mal and occipital carinae complete, meeting 
at a distance above mandibular opening 
about equal to basal width of mandible. 

Pronotum (Fig. 2) medially with a 
t/-shaped carina, which is concave ante- 



Figs. 1-4. Vadum volatum, n. sp. L Dorsal aspect of 2, wings drawn following conventions of Mason (1986). 
2, Pronotum, dorsal aspect. 3, Head. 4, Abdomen of 9. 

riorly and about % as wide as ocellar tri- 
angle, its arms enclosing a shallow, for- 
ward-facing cavity; propleuron with a ven- 
trolateral lobe overlapping pronotum. 
Mesothorax smooth, notauli strong, rugose; 
transscutal groove visible medially; axillae 
broad, roundedly sloping down behind, lat- 
eral margin defined by a strong carina; pre- 
scutellar scrobe very wide and deep, with a 
few transverse costae; post scutellar groove 

broad, transcostate; epicnemial carina 
strong, extending far dorsally; metapleuron 
and propodeum mostly rugose and bearing 
vague carinae. 

Wings (Fig. 1) with full venation; in fore- 
wing vein M reaching R proximad to origin 
of Rs, i.e. 1st discoidal cell sessile; A, with 
a strong dip at which a sclerotized, tubular 
stump of A2 occurs, vein a absent; veins 2RS 
and 2r-m weakly converging anteriorly. Vein 



A. of hind wing absent. Legs long and slen- 
der; mid and hind tibial spurs shorter than 
apical depth of tibia; tarsal claws simple. 

Abdomen (Figs. 1, 4) long and slender, 
about '/3 as wide as thorax and a little less 
than twice as long. Metasoma I in dorsal 
view about 4 times as long as its minimum 
(subbasal) width; base and apex of equal 
width, about 10% wider than minimum; 
spiracles prominent, subbasal; dorsal and 
dorsolateral carinae conspicuous basally; 
ventral edges of tergite widely separated; 
complete first sternite about half as long as 
first tergite; basal strongly sclerotized part 
of sternite I smooth, pointed posteriorly and 
about '/3 as long as tergite I; sternite 2 lying 
half beneath tergite I and half beneath ter- 
gite II; sculpture of tergite I completely ru- 
gose-reticulate. Remainder of metasoma 
smooth, apical terga extensively telescoped, 
metasomal terga 1-5 occupying basal 0.8- 
0.9 of length. Ovipositor (Fig. 4) very long 
and slender, about 1.6 times length of fore- 
wing or twice length of metasoma. Valvulae 
2 with a subapical dorsal notch. Cerci half- 
discoid, articulated. 

Life-history and immature stages un- 

The generic name is neuter, referring to 
shallow water or a shoal; the specific name 
means turning about or rolling. The epithet 
commemorates the type locality, the Becker 
Farm, known as "Rolling Shoals." 

Vadum volatum Mason, New Species 

Figs. 1-4 

Clypeus (Fig. 3) confused punctate ba- 
sally; median bilobate elevation shining, 
impunctate; apicolateral margins strongly 
depressed and narrowly decurved. Face and 
genae smooth, with sparse, fine punctures; 
vertex very sparsely punctate, frons im- 
punctate, shining. Antennae long and slen- 
der, about 25% longer than forewing; basal 
flagellomeres about 4 times longer than wide, 
apical ones about 1.5 times longer than wide, 
very little taper; flagellum broken at article 

29; first flagellomere about 75% as long as 

Sides of pronotum broadly smooth above 
and below with a broad central rugulose area. 
Scutum and scutellum shining, punctate; 
notauli broad, rugulose near pronotum, deep 
and transcostulate centrally, merging into a 
large rugose area behind; mesopleuron 
mostly smooth but bearing a broad, sinuate 
rugulose groove from upper epicnemium to 
middle coxa. 

Color.— Thorax and legs fulvous with pi- 
ceous to castaneous prothorax, propodeum, 
hind tibia and tarsus. Head and abdomen 
black; mouthparts fulvous. Wings hyaline 
with brown veins and stigma. 

Variation. — Very little in the few speci- 
mens available; flagellum of 28-32 articles; 
length of forewing 3.3-4.5 mm, of head and 
body 4-6 mm, of ovipositor sheath 5-7.5 
mm; metapleuron and notauli sometimes 
piceous. Both males have forewing about 4 
mm long and broken antennae, otherwise 
very similar to female in all somatic char- 

Specimens. — 2 3, 5 9, as follows: Holotype 
9, Missouri, Williamsville, J. T. Becker, 
Malaise Trap. June 1-16, 1969. CNC No. 
19357. Paratypes: 1 3, 2 9, same data but 1 
9 dated 16-26 June (CNC). Texas, Brazos 
Co., College Stn., R. Wharton, Malaise Trap. 
1 3, May 4, 1983; 2 9, April 11-15 and 12- 
25, 1981 (Tx. A.M.U.). 


Vadum resembles the cosmopolitan gen- 
era Diospilus and Taphaeus in most of its 
features and appears closely related. Vadum 
differs from the many species of Diospilus 
and Taphaeus by the following four fea- 
tures, each of which appears derived in Vad- 
um: (character state for Diospilus and Tap- 
haeus in parentheses) 1, apical margin of 
clypeus (Fig. 3) depressed on the lateral 0.45 
so that there appears to be an elevated me- 
dian bilobate tooth (margin of clypeus not 
depressed and bearing no such median 



tooth), 2, second flagellomere distinctly 
longer than the first (first longer than sec- 
ond), 3, metasomal tergite I unusually elon- 
gated, the base and apex equally wide, 
length/apical width = 3.0-4.0 (metasomal 
tergite I broadening apically, length/apical 
width = 1.0-1.5), 4, in forewing vein a ab- 
sent (vein a present). Vadum might be re- 
garded as no more than a derived species 
o{Diospilus were it not for the labial palpus 
which has four articles of subequal length, 
whereas all the species of Diospilus (as well 
as all other Diospilini known to me, in- 
cluding Aspigonus, Baeacis and Taphaeus) 

have only three articles in the labial palpus, 
clearly a derived condition. Thus I believe 
the evidence indicates Vadum to be a sister 
group to Diospilus and the other three gen- 
era named above. 

I thank my colleagues M. J. Sharkey and 
J. R. Barron for reviewing the manuscript 
and making useful suggestions. 

Literature Cited 

Mason, W. R. M. 1986. Standard drawing conven- 
tions and definitions for venational and other fea- 
tures of wings of Hymenoptera. Proc. Entomol. 
Soc. Wash. 88: 1-7. 

89(2), 1987, pp. 329-343 




D. Adamski and R. L. Brown 

Department of Entomology, Drawer EM, Mississippi State University, Mississippi State, 
Mississippi 39762. 

Abstract.— k new species of Blastobasidae, Glyphidocera juniperella Adamski, from 
southeastern United States is described. This species feeds on Juniperus and is a pest on 
ornamental varieties. Illustrations of the adult, male and female genitalia, and larval 
chaetotaxy are provided. Scanning electron micrographs of egg, larva, pupa, and abdom- 
inal sex scales of male adult are included. 

Glyphidocera and its type-species, G. au- 
dax, were described from Saint Vincent Is- 
land in the West Indies by Walsingham 
( 1 892). Thirty-nine species occur in the New 
World tropics and 9 species occur in Amer- 
ica north of Mexico. Host plants and biol- 
ogy of these species are unknown. 

Many Glyphidocera are uniformly brown 
and usually possess relatively broad and 
truncate forewings. The genitalia are char- 
acterized by the following: male with valvae 
narrowed basally, abruptly broadened dis- 
tally, apex protracted; costa with fingerlike 
projection at base; gnathos projecting from 
beneath tuba analis; aedeagus with medium 
to large comutus; female usually with duc- 
tus bursae sclerotized and apically expand- 
ed; ductus seminalis spiralled and originat- 
ing from posterior end of corpus bursae; 
corpus bursae with two patches of denticles 
at the anterior and posterior ends, and usu- 
ally with a sclerotized plate near constricted 
end of ductus bursae. 

This study resulted from the discovery of 
larvae of a new species of Glyphidocera 
feeding on nursery stock varieties o{ Junip- 
erus horizontalis Moench, in Mississippi and 
Florida. The biology of this species has been 

studied by Schiffhauer and Mizell (accepted 
for publication), and insecticides have been 
evaluated for its control by Mizell and 
Schiffhauer (accepted for publication). 

Materials and Methods 

The adult, egg, larva, and pupa were ex- 
amined with an incandescent light source 
(reflected light). The Methuen Handbook of 
Colour (Komerup and Wanscher, 1978) was 
used as a color standard for the description 
of the adult. Genitalia were dissected as de- 
scribed by Clarke (1941), except Mercuro- 
chrome and chlorazol black were used as 
stains. All preparations were examined with 
dissecting and compound microscopes. 

The ultrastructure of egg, larva, pupa, and 
abdominal sex scales of males was studied 
with a Hitachi HH-S-2R scanning electron 
microscope at an accelerating voltage of 20 
kV. For SEM examination, immature spec- 
imens were fixed in 3% glutaraldehyde in 
0. 1 M potassium phosphate buffer (pH 7.3), 
rinsed in phosphate (pH 7.3), and postfixed 
in 2% osmium tetroxide in 0. 1 M potassium 
phosphate (pH 7.3). After dehydration in 
ethyl alcohol, specimens were critical point 
dried, mounted on stubs with silver paint 



Fig. 1 . Holotype of Glyphidocera juniperella Adamski (male) (9.2 x ). 

and paste, and coated with gold-palladium 
in a Polaron E5100 sputter coater. 

Abbreviations used in this paper are: 
ANT, antenna; AX TB, axillary tubercle; C, 
cremaster; FR CL S, frontal clypeal suture; 
GL, galea; L3 (Fig. 28), metathoracic leg; 
LB, labium; LBP, labial palp; MD, man- 
dible; MX, maxilla; MXP, maxillary palp; 
?, pit (chordotonal organ, pore, other); PIV 
A, pivot area; PR SC, proleg scars; SP, spi- 
racle; SPIN, spinneret; TON FIB PL, tonofi- 
brillary platelets. 

Glyphidocera juniperella Adamski, 
New Species 

kduW. — Head: Scales light-brown or 
greyish-orange basally, greyish-orange api- 
cally; frons usually lighter than vertex; an- 
tenna dorsally with dark-brown and grey- 
ish-orange scales intermixed, ventrally with 
greyish-orange; labial palpus medially light 
greyish-orange, or light greyish-orange scales 
intermixed with dark-brown scales, later- 

ally dark-brown, apex of second segment 
usually greyish-orange. 

Thorax: Tegulae, mesonotum, and meta- 
scutum dark-brown or with dark-brown 
scales intermixed with greyish-orange scales; 
color of legs similar to labial palpi. 

Forewing {V\g. 1): 7.0-8.5 mm long; dor- 
sal surface with greyish-orange scales inter- 
mixed with dark-brown scales, appearing 
uniformly light-brown without microscope; 
one to two small dark-brown spots present 
in discal cell near distal end and near mid- 
dle; some specimens with dark-brown spots 
near base and on Cu near middle of wing; 
fringe scales brownish-grey. Ventral surface 

Hindwing: Dorsal surface light orange- 
grey at base, becoming greyish-orange api- 
cally; ventral surface light brownish-grey, 
intermixed with dark-brown on costa. 

Abdomen: Greyish-orange scales inter- 
mixed with dark-brown scales; male with 
irregular, transverse row of sex scales on 




Fig. 2. Dorsal view of abdominal terga 1-4 of male Glyphidocera juniperella. Sex scales are indicated by 
arrows pointed towards anterior margins of terga 3-4. Line scale = 1 mm. 

anterior margins of terga III and IV (Fig. 2); 
each scale with sub-parallel sides, obtuse 
apex, with base abruptly curved to short 
pedicel (Fig. 3); interridge grooves deep, fe- 
nestrae oval, irregularly spaced, each cir- 
cumscribed by a wide rim; some cross ribs 
medially fused and parallel with longitudi- 
nal ridges (Fig. 4). 

Male genitalia (Fig. 5): Tegumen with 
heavily sclerotized anterior margin, scaled 
laterally below gnathos; uncus without scales 
or setae, dorsally sclerotized and confluent 
with tegumen, ventrally sclerotized to near 
base; gnathos fused and projecting dorsally 
from below tuba analis, sparsely setose; ae- 
deagus long, comutus medium sized; juxta 
dorsally rolled around base of aedeagus, 
ventrally narrowed and elongated; valva 
heavily sclerotized except for ventral half of 
cucullus and medial groove from base to 
cucullus; costa sparsely setose, basally with 

fingerlike, apically setose projection; cucul- 
lus with ventral half densely setose, dorsal 
half relatively bare; sacculus with single row 
of 5-7 setae near middle and numerous se- 
tae on ventral margin. 

Female genitalia (Fig. 6): Sternum and 
tergum VII lightly sclerotized and evenly 
scaled; sternum and tergum VIII with nu- 
merous microtrichia that increase in density 
toward posterior margins, with several long, 
simple setae restricted to posterior half of 
sclerites; papillae anales with two types of 
setae: type I setae long, simple, directed pos- 
teriorly and posterolaterally; type II setae 
(visible at high magnification) minute, re- 
curved, and directed laterally; antrum with 
two internal, basal prongs, each directed 
posterolaterally; corpus bursae with smooth, 
sub-circular plate near antrum, anterior 
cluster of denticles near inception of ductus 
seminalis, and posterior cluster of denticles 



Figs. 3, 4. SEM of abdominal sex scales of male Glyphidocera juniperella. 3, 4000 x . 4, 20,000 ; 



Fig. 5. Genitalia of male Glyphidocera juniperella (aedeagus is detached). D. Adamski Gen. Slide no. 2854. 
Scale = 1 mm. 

at the posterior end; ductus seminalis spi- 
ralled, bearing cluster of denticles near base. 
Egg. — Eggs laid singly or in imbricate 
clusters on scales of branchlets of Juniperus 
spp. Egg (Figs. 7-8) width nearly two-thirds 
the length; broadly raised along the median 
longitudinal axis and flattened towards the 
outer margin. The surface is characterized 
by slightly raised convolutions. Aeropyles 

appear to be distributed nonrandomly over 
the dorsal surface. A distinct micropylar area 
has not been detected. Color changes of the 
embryo (ranging from yellow-orange to 
dark-brown) are readily observable through 
the translucent chorion. 

Larva. -Length 12.0-15.6 mm [lOn]. 
Body greyish-brown with a velvety textured 
appearance due to slightly raised stellate cu- 



Fig. 6. Genitalia of female Glyphidocera juniperella. D. Adamski Gen. Slide no. 2855. Line scale = 1 mm. 



Figs. 7, 8. SEM of egg o{ Glyphidocera juniperella. 7, 150x. 8, 750 x (arrows point to aeropyles). 



ticular projections on the surface; head cap- 
sule, prothoracic shield, anal plate, pina- 
cula, and legs dark brown. Head (Figs. 9- 
10): hypognathous; epicranium with ridges 
arranged polygonally, ridges on mouthparts 
indistinct; adfrontal sclerites broad (Fig. 1 7), 
delimiting frons dorsolaterally; frons closed; 
Fl, F2, and C3 sub-equal in length; Fl clos- 
er to C3 than F2, in straight line with C2; 
PI long, closer to P2 than A2; Al closer to 
A2 than to A3; CI arising from small 
depression on ventral lateral margin of clyp- 
eus, C2 closer to CI than C3; labrum with 
distal margin modified into two lobes pro- 
jecting medially, with 12 setae, two medial 
pairs, with outer pair at least four times 
length of inner pair, one pair on each lateral 
margin, and two pairs on distal margin; 
mandibles (Figs. 10, 15-16) slightly asym- 
metrical, with pair of setae on outer surface 
(basal seta longer than distal seta (Fig. 10)); 
labium strongly ridged distally, with micro- 
trichia proximally; submental pit absent; 
spinneret bulbous at base, narrowed distal- 
ly; labial palpus two segmented, each seg- 
ment with a dorsally directed apical seta; 
maxilla (Figs. 10-11, 13) prominent; sen- 
silla types and arrangement on medial lobe 
and apex of palpus (Figs. 11, 13), similar to 
those of Choristoneura fumiferana (Clem.) 
(Albert, 1980), Heliothis zea "(Boddie), (Ave, 
1981), and other ditrysian Lepidoptera 
(Grimes and Neunzig, 1986a, b). Stemmata 
III and IV approximate, associated setae as 
in Fig. 21. Sensilla types on antenna (Fig. 
14) similar to other Lepidoptera (Schoon- 
hoven and Dethier, 1966). Prothorax (Fig. 
18): prothoracic shield medially bissected 
by ecdysial line; Dl and D2 parallel or sub- 
parallel, Dl slightly more than half the length 
of D2; LI at least twice the length of L2, 
closer to L2 than L3, and slightly below line 
between L2 and L3, L2 and L3 sub-equal 
in length; SVl and SV2 widely divergent, 
SV2 slightly shorter than SVl and pointed 
posteriorly; distance between SVl and SV2 
less than distance between LI and L2. Me- 

sothorax (Fig. 18): Dl anterodorsal to D2, 
on same pinaculum; Dl slightly more than 
half the length of D2, SD2 slightly more 
than half the length of SDl; L2 and LI di- 
vergent, on same pinaculum; L2 more than 
twice length of LI; L3 posterodorsal to LI; 
MVl on small pinaculum, located within 
fold on antero ventral margin of segment be- 
tween L and SV groups. Metathorax as de- 
scribed for mesothorax. Abdomen (Figs. 1 8- 
20, 22-26): prolegs on A3-6 and AlO and 
of equal size; crochets uniserial and bior- 
dinal (Fig. 26); Dl and D2 parallel or sub- 
parallel on A 1-9; Dl anterior to and widely 
separate from D2 on A 1-8, and antero ven- 
tral and closer to D2 on A9; SDl and SD2 
dorsal to spiracle on A 1-8, SD2 minute, 
detectable only at high magnification (Figs. 
22-23); SDl and SD2 on A 1-8 approximate 
to an invagination on pinaculum; LI and 
L2 in nearly straight line with spiracle and 
SD 1 on A 1 , anterior to line between spiracle 
and SDl on A2-8, L3 pinaculum postero- 
ventral to LI and L2 pinaculum on A 1-8, 
ventral to LI and L2 pinaculum on A9, L3 
in straight line with L2 and SV3 on A9; SV 
bisetose on Al and A7, trisetose on A2-6, 
unisetose on A8-9; VI pinacula in or near 
straight line between each pair of SV's on 
A7-8, slightly anterior to line between SV's 
on A9, Vl's slightly closer on A9 than A7- 
8; AlO with Dl's athwart (Fig. 20); D2's 
stout, recumbent, and directed ventrally 
(Figs. 25, 27). Small, slightly pigmented, ir- 
regularly shaped and sized, tonofibrillary 
"platelets" are located in transverse folds of 
T2-3, on intersegmental membrane of T2- 
A9 on two irregular lines, one between D 
and SD setal groups, and one between spi- 
racles and L setal groups (Figs. 22, 24). 

Pupa. — Pupa (Figs. 28-34): scabrous, an- 
teriorly truncate, widened in thoracic re- 
gion, narrowed posteriorly to a bifurcate 
cremaster; ecdysial line extending from pos- 
terior margin of metathorax to anterior 
margin of vertex; axillary tubercles (cocoon 
cutter) located on each anterolateral margin 



Figs. 9-14. SEM of larva of Glyphidocera juniperella. 9, Frontal view of head capsule, 55 x. 10, Ventral 
view of head capsule, 150x. 1 1, Maxillae, 850 x. 12, Labium, lOOOx. 13, Sensilla on apex of maxillary palpus, 
5000 X. A2 = sensillum styloconicum; Al, A3, Ml, M2, LI, L2, and L3 = sensilla basiconica; SD = sensillum 
digitiform; 14, sensilla on apical portion of antenna, 2500 x, 1 = sensilla basiconica, 2 = sensilla chaetica, 3 = 
sensillum styloconicum, 4 = sensillum trichodeum. 





H2 L3 


















Figs. 15-2L Larva of Glyphidocera juniperella. 15, Right mandible. 16, Left mandible. 17, Frons and 
associated setae. 18, Setal map of thorax and Al-7. 19, Setal map of A8-10. 20, Setal map of AlO. 21, 
Arrangement of stemmata with associated setae. 

of mesothorax (Figs. 28-29, 31); epicranial 
suture V-shaped; fronto-clypeal suture pres- 
ent; antenna, maxilla, legs 1, 2, and apex of 
3 exposed ventrally, femur of leg 1 exposed 
as short, narrow sclerite, leg 3 extending be- 
yond margin of forewing; terminal six ab- 
dominal segments, pivoting as a unit from 
intersegmental area between fourth and fifth 

segments, pivotal area with narrow ridges 
separated by areas with irregular rows of 
small pits (Figs. 29-32); A5-6 with scars of 
abdominal prolegs present; pupal setae long 
and straight, setae on cremaster are apically 
recurved (Figs. 30, 33-34). 

Holotype.— (5, Miss.[issippi], Monroe Co., 
Hamilton, Amfac Nursery, 25 Aug. 1981, 



Figs. 22-27. SEM of larvae of Glyphidocera juniperella. 22, Portion of A6 showing positional relationship 
between spiracle and SD group and tonofibrillary platelets, 200 x . 23, SDl and SD2 on Al. Note invagination 
(labeled "?"), 100 x. 24, Tonofibrillary platelets and stellate integument (Al), 1000 x. 25, A8-10, 50x. 26, Left 
proleg on A5, 500 x . 27, Anal plate with large recumbent D2 setae and AlO prolegs, 100 x . 



Figs. 28-30. SEM of pupa of Glyphidocera juniperella. 28, Lateral view, 30; 
ventral view, 30 x . 

29, Dorsal view, 30 x . 30, 

David Tatum [Coll.]; iss. 1 4-IX-8 1 , ex Gold 
Coast Juniper. The holotype is deposited in 
the U.S. National Museum on indefinite loan 
from the Mississippi Entomological Mu- 
seum (MEM). Data are given as on labels 
except for bracketed information. 

Paratypes. — Mississippi: Monroe Co., 

Hamilton, 25 Aug. 1981, R. L. Brown 
[Coll.], ex Juniperus, 2 S\ David Tatum 
[Coll.], ex Gold Coast Juniper, em. 26-28 
Aug. 1981, 2 (5, D. Adamski genitalia slide 
no. 577; 5 9, D. Adamski gen. si. nos. 542, 
580, 582, and 2855; em. 29 Aug. 1981, 3 3, 
D. Adamski gen. si. nos. 579, 585, and 2888; 



Figs. 31-34. SEM of pupa of Glyphidocera juniperella. 31, Auxiliary tubercle (= cocoon cutter), 400 x. 32, 
Area demarcating pivotal area of pupa, 450 x . 33, Ventral view of cremaster, 1 50 x . 34, Dorsal view of cremaster, 

3 9, D. Adamski gen. si. no. 58 1 ; em. 3 Sept. 
1981, 6 6, D. Adamski gen. si. nos. 587, 
2853, 2854, 2887, USNM gen. si. nos. 
1 1426, 1 1427; 3 9, D. Adamski gen. si. no. 
2856, USNM gen. si. no. 1 1429; em. 5 Sept. 
1981, 4 5, D. Adamski gen. si. nos. 578, 
584, and 586; 1 9, USNM gen. si. no. 1 1428; 
iss. 12-IX-81, 4 (3, 2 9. Paratypes of male 
and female are deposited in the British Mu- 
seum (Natural History). Other paratypes are 
deposited in the Mississippi Entomological 
Museum (MEM). 

Other specimens examined. — F/orzWa.- 
Gadsden Co., Quincy, larva coll. 1 Apr. 
1981, Juniperus horizontalis ''wiltoni,'' R. 
F. Mizell 2 <5. Maryland: Prince George's 

Co., Adelphi, VIII-8-1970, R. W. Hodges 
2 9; USNM gen. si. no. 81436. Mississippi: 
Hinds Co., Clinton, 20 Jul. 1963, and 24 
Jul. 1963, Bryan Mather 2 9; Clay Co., West 
Point, Gold Coast Juniper, 5-23-80, J. D. 
Solomon; Hopkins no. SI 705, 1 9. Larvae— 
Rorida, Gadsden Co., Quincy, ex Juniperus 
horizontalis ""wiltoni,'' Imperial Nursery, 1 
Apr. 1984, R. F. Mizell, 10 fifth instars, 14 
early instars. 


Based upon the examination of type- 
specimens of all Glyphidocera species at 
the United States National Museum and 
British Museum, G . juniperella is most sim- 



ilar in wing coloration and pattern to G. 
barythyma Meyrick, described from For- 
estburg, Texas and G. rhypara Walsingham, 
described from Guerrero, Mexico. How- 
ever, specimens are paler in the latter two 
species. Males of G. barythyma can be dis- 
tinguished from those of G. juniperella by 
the presence of abdominal sex scales only 
on tergum 3. G. juniperella has sex scales 
on terga 3 and 4. The male genitalia of G. 
barythyma have a cucuUus that is broader 
at the apex, a shorter fingerlike projection 
at the base of the costa, and a larger and 
toothed comutus. Males of G. rhypara can 
be distinguished from G. juniperella by 
presence of a narrow valval with a rounded 
apex and a dense setal cluster on the inner 
surface of the cucullus. Female rhypara have 
a more narrow antrum and broader eighth 
sternum. In addition, the denticles within 
the corpus bursae in rhypara are stouter and 
are not basally attached to each other as in 
G. juniperella. 


Clarke (1969) was the first to illustrate 
abdominal sex scales and scale tufts on male 
Glyphidocera. G. juniperella males lack ab- 
dominal scale tufts; however, they do pos- 
sess tergal sex scales. These scales normally 
cannot be seen on pinned specimens be- 
cause they are covered by folded interseg- 
mental membrane, but they are easily de- 
tected on dissected specimens. The 
characteristic exterior rims of fenestrae in 
these sex scales also occur at various sex 
scales of Tortricidae (Brown and Miller, 
1983). The invagination or pit proximal to 
SDl and SD2 on the larval abdomen may 
demarcate muscle attachments of dorsal 
chordotonal organs (Kristensen, pers. 
comm.), a pore, or some other structure of 
unknown function. 


We thank David Tatum, Mississippi Di- 
vision of Plant Industry, for calling our at- 

tention to this species through his initial 
collection; Russell F. Mizell, of the Agri- 
cultural Research Center, University of 
Florida, for providing us with immatures as 
well as reared material; Ronald W. Hodges, 
Systematic Entomology Laboratory, Agri- 
cultural Research Service, at the U.S. Na- 
tional Museum, and Klaus Sattler of the 
British Museum (National History) for their 
cooperation provided during the examina- 
tion of type material; and Greta E. Tyson 
and Michael Sullivan, of the Electron Mi- 
croscopy Center, Mississippi State Univer- 
sity, for their help in the preparation of spec- 
imens and photographic plates. This 
research was supported in part by grants 
from NSF Grant BSR85-01212 and Sigma 

Literature Cited 

Albert, P. J. 1980. Morphology and innervation of 
mouthpart sensilla in larvae of the spruce bud- 
worm, Choristoneura fumiferana (Clem.) (Lepi- 
doptera: Tortricidae). Can. J. Zool. 58: 842-851. 

Ave, D. A. 1981. Induction of changes in the gus- 
tatory response by individual secondary plant 
compounds in larvae of Heliothis zea (Boddie) 
(Lepidoptera, Noctuidae). Ph.D. dissertation. De- 
partment of Entomology, Mississippi State Uni- 
versity. 89 pp., 4 tables, 37 figs. 

Brown, R. L. and P. R. Miller. 1983. Studies of Lep- 
idoptera hindwings with emphasis on ultrastruc- 
ture of scales in Cydia caryana (Fitch) (Tortrici- 
dae). Entomography 2: 261-295. 

Clarke, J. F. G. 1941. The preparation of slides of 
the genitalia of Lepidoptera. Bull. Brooklyn Ento- 
mol. Soc. 36: 149-161. 

. 1969. Catalogue of the Type Specimens of 

Microlepidoptera in the British Museum (Natural 
History) described by Edward Meyrick. Vol. VII. 
British Museum (Natural History), London. 531 

Grimes, L. R. and H. H. Neunzig. 1986a. Morpho- 
logical survey of the maxillae in last stage larvae 
of the suborder Ditrysia (Lepidoptera): Palpi. Ann. 
Entomol. Soc. Am. 79: 491-509. 

. 1986b. Morphological survey of the maxillae 

in last-stage larvae of the suborder Ditrysia (Lep- 
idoptera): Mesal lobes (laciniogaleae). Ann. Ento- 
mol. Soc. Am. 79: 510-526. 

Komerup, A. and J. H. Wanscher. 1978. Methuen 



Handbook of Colour. 2nd ed. Methuen and Co., 
Ltd.. London. 243 pp. 

Mizell, R. F. Ill and D. E. Schiffhauer. Evaluation of 
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cinae). Fl. Entomol. (accepted for publication). 

Schiffhauer, D. E. and R. F. Mizell IIL Bionomics of 
Glyphidocera juniperella Adamski, (Lepidoptera: 
Blastobasidae: Symmocinae) a newly discovered 

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cepted for publication). 

Schoonhoven, L. M. and Dethier, V. G. 1966. Sen- 
sory aspects of host-plant discrimination of lepi- 
dopterous larvae. Arch. Neerlandaises Zool. 1 6(4): 

Walsingham, Lord (Thomas de Grey). 1892. On the 
Micro-lepidoptera of the West Indies. Proc. Zool. 
Soc. Lond. 1892: 492-549. 


89(2), 1987, pp. 344-350 




R. E. Orth 

Department of Entomology, Division of Biological Control, University of California, 
Riverside, California 92521. 

Abstract— Pherbellia fisheri is described from Northwest Territories, Canada. New 
distributional information is recorded for P. griseicollis (Becker) and P. hackmani Roz- 
kosny is reported for the first time from North America. The male genitalia of all three 
species are illustrated and maps of the North American distribution records are given. 

In America north of Mexico the genus 
Pherbellia consists of 4 1 species including 
the new species described here, and P. hack- 
mani Rozkosny, recorded here from North 
America for the first time. Not among the 
above 41 species is P.fusca (Cresson), which 
recently has been considered a nomen du- 
bium (Knutson et al., 1986). 

This study (1) recognizes Pherbellia fish- 
eri, new species, (2) extends the known dis- 
tribution of P. hackmani, described from 
Europe, to North America, (3) expands the 
known distribution of P. griseicollis (Beck- 
er) in North America, (4) provides illustra- 
tions of the terminalia of the above three 
species, and (5) maps the localities in North 
America where the species have been re- 

Seven North American species including 
one subspecies of Pherbellia are now known 
to be Holarctic: P. albocostata (Fallen), P. 
argyra Verbeke, P. griseicollis, P. griseola 
(Fallen), P. hackmani, P. nana nana (Fal- 
len), and P. obscura Ringdahl. In the Palae- 
arctic, these seven species have essentially 
a Fennoscandian distribution; some of them 
extend into other parts of Europe, and to 
North Africa, Soviet Central Asia, Siberia 
and China. 

Several attempts have been made to sep- 

arate the genus Pherbellia into subgenera. 
The most recent division was by Rozkosny 
(1964). The species discussed herein are 
in the subgenus Chetocera Robineau-Des- 
voidy, as defined by Rozkosny. Rozkosny 
(1984) further alludes to a P. griseicollis 
group in which he listed P. griseicollis and 
P. sordida (Hendel). Pherbellia fisheri is a 
member of that group and perhaps P. hack- 
mani may also belong there. Furthermore, 
the Pherbellia fuscipes group (species with 
a row of hairs on the posterior margin of 
the anepistemum, as designated by Steyskal 
(1961), belongs to Chetocera, as defined by 
Rozkosny (1964). 

Pherbellia fisheri, P. griseicollis, P. hack- 
mani and P. sordida share the following 
combination of characters: 

1. Midfrontal stripe less than Vi distance 
from anterior ocellus to frontal margin. 

2. Two fronto-orbital bristles. 

3. Wing not patterned. 

4. First vein surpassing level of anterior 

5. Anepistemum bare. 

6. Anepimeron with 2 (rarely 3) large bris- 
tles and several fine setae. 

7. Katepistemum with only fine setae dor- 



Pherbellia fisheri Orth, New Species 

Figs. 1-3, 8 

Holotype male. — Height of head -h width. 
Medifacies yellowish, pruinose; facial 
grooves subshiny; parafacies and cheeks 
pruinose, pale yellow. Frons testaceous, yel- 
lowish anteriorly. Midfrontal stripe extend- 
ing less than Vi distance from anterior ocel- 
lus to frontal margin. Ocellar triangle and 
orbital plates tannish grey, tomentose. Or- 
bital plates tapered anteriorly, extending be- 
yond midfrontal stripe. Orbito-antennal spot 
lacking; narrow white tomentose stripe along 
upper orbital margin. Two pairs fronto-or- 
bital bristles; anterior bristle approximately 
% as long as posterior bristle; ocellars, post- 
ocellars, and inner and outer verticals well 
developed. Occiput tannish grey, tomen- 
tose. Short black setae on lower % of cheeks, 
on anterior '/2 of frons, between ocellar and 
postocellar bristles, on orbital plates, and in 
mid-cervical patch. Lateral occipital mar- 
gins with somewhat stronger setae and bris- 
tles. Antenna testaceous; aristal hairs ap- 
proximately as long as width of first segment 
of arista. 

Thorax greyish brown, tomentose, slight- 
ly mottled, with 4 indistinct brown longi- 
tudinal stripes. 

Pleura tannish grey, tomentose. Anepi- 
stemum bare, with upper posterior surface 
more brownish. Anepimeron with cluster of 
small bristles and 2 much longer, stronger 
bristles situated mid-anteriorly. Katepister- 
num with fine setae over much of surface 
and well-developed setae ventrally. Pro- 
sternum bare. 

Coxae greyish white, tomentose. Forefe- 
mur brownish, infumated. Mid- and hind- 
legs tawny, slightly infumated; hindfemur 
slightly darkened distally. Tarsal segments 
tawny except brown 4th and 5th segments 
on fore- and midlegs. 

Wing length 4.2 mm. Membrane greyish 
yellow, hyaline, costal margin and veins 
brownish yellow, area around crossveins 
clouded. No stump veins; first vein sur- 
passing level of anterior crossvein; anal vein 

reaching wing margin. Halter, squama, and 
squamal cilia pale yellow. 

Abdominal segments greyish brown, 
slightly infumated dorsally; terminalia as in 
Figs. 2 and 3. 

Allotype female. — Similar to holotype 
except for reproductive structures. Wing 
length 5.0 mm. 

Diagnosis.— Among Pherbellia species, P. 
fisheri appears to be most closely related to 
P. griseicollis and P. sordida. Identification 
should be made by examining male termi- 
nalia and using the locality maps. Pherbellia 
fisheri is set apart from both species by the 
smaller, less deeply emarginated ventral 
posterior lobe on the anterior surstylus (for 
illustrations of P. sordida see Rozkosny 

Holotype. — 3, CANADA, Northwest 
Territories, Aklavik, July 27, 1931, O. 
Bryant, Lot 234. Deposited in United States 
National Museum of Natural History. 

Allotype. — 2, same data as holotype ex- 
cept, July 27, 1 932, Lot 306. Deposited with 

Paratypes. -CANADA. Alberta: Banff, 
July 19, 1915 (1 9), N. B. Sansome, May 
25, 1922 (1 2, 3 <5), May 26, 1922 (2 9), June 
1, 1922 (1 3), June 21, 1922 (1 9), C. B. D. 
Garrett; Lancaster Park, July 28, 1963 (1 
$), J. R. Vockeroth; Waterton Lakes Na- 
tional Park, July 7-12, 1980 (1 <5), H. J. 
Teskey. Manitoba: Churchill, June 25, 1930 
(1 (5), collector ? Northwest Territories: Ak- 
lavik, Sept. 12, 1930 (1(5) Lot 142, Sept. 18, 
1930 (1 9) Lot 1 53, 0. Bryant; VSX. Alaska: 
Umiat, July 3, 1959 (1 9, 1 <5), July 7, 1959 
(3 9, 3 3), J. E. H. Martin. 

Etymology. — I take pleasure in naming 
this species in honor of T. W. Fisher, whom 
I have enjoyed working with for many years. 

Distribution. — This boreal species is 
known from widely separated areas in Alas- 
ka and Canada. The northernmost collect- 
ing site is Umiat, Alaska, 69°25', and the 
southernmost is Waterton Lakes National 
Park, Alberta, 49°06'. 

Discussion.— The illustrations of the ter- 



0.^ m 

Figs. 1-3. Pherbelliafisheri, new species. 1, Head, holotype male. 2, Terminalia, ventral view, paratype male. 
3, Terminalia, sinistral view, inverted, paratype male. 

minalia of Palaearctic Pherbellia sordida by 
Rozkosny (1984) resemble P. fisheri. A 
specimen of P. sordida collected in Wus- 
tung, Poland, by O. Duda, May 24, 1921, 
was borrowed from R. Rozkosny, J. E. Pur- 
kyne University, Brno, Czechoslovakia, for 
examination. Comparison of the terminalia 
of P. sordida with P. fisheri revealed them 
to be distinct. 

Pherbellia fisheri was independently rec- 
ognized as undescribed by L. Knutson sev- 
eral years ago. Upon receipt of my manu- 
script for review he informed me of his 
findings and suggested that I proceed with 
its description. He also provided most of 
the specimens used in this paper. I gratefully 
acknowledge his contribution. 

The northernmost collection sites of P. 



fisheri (Umiat, Alaska, and Aklavik, North- 
west Territories) lead one to speculate that, 
as with other similarly distributed species, 
P. fisheri may also occur in Fennoscandia. 
Closer study of material labeled P. grisei- 
collis from northern Europe may reveal the 
presence of the new species there. 

Pherbellia griseicollis (Becker) 
Figs. 4, 6, 8 

Externally, Pherbellia griseicollis and P. 
fisheri are so similar that I cannot separate 
them with a great degree of confidence. In 
general, the dorsum is greyer and the legs 
are slightly darker than P. fisheri. 

Distribution. — In the Palaearctic P. gris- 
eicollis is known to occur in Fennoscandia, 
the Murmansk region, the Karelian ASSR 
of the USSR, and in Western Siberia (Roz- 
kosny, 1984). Prior to this study, P. gris- 
eicollis was known only from Prudhoe Bay, 
Alaska, in North America (Knutson et al., 
1986). It is now known to be rather wide- 
spread with the southern limit of its distri- 
bution near Lima, Montana, approximately 
45°N latitude. 

Collection records.— CANADA. Alberta: 
Banff, July 25, 1924 (1 <3), E. Hearle; Banff 
National Park, June 4, 1955 (1 9), June 9, 
1955 (1 9), G. E. Shewell, July 9, 1955 (2 
9), J. R. McGillis. British Columbia: Buck- 
inghorse Provincial Campground, Alaska 
Hwy. DC-175 (281.6 km), June 27, 1978 (1 
(5), P. H. Amaud, Jr.; Lac le Jeune, June 25, 
1973 (2 9), June 27, 1973 (1 9, 1 <5), H. J. 
Teskey. Manitoba: Warkworth Creek, nr. 
Churchill, June 10, 1952 (1 5), G. E. Shew- 
ell. Northwest Territories: Aklavik, June 7, 
1931 (1 5), July 25, 1931 (1 3), O. Bryant; 
Yellow Knife, Kam Lake, June 20, 1966 (1 
(3), G. E. Shewell; 21 mi E of Tuktoyaktuk, 
June 17, 1971 (19), June 21-25, 1971(15), 
8-12 July, 1971 (1 9), 17-21 July, 1971 (2 
9), 20-25 July, 1971 (1 9, 1 3), D. M. Wood. 
Yukon: Herschel Island, 24-28 July, 1971 
(1 9, 1 3), D. M. Wood. VSX. Alaska: Circle, 
July 26, 1971 (1 3), B. A. Foote; N. Coast, 
Prudhoe Bay, June 8, 1971 (1 9), June 16, 

1971 (13), June 20, 1971 (13), July 4, 1971 
(1 9, 1 3), M. Deyrup; Umiat, Aug. 7, 1959 
(13), R. Madge. Idaho: Idaho Co., Lolo Pass, 
June 18, 1965 (1 3), R. L. Westcott. Mon- 
tana: Beaverhead Co., 9 mi S of Lima, June 
26, 1966 (1 3), B. A. Foote. 

Pherbellia hackmani Rozkosny 
Figs. 5, 7, 8 

Externally, P. hackmani is set apart from 
P. fisheri, P. griseicollis and P. sordida by a 
more intense clouding of the membrane 
bordering the anterior and posterior cross- 
veins and a yellowish brown gross aspect. 
The male terminalia are as in Figs. 5 and 7. 
The anterior surstyli are similar to P. argyra 
Verbeke (see Fisher and Orth, 1983). 

Distribution. — In the Palaearctic this 
species is known only from the type series 
(Rozkosny, 1984). The holotype male was 
collected at Mukkavuoma, Tome Lappmark, 
Sweden. There are ten additional paratypes 
from Sweden, Finland, and western Siberia. 

This study presents the first records of 
Pherbelli hackmani in North America. It is 
a boreal species, and has been recorded south 
of 60°N latitude only at King Salmon, Alas- 
ka (58°40'N, 156°40'W). 

Collection records.— CANADA. North- 
west Territories: Aklavik, June 15, 1930 (1 
9, 1 3), O. Bryant; 21 mi E of Tuctoyaktuk, 
July 17-21, 1971 (1 3), D. M. Wood; Tun- 
unuk, Aug. 15, 1930 (1 9, 1 3), O. Bryant. 
Yukon: Dickson Lake, Mt. Mye, 133°08'- 
62°21', 5000 ft., June 14, 1960 (1 9), J. E. 
H. Martin; Firth River, Aug. 3, 1956 (2 9), 
R. E. Leech; Herschel Is., July 22, 1953 (1 
3), J. S. Waterhouse, (1 9), C. D. Bird; North 
Fork Pass, Ogilvie Mts., 4100 ft., June 11, 
1962 (1 3), June 12, 1962 (1 9), June 20, 
1962 (1 3), R. E. Leech, June 18, 1962 (1 
9), July 1, 1962 (1 9), P. J. Skitsko. USA. 
Alaska: Cape Thompson, June 10, 1960 (sex 
?), W. C. Hanson; Cape Thompson, Crow- 
bill Mountain, June 6, 1960 (1 3), W. C. 
Hanson; Gulkana, Paxon Lodge, Aug. 4, 
1951 (2 9, 1 3), W. R. M. Mason; Isabella 
Pass, Mi. 206 Richardson Highway, 2900 


Figs. 4-7. 4, 6, Pherbellia griseicollis. male, 21 mi E of Tuktoyaktuk, Northwest Territories. 4, Terminalia, 
ventral view. 6, Terminalia, sinistral view, inverted. 5, 7, Pherbellia hackmani, male. Summit Lake, Isabella 
Pass, Alaska. 5, Terminalia, ventral view. 7, Terminalia, sinistral view, inverted. 





Fig. 8. Collection sites for Pherbellia griseicollis, P. hackmani, and P. fisheri. 



ft., July 17,1962(1 2), P. J. Skitsko; Isabella 
Pass, Summit Lake, July 9, 1951 (2 9), P. J. 
Skitsko, Aug. 3, 1951 (1 9, 1 <5), Mason- 
McGillis; King Salmon, Naknek River, July 
9, 1952 (1 3), J. B. Hartley, July 13, 1952 
(1 6), Aug. 4, 1952 (1 5), W. R. Mason; 
Kotzebue, June 24, 1951 (1 9), R. I. Sailer; 
Nome, July 9, 1951 (1 <5), D. P. Williams; 
Umiat, June 10, 1947 (1 9), C. Smith, July 
3, 1959 (1 9), July 10, 1959 (1 <3), J. E. H. 
Martin, July 10, 1959 (1 S), July 23, 1959 
(1 9), Aug. 3, 1959 (2 9, 2 <3), Aug. 7, 1959 
(1 9), Aug. 12, 1959 (1 9), R. Madge; Una- 
lakleet, June 11, 1961 (2 9), June 18, 1961 
(2 5), R. Madge. 


I thank the following individuals and in- 
stitutions for their assistance or loan of ma- 
terial: H. J. Teskey, Biosystematics Re- 
search Centre, Agriculture Canada, Ottawa, 
Ontario; L. Knutson, Biosystematics and 

Beneficial Insects Institute, USDA, Belts- 
ville, Maryland; A. D. Bratt, Calvin College, 
Grand Rapids, Michigan. A special thanks 
to T. W. Fisher, University of California, 
Riverside, and L. Knutson for reviewing the 

Literature Cited 

Fisher, T. W. and R. E. Orth. 1983. The Marsh Flies 
of California (Diptera: Sciomyzidae). Bull. Calif 
Insect Surv. 24, Univ. Calif Press, vii + 117 pp. 

Knutson, L., R. E. Orth, T. W. Fisher, and W. L. Mur- 
phy. 1986. Catalog of Sciomyzidae (Diptera) of 
America north of Mexico. Entomography 4: 1-53. 

Rozkosny, R. 1964. Zur Taxonomie der Gattung 
Pherbellia Robineau-Desvoidy (Diptera, Scio- 
myzidae). Acta Soc. Entomol. Cechoslov. 61(4): 

Rozkosny, R. 1984. The Sciomyzidae (Diptera) of 
Fennoscandia and Denmark. Fauna Entomol. 
Scand. 14: 1-224. 

Steyskal, G. C. 1961. The North American Scio- 
myzidae related to Pherbellia fuscipes (Macquart) 
(Diptera: Acalyptratae). Pap. Mich. Acad. Sci. Arts 
Lett. 46: 405-415. 


89(2), 1987, pp. 351-355 





Jay a. Rosenheim and J. Kenneth Grace 

Department of Entomological Sciences, 20 1 Wellman Hall, University of California, 
Berkeley, California 94720; JKG Current address: Faculty of Forestry, University of 
Toronto, Toronto, Ontario M5S lAl, Canada. 

Abstract.— An aggregation of Mimumesa mixta nests was located near the end of a 
decayed Douglas-fir timber in a second-story porch deck in Berkeley (Alameda Co.), 
California. Cells were excavated in the rotting wood in approximately linear series parallel 
to the grain and separated from one another by plugs of macerated wood fragments. Nests 
were provisioned with several adult and nymphal cicadellid and delphacid homopterans. 
Both M. mixta and its chrysidid parasite Elampus viridicyaneus Norton were reared from 
the nests. The sex ratio of A/, mixta did not differ significantly from 1:1, while all emerging 
E. viridicyaneus were female. The cocoons of both wasps are described. 

The genus Mimumesa (Hymenoptera: 
Sphecidae) is distributed throughout the 
Nearctic, Neotropical, Palearctic, and Ori- 
ental Regions (Bohart and Menke, 1976). 
These wasps excavate multicellular nests in 
level or sloping ground, decaying wood, and 
plant stems and provision them with del- 
phacid or cicadellid prey (Bohart and Menke, 
1976; Spooner, 1948; Tsuneki, 1959; Petit, 
1979). Gumey (1951) provided the only de- 
scription of the nesting biology of a North 
American species of Mimumesa, that of Mi- 
mumesa nigra (Packard). The discovery of 
a large aggregation of Mimumesa mixta (W. 
Fox) nests provided an opportunity to study 
the biology of this species as well as that of 
its parasite, Elampus viridicyaneus Norton 
(Hymenoptera: Chrysididae). 


The timber containing the nesting aggre- 
gation was collected in Berkeley, Alameda 
County, California on 15 July, 1985. To 

simulate natural conditions the timber was 
maintained in an exterior screened cage in 
a shaded exposure at the Oxford Agricul- 
tural Tract, University of California, Berke- 
ley until the adult emergence occurred dur- 
ing May, 1986. Cell dimensions were 
measured to the nearest millimeter, and host 
and parasite cocoons were measured with a 
calipers accurate to 0.05 mm. Voucher spec- 
imens of both wasp species have been de- 
posited in the R. M. Bohart Museum of 
Entomology, Department of Entomology, 
University of California, Davis. 


Nest location and architecture.— Nests 
were located near one end (approximately 
0.6 m) of a large timber made by glue-lam- 
inating seven Douglas-fir, Pseudotsuga 
menziessi (Mirb.) Franco, 2x4 (inch) 
boards together. The timber was found in a 
stack of lumber that had been removed dur- 
ing the previous week from a second story 





Figs. 1-4. 1, Mimumesa mixta nesting aggregation in decayed end of a 9 x 30 cm timber. 2, Linear series 
of M. mixta cells. 3, Cocoons of M. mixta (left) and Elarnpus viridicyaneus (right). Cocoons pictured in upper 
row are intact, while those below have been sectioned lengthwise to reveal internal structure. Small divisions 
of scale are in mm. 4, Cocoons of E. viridicyaneus (above) and M. mixta (below), showing emergence holes. 
Same scale as previous figure. 

porch deck. No active wasps were observed 
at the time of collection, the entire aggre- 
gation being composed of overwintering in- 
dividuals. The timber was located between 
2.5 and 4 m above ground. At the time of 
collection the end of the timber containing 
the wasp nests was dry but already decayed 
by brown-rot fungi (Basidiomycetes). M. 
mixta was apparently able to excavate the 
decay-softened wood. 

The nest aggregation consisted of over 400 
cells which extended throughout the de- 
cayed portion of the timber but not into the 
sound wood (Fig. 1). Cells were arranged 
end to end in approximately linear series 
parallel to the grain (Fig. 2), each being sep- 
arated from the next by a plug of macerated 
wood fragments. Nests appeared to be in- 
terconnected, and we were unable to deter- 

mine where an individual nest began or end- 
ed. Measurements of 30 randomly selected 
cells gave a mean cell length of 9.3 ± 1.8 
mm and mean width of 4.0 ± 0.6 mm (range 
5-12 mm by 3-5 mm). No cell contained 
more than a single cocoon. 

Host and parasite populations.— After 
adult emergence the wood was dissected to 
provide an estimate of the sizes of the host 
and parasite populations (Table 1). A total 
of 44 M. mixta emerged successfully. An 
additional 1 1 9 M. mixta larvae, pupae, and 
adults were found inside cocoons; some of 
these were alive, and it is unclear to what 
extent additional emergence might have oc- 
curred. Two host cocoons, one containing 
a dismembered adult wasp and the other a 
mass of insect frass, had apparently been 
attacked by fly larvae of the family Sciari- 



Table 1 . No. of specimens ofMimumesa mixta and 
Elampus vihdicyaneus reared or dissected from a nest- 
ing aggregation. 

Species and Stage No. 

Table 2. Species of Homoptera recovered from cells 
of Mimumesa mixta. 

Taxon No. Adults/Nymphs 

Mimumesa mixta 
Emerged adults 9 
Pupae/adults within cocoons 9 

Pupae within cocoons, unsexable' 
Larvae within cocoons 
Emerged cocoons 
Cocoons with fly feeding holes only 

Elampus viridicyaneus 
Emerged adults 9 
Pupae/adults within cocoons 2 
Pupae within cocoons, unsexable' 
Larvae within cocoons 
Emerged cocoons 
Cocoons with fly feeding holes only 

' Pupae that died before sclerotization. 







dae, which left a single, small (ca. 0.7 mm 
diameter), irregular hole in each cocoon. 
These fly larvae were common throughout 
the nest aggregation and appeared primarily 
to be fungivores, only incidentally attacking 
cocoons. The sexes of 63 of the sphecid pu- 
pae could be determined, which when com- 
bined with the data from the adults yielded 
a sex-ratio of 52 females: 56 males or 1: 
1.08, which was not significantly different 
from 1:1 (z = -0.39, P > 0.5). The 131 
host cocoons with emergence holes exceed- 
ed the number of emerged adults by 87, 
indicating that some of the cocoons may 
have remained from one or more previous 

Twenty-two adult E. viridicyaneus 
emerged successfully. Twenty-five larvae, 
pupae, and adults were found inside co- 
coons; some of these appeared to be alive 
and might have emerged later. Three empty 
parasite cocoons had apparently been at- 
tacked by fly larvae. The 22 adults and 12 
pupae whose sex could be determined were 


Delphacodes sp. 


Sorhoanus helvinus 
(Van Duzee) 

59 adults 

Tiaja californica (Ball) 

8 adults 

Euscelidius variegatus 


8 adults 

Amblysellus grex (Oman) 

5 adults 

Endria lassus (Ball) 

2 adults 

Calanana rubralineata 


1 adult 

Reticopsis nubila (Van Duzee) 

1 adult 

Deltocephalinae, undetermined 

10 adults 

Deltocephalinae, undetermined 

5 nymphs 

all females. Barring strongly sex-specific 
mortality in the immature stages, E. viri- 
dicyaneus showed a female-biased sex ratio 
(z = 5.8, P < 0.001). The 192 E. viridicy- 
aneus cocoons with normal emergence holes 
exceeded the number of observed emerged 
adults, suggesting the existence of one or 
more earlier generations of parasites in the 
same nesting aggregation. 

Nest provisions.— M. mixta provisioned 
cells with adult and nymphal Homoptera 
(Table 2). Nine species of cicadellids and 
one delphacid were recovered; 59 of the 100 
prey individuals recovered represented a 
single species of cicadellid, Sorhoanus hel- 
vinus (Van Duzee). Intact prey were found 
both in cells in which the wasp larva had 
died and in cells containing cocoons. Two 
apparently complete cells contained 1 8 and 
1 9 prey items, respectively. 

Mimumesa mixta cocoons.— Cocoons of 
M. mixta (Figs. 3, 4) were oblong, spongy 
in texture, and generally covered with wood 
and fragments of devoured provisions. Co- 
coons averaged 8.5 ± 0.7 mm long by 3.3 ± 
0.3 mm wide (range 6.7-9.7 mm by 2.8- 
3.8 mm, n = 60). The inner surface of the 
cocoon was light brown and very smooth; 



the end opposite that with the generally off- 
center exist hole was coated thinly with the 
dark brown meconium. 

Elampus viridicyaneus cocoons. — Co- 
coons of E. viridicyaneus (Figs. 3, 4) were 
superficially similar to those of M. mixta, 
being oblong and covered with wood frag- 
ments and prey debris. However, they were 
shorter, and the cocoon walls were crisp, 
paper-like, and had a varnished appearance. 
The inner cocoon surface was darker than 
that of M mixta and was roughly textured. 
The exit hole was centered and represented 
the removal of the end of the cocoon. Mean 
cocoon dimensions were 6.7 ± 0.8 mm long 
by 3.2 ± 0.2 mm wide (range 4.9-9.0 mm 
by 2.6-3.88 mm, n = 42). 


Mimumesa m/.\'/<2.— Several Mimumesa 
species are known to nest in the soil, and 
Mimumesa dahlbomi (Wesmael), Mimu- 
mesa dahlbomi pacifica (Tsuneki), and Mi- 
mumesa nigra (Packard) are known to nest 
in decayed wood (Bohart and Menke, 1 976; 
Spooner, 1948; Tsuneki, 1959; Petit, 1979). 
Spooner (1948) suggested that Mimumesa 
unicolor (Vander Linden) and Mimumesa 
lit t oralis (Bondroit) (as Mi mesa celtica 
Spooner) may nest in plant stems, but M. 
unicolor has subsequently been found nest- 
ing in the ground (Petit, 1979) and the bi- 
ology of M. littoralis has not been described. 
The short description by Gumey (1951) of 
a M. nigra nest is the only description of 
the biology of a North American species. 

Gumey (1951) reared a single M. nigra 
male from a cocoon taken from a nest con- 
taining "a dozen or more cells" in the de- 
cayed end of a 2 x 4 (inch) fence board ca. 
30 cm above ground. Although abandoned 
beetle borings are used by M. dahlbomi and 
M. dahlbomi pacifica to gain entry into the 
wood (Bohart and Menke, 1976; Spooner, 
1948; Tsuneki, 1959), Gumey (1951) did 
not mention any evidence of beetle infes- 
tation in the board containing the M. nigra 
nest. Likewise, we found no evidence of 

wood-boring beetle activity in the timber 
containing the M. mixta nests. M. mixta 
may either have excavated directly into the 
soft, decayed end-grain of the board or en- 
tered through cracks in the decayed wood. 
Species of the genus Psen, the only other 
genus in the subtribe Psenina known to nest 
in decayed wood (Bohart and Menke, 1 976), 
apparently excavate directly into stumps 
(Iwata, 1938; Tsuneki, 1959). 

The number of empty host and parasite 
cocoons exceeded the number of newly 
emerged adults, suggesting that both M. 
mixta and its parasite were active at the 
nesting site for one or more previous gen- 
erations. The complete history of the nest- 
ing aggregation cannot, however, be in- 
ferred from our single collection due to the 
possible re-use of old cells by M. mixta. 

Mimumesa unicolor, the only Mimumesa 
species known to be attacked by chrysidid 
parasites, has been reported as a host of 
Chrysis succincta L., Hedychridium ardens 
Cocquebert, and Omalus auratus (L.) (Bo- 
hart and Menke, 1976; Moczar, 1967; Spoon- 
er, 1948). 

Elampus viridicyaneus. —Tht genus 
Elampus contains six species in North 
America (Bohart and Kimsey, 1982) for 
which the biology is almost completely un- 
known (Huber and Pengelly, 1977). The only 
published host record for a North American 
species is for E. viridicyaneus parasitizing 
Hoplisoides costalis (Cresson) (as Psam- 
maecius costalis (Cr.)) (Krombein, 1958). 
Huber and Pengelly (1977) relate that this 
record is based upon a single specimen in 
the U.S. National Museum bearing the label 
"Bred from nests oWorytes (s.l.) from Hun- 
tington, L.L Cocoon March 24, 1924. em. 
April 30, 1924. S. C. Bridwell." Hoplisoides 
costalis is a ground-nesting, membracid- 
provisioning nyssonine (Evans, 1 966), while 
M. mixta, as presented above, nests in de- 
caying wood, provisions with cicadellids and 
delphacids, and is in the subfamily Pem- 
phredoninae. Thus Krombein's (1958) re- 
cord combined with that presented here im- 



plies a host range for E. viridicyaneus that 
is broad in both the systematic and behav- 
ioral/ecological senses. Observations of sand 
grains trapped in the coarse integumentary 
punctations of curated specimens o^ E lam- 
pus spp. (Huber and Pengelly, 1977; Kur- 
czewski and Kurczewski, 1970) support the 
suggestion that the host pool of North 
American Elampus spp. includes at least 
some ground-nesting species. Spooner 
(1948), summarizing the European litera- 
ture, and Moczar (1967) together listed three 
species in the genus Mi mesa as hosts of two 
European Elampus species. These three Mi- 
mesa species, and indeed all the members 
of the genus for which biological informa- 
tion exists, construct nests in the ground 
(Bohart and Menke, 1976). Thus, world- 
wide, the genus Elampus does parasitize 
both wasps that excavate nests in rotting 
wood and wasps that nest in the ground. 
The genus Mimesa is closely related to Mi- 
mumesa, both being members of the sub- 
tribe Psenina (Bohart and Menke, 1976). 

The significance of the female-biased sex 
ratio of emerging £". viridicyaneus is unclear. 
Museum collections of this and other Elam- 
pus species generally include approximately 
equal numbers of both sexes (Bohart and 
Kimsey, 1982; Huber and Pengelly, 1977). 
Highly female-biased sex-ratios for single 
batch rearings of chrysidid parasites are not 
uncommon (e.g. Krombein, 1967; Medler, 
1964) and sex ratios may vary between sites 
(Krombein, 1967). 


We thank the following taxonomists for 
their identifications: R. M. Bohart (Spheci- 
dae), L. S. Kimsey (Chrysididae), Depart- 
ment of Entomology, University of Cali- 
fornia, Davis; J. P. Kramer (Homoptera), 
Systematic Entomology Laboratory, Agri- 
cultural Research Service, USDA; E. I. 
Schlinger (Sciaridae), Department of En- 
tomological Sciences, University of Cali- 
fornia, Berkeley. We are grateful also to R. 

M. Bohart and G. W. Frankie (Department 
of Entomological Sciences, University of 
California, Berkeley) for critical readings of 
earlier drafts of the manuscript. This ma- 
terial is based in part upon work supported 
under a NSF Graduate Fellowship to JAR 
and by the Division of Entomology and Par- 
asitology, U.C. Berkeley (JKG). 

Literature Cited 

Bohart, R.M. and A. S. Menke. 1976. Sphecid Wasps 
of the World. Univ. Calif. Press, Berkeley. 695 pp. 

Bohart, R. M. and L. S. Kimsey. 1982. A synopsis 
of the Chrysididae in America north of Mexico. 
Mem. Am. Entomol. Inst. 33: 1-266. 

Evans, H. E. 1966. The Comparative Ethology and 
Evolution of the Sand Wasps. Harvard University 
Press, Cambridge, Mass. 526 pp. 

Gumey, A. B. 1951. The nesting habits oi Mimesa 
{Mimumesa) nigra (Packard). Proc. Entomol. Soc. 
Wash. 53: 280. 

Huber, J. T. and D. H. Pengelly. 1977. A revision of 
the genus Elampus Spinola {Notozus Auctt.) (Hy- 
menoptera: Chrysididae) in America north of 
Mexico. Proc. Entomol. Soc. Ont. 108: 75-137. 

Iwata, K. 1938. Habits of some Japanese pemphre- 
donids and crabronids (Hymenoptera). Mushi 1 1: 

Krombein, K. V. 1958. Hymenoptera of America 
North of Mexico. Synoptic Catalogue. First sup- 
plement. U.S. Dept. Agric, Agric. Monogr., 
Washington, D.C. 305 pp. 

. 1967. Trap-nesting Wasps and Bees: Life 

Histories, Nests and Associates. Smithsonian Press, 
Washington, D.C. 570 pp. 

Kurczewski, F. E. and E. J. Kurczewski. 1970. An 
annotated list of cuckoo- wasps from Erie County, 
Pennsylvania (Hymenoptera: Chrysididae). Proc. 
Entomol. Soc. Wash. 72: 190-201. 

Medler, J. T. 1964. Parasitism of Eumeninae by cuck- 
oo wasps in trap-nests in Wisconsin. Proc. Ento- 
mol. Soc. Wash. 66: 209-215. 

Moczar, L. 1967. Hymenoptera III. Chrysidoidea. 
Fauna Hungariae 86: 1-1 18. 

Petit, J. 1 979. Note sur Mimumesa sibiricana R. Bo- 
hart (Hym. Sphecidae). Lambillionea 79: 9-14. 

Spooner, G. M. 1948. The British species of psenine 
wasps (Hymenoptera: Sphecidae). Trans. R. Ento- 
mol. Soc. Lond. 99: 129-172. 

Tsuneki, K. 1959. Contributions to the knowledge of 
the Cleptinae and Pseninae faunae of Japan and 
Korea (Hymenoptera, Chrysididae and Spheci- 
dae). Mem. Fac. Lib. Arts, Fukui Univ. (2, Nat. 
Sci.) 9: 1-78. 

89(2), 1987, pp. 356-358 


Karl V. Krombein 

Department of Entomology, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560. 

Abstract.— The Motschulsky types of Bethylidae were studied. Dolus Motschulsky is 
synonymized with Epyris Westwood by designation ofpolitus Motschulsky as type-species; 
the Egyptian politus and apicalis Motschulsky are new combinations in Epyris; the Cey- 
lonese subnitidus Motschulsky and opacicollis Motschulsky are new combinations in Ho- 
lepyris Kieffer, and the former species is synonymized under the latter. Lectotypes are 
designated for politus and opacicollis. The Ceylonese Homalusl amplipennis Motschulsky 
is a new combination in Holepyris. The Ceylonese Goniozus montanus Kieffer, replace- 
ment name for Bethylus distigma Motschulsky not Thomson, is confirmed as a species 
of Goniozus. 

The Russian coleopterist Motschulsky 
published a catalog of Ceylonese insects in 
three parts and a supplement ( 1 8 6 1 -6 5). He 
listed therein previously known species and 
described a number of new insects. Most of 
the new species were Coleoptera but he also 
included some species of Hymenoptera and 
other orders. The type series are in the Zoo- 
logical Museum, Moscow State University, 

Motschulsky described one genus and six 
species of Bethylidae in this catalog of Cey- 
lonese species, four species from Ceylon and 
two from Egypt. The Ceylonese insects came 
from J. Nietner, a coffee planter and ama- 
teur coleopterist. The specimens are not in 
good condition, some having suffered dam- 
age from dermestids or breakage so that ap- 
pendages are often lacking. The specimens 
are glued on small cards. One specimen of 
each species bears a small hand-written la- 
bel "Type" and a larger label bearing Mot- 
schulsky's hand-written identification label. 
The identification labels of the Ceylonese 

species also bear a notation such as "I. or. 
Ceyl. Mt. N.E." indicating that this speci- 
men came from the Oriental island of Cey- 
lon from "Montagnes de Nura-Ellia." Spec- 
imens that I consider to be syntypes do not 
bear identification labels but were placed in 
the collection next to the specimen bearing 
the "type" label. Motschulsky did not des- 
ignate types in his paper. In the two species 
containing several syntypes, I have selected 
as lectotype the specimen bearing the "type" 

Motschulsky Hsted the localities from 
which the bethylids were described as 
"Montagnes de Nura-Ellia" and as "Mont 
Patannas." Nuwara Eliya is a town at an 
altitude of some 1800 m. Two mountains 
nearby are Hakgala and Pidurutalagala at- 
taining elevations of about 2100 and 2500 
m respectively. Patannas is not a mountain; 
patanas are areas of montane grassland oc- 
curring at altitudes above 500 m. 

I am publishing the following synonymic 
notes in advance of a revisionary study of 



the Ceylonese Bethylidae so the informa- 
tion can be included in a world catalog of 
Bethylidae in preparation by Gordon Gordh. 

Subfamily Epyrinae 
Dolus Motschulsky 

The genus Dolus (1863: 27) has been a 
puzzle since its description. Dalla Torre 
(1898: 536) assigned it to the Dryininae; 
(Motschulsky placed all his bethylids under 
the heading Dryinides). Ashmead (1902: 
272) placed it with question as a synonym 
of Mesitius sensu Ashmead not Spinola. 
Kieffer (1908) omitted it in his treatment of 
Bethylidae in Genera Insectorum, but in- 
cluded it as a questionable [zweifelhafte] ge- 
nus in his monograph of the family in Das 
Tierreich (1914). 

Motschulsky described Dolus for four new 
species, two from Ceylon, subnitidus and 
opacicollis, and two from Egypt, politus and 
apicalis. The first two species are congeneric 
with species placed in Holepyris Kieffer, 
1904, and the last two with species placed 
in Epyris Westwood, 1832. In the interest 
of nomenclatorial stability I designate pol- 
itus as the type-species of Dolus, thus sink- 
ing that genus as a synonym of Epyris. 

Dolus politus and apicalis are new com- 
binations in Epyris. The former species was 
described from three conspecific females, 
one of them on a single card bearing "type" 
and identification labels, the other two fe- 
males on a single card without labels. I des- 
ignate as lectotype the specimen bearing the 
"type" label. Dolus apicalis is based on a 
unique female. Neither species is known to 
occur in Sri Lanka. 

It should be noted that Epyris apicalis 
Smith, 1874, becomes a secondary hom- 
onym with the transfer of apicalis Mot- 
schulsky to Epyris. I shall not propose a 
substitute name inasmuch as a junior syn- 
onym may be available for apicalis Smith. 

Holepyris opacicollis (Motschulsky), 
New Combination 

Dolus opacicollis Motschulsky, 1863: 28. 

Dolus subnitidus MoXschwX^ky, 1863: 27-28. 
New Synonym. 

Dolus opacicollis was described from three 
conspecific females from "Montagnes de 
Nura-Ellia et Patannas." One specimen 
bears a label "type" and another label on 
which Motschulsky wrote his identification 
and "I. or. Ceyl. Mt. Pat." Two unlabeled 
females standing next to this specimen are 
syntypes; we know that Motschulsky had a 
series of opacicollis for he gave the length 
as 1-/5-^/4 lines. I designate as lectotype the 
specimen bearing the "type" and identifi- 
cation labels. Dolus subnitidus was de- 
scribed from a single female from "Mon- 
tagnes de Nura-Ellia"; it has suffered some 
dermestid damage. 

The minor differences between the two 
species noted by Motschulsky fall within the 
normal range of variation of this taxon. The 
specimen of subnitidus has the head some- 
what less dull than is typical so I place it as 
a synonym oi opacicollis. I have two females 
collected in Udawattakele Sanctuary, Kan- 
dy, by the Smithsonian's Ceylon Insect 
Project, and a third labeled just Ceylon, col- 
lected by G. H. K. Thwaites, the first Di- 
rector of the Royal Botanic Garden at Per- 
adeniya near Kandy, and accessioned by the 
British Museum in 1867. These three spec- 
imens are very similar to the type of opa- 

Holepyris amplipennis (Motschulsky), 
New Combination 

Homalusl amplipennis Motschulsky, 1863: 

Perisemus amplipennis Dalla Torre, 1898: 


The unique type is a male from "som- 
mites du Mont Patannas." It is not the op- 
posite sex of opacicollis (Motschulsky). 

Subfamily Bethylinae 
Goniozus montanus Kieffer 

Bethylus distigma Motschulsky, 1863: 26- 
27. (Preoccupied in Goniozus by distig- 
mw5 Thomson, 1861.) 



Goniozus montanus Kief[QT, 1908: 15 (New 
name for distigma Motschulsky.) 

The unique type is a female from "Mon- 
tagnes de Nura-Ellia." Motschulsky wrote 
bistigma rather than distigma on his label. 
I can confirm Kieffer's transfer of this species 
to Goniozus. 


I am grateful to my colleague Alexander 
V. Antropov, Moscow State University, 
USSR, for lending the Motschulsky mate- 
rial for study. I thank also Curtis W. Sa- 
brosky. Systematic Entomology Labora- 
tory, U.S. Department of Agriculture (ret.). 

for reviewing the manuscript and making 
helpful suggestions. 

Literature Cited 

Ashmead, W. H. 1902. Family XXXII. -Bethylidae. 

In Classification of the fossorial, predaceous and 

parasitic wasps, or the superfamily Vespoidea. Can. 

Entomol. 34: 268-273, 287-290. 
Dalla Torre, K. W. von. 1898. Chalcididae at Proc- 

totrupidae. Catalogus Hymenopterorum 5: 1-598. 
Kieffer, J. J. 1908. Bethylidae. //; Wytsman, Genera 

Insectorum 76: 1-50. 

. 1914. Bethylidae. Das Tierreich 41: 1-595. 

Motschulsky, V. de. 1863. VI: Hymenopteres. In Es- 

sai d'un catalogue des insectes de Tile Ceylan. Bull. 

Soc. Imp. Nat. Moscou 36: 1 1-73. 


89(2), 1987, pp. 359-362 


Louise M. Russell 

Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, BARC- 
West, Beltsville, Maryland 20705. 

Abstract. —The beech mealybug, Peliococcus serratus (Ferris), lives on Fagus grandifolia 
J. F. Ehrh. in eastern North America. In Maryland the species has two generations a year. 
Adult females settle on the bark of tree trunks and form a covering ovisac in which eggs 
are deposited from June until August and October through November. Eggs laid in summer 
hatch in about 7-14 days while those deposited in the fall overwinter. Mealybugs and 
eggs are destroyed by adverse weather conditions, parasitoids, and predators. Annotated 
citations are given to literature on the species. 

This article portrays, for the first time, 
the biology and habits of the beech mealy- 
bug, Peliococcus serratus (Ferris), in North 
America. It supplements the meager data 
available on a scale insect that lives on 
American beech, Fagus grandifolia J. F. 
Ehrh. (= F. americana Sweet), a valued for- 
est and shade tree. The mealybug merits 
study because of general interest in the ecol- 
ogy of indigenous insects and because it adds 
to the little that is known of the life cycles 
of most mealybugs native to the Nearctic 
fauna. The beech mealybug occurs sparingly 
and would achieve pest status only under 
greatly changed conditions. Although P. 
serratus apparently is restricted to Fagus 
(Fagaceae) in North America, it was re- 
ported from Corylus avellana L. (Betula- 
ceae) in Italy by Tranfaglia (1976). 

Literature on P. serratus is limited to the 
following annotated citations: 

Phenacoccus serratus FQTTis, 1925: 231-232 
(description, 3 collections); Trimble, 
1928: 43 (recorded as rare); Friend, 1932: 
596 (ovisacs described); Britton, 1933: 
375 (recorded); Rau, 1942: 124 (location 

of immatures, attended by ants, reared 
parasites Homalotylus sp. and Leptomas- 
tidea sp.); Herting and Simmonds, 1972: 
1 1 8 (rerecorded parasites listed by Rau). 
Peliococcus serratus (Ferris), Ferris, 1950: 
1 1 8 (redescription, 2 additional collec- 
tions); Baker, 1972: 99 (erroneously re- 
corded from birch); Tranfaglia, 1976: 
134-136 (redescribed from Corylus av- 
ellana in Italy, biology, habits, attended 
by ants, Crematogaster sp.), 1981: 9 (re- 
ferred to 1976 report); Drooz, 1985: 94 
(distribution in U.S.). 


My study of Peliococcus serratus extend- 
ed from October 1977 through September 
1986 in a forested area in Silver Spring, 
Montgomery County, Maryland. About 50 
trees were scrutinized to a height of 4.5 m 
with the unaided eye and to 9.1 m with 
binoculars. Tree trunks harboring mealy- 
bugs or ovisacs measured 15.2-20.3 cm in 
diameter at base. Insects or ovisacs were 
located 0.3-2.4 m from the ground on tree 
trunks. Mealybugs without ovisacs were ob- 



Fig. 1 . Ovisacs of Peliococcus senatus in knothole on bark of tree trunk. 

served on leaves and twigs 2.4 m above 
ground although they unquestionably were 
present at a much greater height. Trees were 
examined monthly all year and, if infested, 
usually were scrutinized weekly or daily 
April through November. Mealybugs or 
ovisacs were found on only four trees, one 
of which was located 1 km from the others 
and on which ovisacs were found only in 
1986. The other trees were in close prox- 
imity, one being 3.6 m from two that were 
1.5 m apart. Mealybugs or ovisacs were 
present on one tree each of the 9 years, on 
one in 1981, 1983, 1985, 1986, and on the 
other only in 1985. Tops of the two closest 
trees intertwined. Specimens were not seen 
from October 1983 to November 1984, the 
longest period in which mealybugs or ovi- 
sacs were not observed. After examination 
of the trees, immatures, adult females, and 
ovisacs were brought into the laboratory for 
further study. 


Ovisacs are formed gradually in 3-6 days 
and when complete are subrectangular, 5- 
8 mm long, and pure white (Fig. 1). Those 
formed in the summer tend to be smaller 
than those made in the fall. The outside is 
rather feltlike, the inside is fluffy and fila- 
mentous, and they are thicker dorsally than 

When females are ready to oviposit, they 
crawl down the tree trunk from the tree can- 
opy and usually settle in a knothole, scar, 
crevice or other rough, protecting place. Oc- 
casionally, however, they settle on smooth 
bark or move to a second location even after 
the ovisac is formed. At this time the fe- 
males are deep pink or dark purple and have 
a thin, white, waxy substance arranged in 
transverse rows across the body. A large fe- 
male in this condition on 6 October 1985 
had a little white fuzzy material on the body. 



Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 







Fig. 2. Seasonal history of Peliococcus senatus in Maryland. Solid line denotes first generation, broken line 
second generation. 

more posteriorly than anteriorly, on 7 Oc- 
tober. The buildup continued through 11 
October and on 1 2 October the ovisac was 
complete. This female started laying eggs 3- 
4 days after the ovisac was completed. On 
30 October the ovisac was removed and 
brought to the laboratory. The shrivelled, 
but soft body of the female was located at 
one end and 225 eggs filled the remainder 
of the ovisac. The female died, and a few 
eggs hatched within 3 days but most of them 
collapsed. The development of this female, 
ovisac, and eggs was analogous to that of 
other specimens. 

Eggs deposited in October and November 
overwinter and hatch from late April to ear- 
ly May of the following year (Fig. 2). Crawl- 
ers travel up tree trunks and onto the 
branches where they and later stages occur 
on the lower surface of leaves and on bark 
in twig axils. In this location adults develop 
in approximately 4 weeks. From late June 
into July and occasionally into August, fe- 
males come down the tree trunks, form ovi- 
sacs, and lay eggs which hatch in 7-14 days. 
Only 55-60 eggs were found in summer ovi- 
sacs and only 60 eggs were counted in 1 
ovisac formed in October. However, 200- 
225 eggs were found in most ovisacs formed 
in the fall. Initially the eggs are bright yellow 
and some remain yellow, but the majority 

turn pink, and some, presumably dead, be- 
come black. 

Descending and ovisac-forming females 
were not found in September, but second 
stage specimens and adult females were 
present on leaves and twigs at this time. 
Owing to the location of the insects in the 
tree canopy, I was unable to determine the 
length of the first and second stages. Adult 
males were not found, but 6 male cocoons, 
possibly of P. serratus because no other co- 
coon-forming species was present, were col- 
lected in July and September. 

Rau ( 1 942) noted that P. serratus was oc- 
casionally attended and shielded by ants, 
and Tranfaglia ( 1 976) wrote that the species 
is habitually cultured by a formicid for the 
production of honeydew. I have not ob- 
served these activities although I have seen 
ants on the same tree trunks as the mealy- 

P. serratus exists under hazardous con- 
ditions. Insects and ovisacs are dislodged 
by strong winds and heavy rains almost in- 
stantly, and they are attacked by hymenop- 
terous parasites (Rau, 1942; Herting and 
Simmonds, 1972). I have observed adult 
hymenopterous parasites, coccinellid and 
chrysopid larvae on tree trunks near mealy- 
bugs, syrphid larvae near and inside mealy- 
bugs, and both hibernating and predaceous 



mites in ovisacs. Ovisacs also are some- 
times covered with fungus. Ovisacs, females 
and eggs may be totally destroyed by one or 
more of these agents within 8 hours. 

The distribution of P. serratus as indi- 
cated by literature citations and/or the pres- 
ence of specimens in the National Collec- 
tion of Coccoidea at Beltsville, Maryland is 
as follows: 

Canada. -Ontario (Ferris, 1925, 1950). 

United States. — Connecticut: (Friend, 
1932), (Britton, 1933), USNM; District of 
Columbia: USNM; Maryland: (Ferris, 
1950), USNM; Massachusetts: USNM; New 
Hampshire: USNM; New Jersey: USNM; 
New York (Ferris, 1925, 1950), USNM; 
Ohio: (Ferris, 1925, 1950), USNM; Penn- 
sylvania: (Trimble, 1928), USNM; Tennes- 
see: USNM; Virginia: (Drooz, 1985), 

Europe. — Italy: Campania (Tranfaglia, 


I gratefully acknowledge the assistance of 
members of the Systematic Entomology 
Laboratory, BBII, Agricultural Research 
Service as follows: J. M. Kingsolver for re- 
viewing the manuscript, D. R. Miller for 
identifying the mealybugs, and M. B. Sto- 
etzel for photographing the ovisacs and giv- 
ing helpful suggestions. I thank J. A. Da- 
vidson, University of Maryland, College 
Park, Maryland, and Michael Kosztarab, 

Virginia Polytechnic Institute and State 
University, Blacksburg, Virginia for review- 
ing the manuscript. 

Literature Cited 

Baker, W. L. 1972. Eastern Forest Insects. U.S. Dept. 
Agr. For. Serv. Misc. Publ. 1 175. 642 pp. 

Britton, W. E. 1933. Entomological features of 1932. 
Conn. Agr. Exp. Sta. Bull. 349: 369-381. 

Drooz, A. T. 1985. Insects of Eastern Forests. U.S. 
Dept. Agr. For. Serv. Misc. Publ. 1426. 608 pp. 

Ferris, G. F. 1925. Notes on Coccidae XI (Hemip- 
tera). Can. Entomol. 57(9): 228-234. 

. 1950. Atlas of the Scale Insects of North 

America, (sen 5) V: vii + 278 pp. Stanford Uni- 
versity Press, Stanford, California. 

Friend, R. B. 1932. A new scale insect on beech. 
Conn. Agr. Exp. Sta. Bull. 338: 596. 

Herting, B. and F. F. Simmonds. 1972. A Catalogue 
of Parasites and Predators of Terrestrial Arthro- 
pods. Commonwealth Inst. Biol. Control. 210 pp. 

Rau, G. J. 1942. The Canadian apple mealybug, 
Phenacoccus aceris Signoret, and its allies in 
northeastern America. Can. Entomol. 74(7): 1 18- 

Tranfaglia, A. 1976. Studi sugli Homoptera Coccoi- 
dea IV. Su alcune cocciniglie nuove o poco con- 
osciute per I'ltalia (Coccidae, Eriococcidae, Pseu- 
dococcidae). Boll. Lab. Entomol. Agr. "Filippo 
Silvestri" Portici 33: 128-143. 

. 1981. Studi sugli Homoptera Coccoidea V. 

Notizie morphosistematiche su alcune specie di 
cocciniglie con descrizione di tre nuove specie di 
Pseudococcidi. Boll. Lab. Entomol. Agr. "Filippo 
Silvestri" Portici 38: 3-28. 

Trimble, F. M. 1928. Scale insects of Pennsylvania 
(Homop.: Coccidae). Entomol. News 39(2): 42- 

89(2), 1987, pp. 363-366 




R. G. Fennah 

Commonwealth Institute of Entomology, % British Museum (Natural History), London 
SW7 5BD, United Kingdom. 

Abstract.— Tht genus Gastrinia St^l (Homoptera: Fulgoroidea) is redefined and trans- 
ferred from Issidae to Nogodinidae, and a new subfamily is proposed for its reception. 
Supplementary characterization is provided for the type species, G. vaginata StM, and a 
new species, G. phidon, is described from Minas Gerais, Brazil. 

StM (1866a) referred his genus Gastrinia 
to his new subfamily Tropiduchida, and 
Melichar (1914) placed it in his new tribe 
Hiraciini. In 1982, the present writer ex- 
cluded both Hiracia and Gastrinia from the 
Tropiduchidae and transferred them to the 
Issidae. Further study of these genera has 
confirmed the conclusion that Hiracia is an 
issid, but has revealed that Gastrinia differs 
from both Issidae and Nogodinidae in the 
proportions of the abdominal laterotergites 
and pleurites, the shape of the spiracles, and 
the structure of the aedeagus and the ovi- 
positor. However, the number and arrange- 
ment of the post-tibial and post- tarsal spines, 
the relationship of the basal sclerite of the 
tegmen with the base of the clavus, the fre- 
quency of branching and direction of the 
wing- veins and the proportions of the gen- 
ital styles are broadly similar to correspond- 
ing features to be found in Nogodinidae, 
though not in combination. In view of the 
magnitude of the differences that separate 
Gastrinia from all other Nogodinidae, a new 
subfamily is now erected for its accom- 
modation, characterized as follows. 

Family Nogodinidae 
Gastriniinae, New Subfamily 

Habitus broad and depressed. Pronotum 
much wider than head, with 2 carinae be- 

tween eye and tegula. Tegmen with basal 
cell absent, basal sclerite narrow and straight. 
Post-tibia with 10-12 long teeth apically; 
basal metatarsal segment rather long, with 
coarse teeth apically. Abdominal spiracles 
elongate, narrow, embedded in lateroter- 
gites strongly obliquely to lower margin; la- 
terotergites IV-VI not much longer than 
broad, and much broader than pleurites. 
Genital style elongate, with narrow sides and 
a short process dorsally at apex. Aedeagus 
with a short basal collar. 

With the exception of the number of api- 
cal teeth on the post-tibia, none of the above 
characters occurs in the nominate subfam- 

Gastrinia St^l 

Gastrinia Stai, 1859: 319. Type species, 
Gastrinia vaginata StSl. 

Vertex pentagonal. Frons strongly de- 
flexed ventrocaudad, median disc promi- 
nent at base, sublateral carinae curving 
mesad basally and not meeting anterior 
margin of vertex. Post-clypeus with sides 
very narrow, and visible in anterior view. 
Antennae with second segment globose or 
nearly so, infuscate except at apex, with pal- 
lid disc sensilla, microsetae minute, mac- 
rosetae apparently absent. Ocelli present. 
Rostrum distally overlapping post-coxae. 



Figs. 1-13. 1-12, Gasthnia phidon. 1, Head and thorax, dorsal view. 2, Head, thorax and base of tegmen, 
side view. 3, Frons and clypeus, anteroventral view. 4, Basal part of third antennal segment. 5, Tegmen. 6, Apex 
of post-tibia and basal 2 segments of metatarsus, ventral view. 7, Laterotergite with spiracle (It), pleurite (pi) 
and lateral part of stemite (s) of fourth abdominal segment. 8, Ninth abdominal segment and anal segment of 
male, anterodorsa! view. 9, Male genitalia, right side. 10, Aedeagus, ventral view. 11, Apex of aedeagus, left 
side, with spinose process slightly displaced outward. 12, Phallobase (suspensorium), posteroventral view, with 
basal part of phallus emerging from orifice. 13, Gastrinia vaginata. Female genitalia, right side. 

Pronotum much wider than head, and 
broader than long in middle (about 4:1), 
disc slightly produced anteriorly, posterior 
margin shallowly concave; 2 carinae be- 
tween eye and tegula. Mesonotum almost 
flattened, with lateral carinae strongly con- 
verging cephalad. Tegula large, exposed, and 
abruptly decurved. Post-trochanters rock- 
ing mesad-laterad. Post-tibia with 4 spines 
laterally, 10-12 apically. Basal metatarsal 
segment with 10-13 coarse teeth an a tract 
of sparse long setae; second segment with 2 
teeth; apical margin between them shallow- 
ly convex. Tegmen coriaceous, distally sub- 
tectiform, costal and commissural margins 
parallel, and apical margin angulate, scarce- 
ly surpassing abdomen, Sc+R scarcely bent 
at junction with M basally, M and Cu, sim- 
ple to beyond level of apex of clavus, an 
oblique row of veinlets from apex of clavus 

to M near apical margin. Basal cell obsolete, 
basal sclerite narrow, straight. Wing-tuck- 
ing process absent. Wing ample, with apical 
margin not deeply cleft, and costal margin 
straight. Venation regular, not bent ante- 
riorly in distal half, M with 2-3 branches 
apically, Cu, with 3-5 branches, apical cells 
almost parallel-sided, much longer than 
broad. Abdomen strongly dorsoventrally 
compressed; tergites narrowly divided by 
membrane along middle line, stemites not 
divided medially, laterotergites large, little 
longer than broad, and several times broad- 
er than pleurites; spiracles greatly elongated, 
embedded in laterotergites strongly 
obliquely to lower margin. Pygofer rela- 
tively short, with a small angulate medio- 
ventral process. Aedeagus long, straight and 
tubular, supported in basal quarter by a short 
tubular suspensorium. Genital style elon- 



gate, with narrow sides and a short process 
dorsally at apex. Anal segment of female 
elongate, with lateral margins deep and de- 
cumbent. Ovipositor elongate, slender and 

Gastrinia vaginata StSl 
Fig. 13 

Gastrinia vaginata StM, 1859: 319. 
Hiracia lacerdae Signoret, 1861: 57. (syn.) 
StM, 1866b: 393. 

Female (supplementary description).— 
Length with tegmen, 15 mm; tegmen, 1 1.8 

Vertex broader than long (almost 1.2:1). 
Frons longer medially than broad (1.2:1), 
medially ecarinate. Rostrum with apical 
segment longer than broad in side view (6.6: 1). 
Basal metatarsal segment with 1 3 teeth api- 
cally, longer dorsally in middle line than 
wide between tips of outermost teeth (1.9:1), 
and than length of outer apical spine mea- 
sured from level of apex of mid-dorsal mar- 
gin (2.1:1). Second metatarsal segment lon- 
ger dorsally than wide between apical spines 
(about 1.2:1). Tegmen longer than broad 
(3.0: 1), cell PCu longer than common claval 
vein (about 1.3:1). 

Tegmen with 2 narrow oblique bands from 
Sc to basal angle of clavus and from Sc to 
apex of clavus, reddish brown. 

Ninth abdominal segment very short dor- 
sally, overlapped by seventh and eighth ter- 
gites. Anal segment in side view longer than 
deep dorsoventrally (about 4.5:1), narrow- 
ing and weakly ascending distad and nar- 
rowly rounded apically; sides decumbent, 
ventral surface deeply hollowed to ensheath 
ovipositor; anal foramen situated at middle. 
Ovipositor slender, much longer than deep 
in side view (about 18:1), of almost equal 
width throughout, and weakly ascending 
distad. Seventh stemite short, not quite 
overlapped by sixth, with hind margin 
weakly concave; area between this and base 
of ovipositor apparently consisting of tough 
membrane, and with a weak median ridge. 

Material examined. 1 $, Brazil, Bahia (coll. 

This specimen, determined by St^l as be- 
longing to his G. vaginata, (Melichar 1914: 
210) agrees with the description and illus- 
tration of Hiracia lacerdae given by Sig- 
noret, and also agrees with the description 
of Gastrinia vaginata St^l based on a female 
from Bahia, and with the figure of G. (= Am- 
fortas) vaginata StSl given by Melichar 
(1914: fig. 30). It is one of two specimens 
from Bahia in the Signoret Collection in the 
Naturhistorisches Museum, Vienna, and the 
only one that agrees with the original de- 
scription of H. lacerdae. It is here desig- 
nated as the lectotype, and has been labelled 
as such. 

The status of the second female from Ba- 
hia in the Signoret Collection, labelled "Ba- 
hia, Coll. Signoret, det. Signoret," and con- 
sidered to be specifically distinct from G. 
vaginata by Melichar must remain unset- 
tled until further material is available for 
study. It is not mentioned by Signoret, and 
its size and tegminal marking are not cov- 
ered by the description given for H. lacer- 

The length/greatest breadth ratios of the 
frons, apical segment of the rostrum, basal 
metatarsal segment, second metatarsal seg- 
ment and tegmen are 1.2:1, 5.1:1, 1.6:1, 1.2:1 
and 2.9:1, respectively, and the ratio of 
lengths of tegminal cell PCu/common claval 
vein, about 1.3:1. 

Gastrinia phidon Fennah, New Species 

Figs. 1-12 

Male. — Length with tegmen, 9.0 mm; teg- 
men, 7.0 mm. Vertex broader than long 
(slightly less than 1.1:1). Frons longer me- 
dially than broad (1.2:1), feebly medially 
carinate in distal half. Rostrum with apical 
segment longer than broad in side view 
(4.9:1). Basal metatarsal segment with 12 
teeth apically, longer dorsally in middle line 
than wide between tips of outermost teeth 
(1.5:1), and than length of outer apical spine 



measured from level of apex of mid-dorsal 
margin (1.9:1). Second metatarsal segment 
longer dorsally than wide between apical 
spines (1.1:1). Tegmen longer than broad 
(2.6:1), cell PCu longer than common claval 
vein (2.6:1). Wing with Sc 2-3 branched, R 
simple, M and Cu, each with 3 branches, 
PCu forked basad of level of Cu, fork. 

Dorsally yellowish brown, mottled with 
paler spots. Mesonotum with a suffusion in 
each lateral field and a small round spot near 
each posterolateral margin of disc, piceous. 
Lower surface of thorax and legs more or 
less reddish brown, mottled with pale round 
spots; abdomen ventrally pale brown with 
fuscous spots at base of setae. Tegmen light 
yellowish brown, with light reddish brown 
markings in depressions; venation coarse, 
with supernumerary irregular veinlets, 
mostly concolorous, but with a few spots on 
main veins in corium and clavus and a larg- 
er spot close to Ml +2 subapically, dark red- 
dish brown or piceous. Wing uniformly 
dilute fuscous, apical veins almost concol- 
orous, but orange brown on margin at apices 
of R, M and Cu,; longitudinal veins slightly 

Anal segment of male relatively large, in 
dorsal view widest near base, with lateral 
margins curving to deeply rounding apex; 
anal orifice in distal half; anal style very 
short. Pygofer with lateral margins sinuate, 
dorsolateral angles obscure; medioventral 
process broader at base than long. Aedeagus 
porrect caudad, with lateral margins of 
phallobase meeting at an obtuse angle below 
phallus in basal quarter; a pair of delicate 
spinose processes ventrolaterally near apex 
of aedeagus, each directed cephalad and lying 
close against ventrolateral surface. Genital 
style in side view about 3.5 times as long 
as wide, and widest near base, stiffened with 
a shallow ridge internally, extending from 
base to apex; dorsal and ventral margins 
gradually converging distad in basal three- 
quarters, thence subparallel; apical margin 
shallowly rounded, shortly produced dor- 

sally in a peg-like process. Length, 9.0 mm; 
tegmen, 7.0 mm. 

Holotype 3. — Brazil: Minas Gerais, Pedra 
Azul, xii. 1970, (F. M. Oliviera) in British 
Museum (Natural History). 

This species differs from G. vaginata and 
the second specimen ofGastrinia in the Sig- 
noret collection in the ratio of the basal width 
of the frons to the width at the frontoclypeal 
suture (about 1.1:1 in G. phidon and 1.3:1 
in the other two), the ratio of length to width 
in the tegmen (2.6: 1 in G. phidon and 3.0: 1 
and 2.9:1 in the others) and the relative 
lengths of cell PCu and the common claval 
vein in the tegmen (2.6:1 in G. phidon and 
about 1.3:1 in the others). 

The name phidon is a classical personal 
name, and is used in apposition. 


I thank W. J. Knight (British Museum 
(Natural History)) for the privilege of study- 
ing the Fulgoroidea in his charge, A. Kal- 
tenbach (Naturhistorisches Museum, Vi- 
enna) for the loan of the female of Hiracia 
lacerdae Signoret referred by St^l to Gas- 
trinia vaginata Stdl and for supplying in- 
formation on a second female in the Sig- 
noret collection, and two anonymous 
reviewers for helpful comments on the 

Literature Cited 

Fennah, R. G. 1982. A tribal classification of the 

Tropiduchidae (Homoptera: Fulgoroidea), with the 

description of a new species on tea in Malaysia. 

Bull. Entomol. Res. 72: 631-643. 
Melichar, L. 1914. Monographic der Tropiduchiden. 

Verh. Naturforsch. Ver. Brunn 53: 1-145. 
Signoret, V. 1861. Description de quelques Hemip- 

teres nouveaux. Ann. Soc. Entomol. Fr. (4)1: 55- 

Stai, C. 1859. Novae quaedam Fulgorinorum formae 

speciesque insigniores. Berl. Entomol. Zeit. 3: 313- 

. 1866a. Hemiptera Homoptera Latr. Hemip- 

tera Africana 4: 1-276. 
. 1866b. Analectahemipterologica. Bed. Ento- 

mol. Z. 3: 319. 

89(2), 1987, pp. 367-368 


The Prevalence of Icosta americana (Diptera: Hippoboscidae) on 
Ruffed Grouse {Bonasa umbellus) in Wisconsin 

Little has been published regarding the 
distribution of Icosta americana (Leach) 
from ruffed grouse, Bonasa umbellus L., in 
Wisconsin. MacArthur (1948. Bull. Public 
Mus. Mil. 8: 367-440) reported americana 
from a "partridge," locality unknown and 
Bequaert (1955. Entomol. Am. 34-35(N.S.): 
1-611) reported infected grouse from three 
counties. Because of the lack of information 
from other areas of the state, our study was 

From 1968 through 1985, 206 ruffed 
grouse were collected by the senior author 
and his father from seven Wisconsin coun- 
ties and examined for americana. Preva- 
lence of infection in the samples was: 11/ 
80 Richland Co.; 1/98 Portage Co.; 0/17 
Adams Co.; 0/4 Chippewa Co.; 0/2 Forest 
Co.; 0/4 Monroe Co.; and 0/ 1 Waushara Co. 
Seven grouse from Richland Co. harbored 
one americana each and four birds carried 
two flies each, while the infected Portage 
Co. grouse had one. Infection rate in south- 
western Richland Co. (13.7%) was consid- 
erably higher than the approximately 1% in 
the more central Portage Co. 

In the 1984 and 1985 seasons Wisconsin 
hunters were asked to collect any flies seen 
on ruffed grouse. Six additional hippobos- 
cids were collected: two from Tremapealeau 
Co., and one each from Grant, Richland, 
Jackson, and Rusk counties. All of these are 
in the lower two-thirds of Wisconsin with 
the exception of Rusk which is in the upper 

The reason for this distribution is un- 
known and may be related to one or more 
ecological factors. Bequaert (op. cit.) sug- 
gested americana may not occur north of 
48° 10' latitude, approximately the Cana- 

dian border in central United States. Ben- 
nett (1961. Can. J. Zool. 39: 379-406) found 
only a "few" specimens o^ americana from 
more than 400 ruffed grouse in Ontario and 
63 ruffed grouse and spruce grouse {Can- 
achites canadensis L.) taken between July 
and November in Algonquin Park, Ontairo 
just slightly south of 48°10' north latitude. 

In Wisconsin there is a tension zone di- 
viding the state into two floristic provinces 
(Curtis, 1959, The Vegetation of Wisconsin. 
The University of Wisconsin Press, Madi- 
son, WI). The southwest third of the state 
contains southern hardwood forests with 
some prairie elements, and the northern half 
features coniferous-hardwood forests with 
a few boreal elements. The narrow tension 
zone separating each province contains 
members of each. 

Bequaert (op. cit.) records americana from 
ruffed grouse in Clark, Marathon, and Rusk 
counties. Although these counties are just 
north of the tension zone, our data suggest 
this fly is more common on grouse in or 
below the tension zone. 

Bequaert (op. cit.) believed that native 
galliform birds were the original hosts of 
americana, and that raptors later acquired 
the fly by preying on game birds. Converse- 
ly, Bennett (op. cit.) believed americana is 
primarily a parasite of birds of prey and 
occur only accidentally on galliformes. 

By definition, accidental parasites are rare, 
do not breed on their host, and remain with 
the host only for a short period of time. That 
13.7% of ruffed grouse from Richland Co. 
have this parasite argues against its being 
accidental on grouse. Also, in three in- 
stances female americana were captured 
alive and viable. When placed in a jar, they 



larviposited. Female americana carrying 
larvae and a male were collected as late as 
January 28. 

These data suggest americana is common 
on grouse in southern Wisconsin. This 
species most likely breeds on the host, and 
female americana may overwinter with lar- 
vae in utero. 

Voucher specimens of two americana 
from ruffed grouse are deposited in the Na- 
tional Museum of Natural History in Wash- 
ington, D.C. The remainder of the speci- 

mens are in the Museum collection at the 
University of Wisconsin-Stevens Point. 

We thank D. J. Taft for his help in col- 
lecting americana. 

Stephen J. Taft, Department of Biology, 
University of Wisconsin-Stevens Point, Ste- 
vens Point, Wisconsin 54481 and Susan 
Marcquenski, Wisconsin Department of 
Natural Resources, Box 7921, Madison, 
Wisconsin 53707. 


89(2), 1987, p. 368 

Book Notice 

The Insect and Spider Collections of the 
World, by Ross H. Amett, Jr. and G. A. 
Samuelson, assisted by John B. Heppner, 
Gordon M. Nishida, J. Charles Watt, and 
Robert E. Woodruff. E. J. Brill Publishers, 
New York, NY. 1986. 220 pp. Cost: 
$ 1 9.95 (paper with plastic comb binding). 

Part I has a list and description of 9 1 8 
public collections known to exist; 377 of the 
major collections are described in detail. 
They are arranged alphabetically by coun- 
try, state/province, and city. Every country 
in the world is listed— those without known 
collections are indicated. The mailing ad- 
dress of each museum is cited, and a four- 
letter coden is assigned to each. If a ques- 
tionnaire was returned, the names of cura- 
tors, phone numbers, details about the size 

and content of the collection, and data about 
primary types and special collections are 

Part II has a Hst and descriptions of 2 1 1 
private collections, arranged alphabetically 
by owner's name, with pertinent data. A 
coden is assigned to each collection. 

To systematists who borrow specimens 
for research and to collectors who would 
like to have their collections identified and 
used in research, this list will be most wel- 
come. (E. J. Brill, founded in 1683 in The 
Netherlands, now has an office in USA: Suite 
404, 225 W. 57th St., NYC 10019.) 

T. J. Spilman, Systematic Entomology 
Laboratory, BBII, Agricultural Research 
Service, U.S.D.A., % U.S. National Mu- 
seum NHB 168, Washington D.C. 20560. 


89(2), 1987, p. 369 


New Distributional and Rearing Records for 
Neotropical Flower Flies (Diptera: Syrphidae) 

The biology of most Neotropical flower 
flies is poorly known. In this paper, I report 
new information on the distribution, flower 
visitation, or larval habitat of 4 syrphids. 

Copestylum sexmaculatum (Palisot de 
Beauvois) (= C. pallens of authors). — First 
state record: Illinois, Pope County, Bell 
Smith Springs Rec. Area, 19 July 1975, 1 <5 
feeding on blossoms of Pycnanthemum ten- 
uifolium Schrad. (Labiatae). Wirth et al. 
(1965. In Stone et al., Cat. Diptera America 
North of Mexico, Agric. handbook 276: 602) 
and Thompson (1981. Mem. Entomol. Soc. 
Wash. 9: 138-139) reported this fly from 
North Carolina, Florida to Arizona, and the 
West Indies. 

Meromacrus panamensis Curran. — First 
United States record: Texas, Cameron 
County, Sabal Palm Grove Sanctuary, 14 
December 1984, 1 $ feeding on blossoms of 
Xylosma flexuosa (H.B.K.) O. Ktze. (Ha- 
courtiaceae). Thompson (ibid.: 157; 1985, 
pers. comm.) previously noted that this 
species is distributed from Mexico to Brazil. 

Meromacrus pratorum (Fabricius).— First 
rearing record: Puerto Rico, El Yunque Rec. 
Area, 1 <5 reared from larva collected in fluid 
and debris of upright flower bract of Heli- 
conia sp. (Musaceae) on 29 May 1981. Sack 
(1921. Arch. Naturgesch. (Abt. A) 87(3): 
144-145) reported that rat-tailed larvae of 
M. pratorum live in sediment of wet rot 
pockets of trees. His identification was ap- 
parently incorrect because Thompson (1981: 
157) found M. pratorum to be endemic to 
the West Indies. Snow (1958. Ecology 39: 

83-88) observed larvae of M. acutus (Fa- 
bricius) in decaying stumps filled with water. 
The larval habitat of M. pratorum in Puerto 
Rico more closely resembles that of Qui- 
chuana angustiventris (Macquart) reared 
from Heliconia bracts by Seifert and Seifert 
(1976. J. N.Y. Entomol. Soc. 94: 233-242; 
1979. Ecology 60: 462-467) than that of 
other Meromacrus species. Additional study 
is needed to determine the diversity of lar- 
val habitats used by species of Meromacrus 
and related genera. 

Polybiomyia macquarti Shannon. — First 
rearing record: Texas, Cameron County, Sa- 
bal Palm Grove Sanctuary, 2 9 reared from 
larvae collected from sap-soaked detritus in 
small rot pocket in trunk of Leucaena pul- 
verulenta (Schlecht.) Benth. (Leguminosae) 
on 14 December 1984. Other cerioidine 
species reared by Johnson (1893. Entomol. 
News 4: 91), Banks (1902. Proc. Entomol. 
Soc. Wash. 5: 310) and Maier (1982. Proc. 
Entomol. Soc. Wash. 84: 603-609) have 
similar larval habitats. 

I thank F. Christian Thompson (System- 
atic Entomology Laboratory, Agricultural 
Research Service, USDA) for identifying 
most of the syrphids. The stafl'of the Sabal 
Palm Grove Sanctuary, Brownsville, Texas 
provided valuable biological information. 

Chris T. Maier, Department of Ento- 
mology, The Connecticut Agricultural Ex- 
periment Station, P.O. Box 1106, New Ha- 
ven, Connecticut 06504. 


89(2), 1987. p. 370 


Anomalous Tubercle Patterns Found on Lymantria dispar (L.) 

Caterpillars in the Field in Maryland 

(Lepidoptera: Lymantriidae) 

The gypsy moth caterpillar is conspicu- 
ous by having a double row of colored spots 
along its dorsal surface during and after the 
fourth instar. The anterior 5 pairs are blue, 
while the posterior 6 pairs are a dull-to- 
bright red. These spots are dorsal tubercles 
that form the basal portion of hair tufts, and 
they represent outgrowths of the integument 
bearing numerous setal sockets (Traxler, J. 
1977. N.Y. Entomol. Soc. 85: 71-97). 

In the spring of 1986, I had occasion to 
observe gypsy moth caterpillars from near- 
ly 50 locations in Northern Maryland, 
amounting to several thousand collected, 
and many thousands more observed in the 
field either by myself, or by collaborators 
Kathleen Tatman, Kevin Boyd, and Anne 
Wieber. From these, 17 individual cater- 
pillars were collected having anomalous tu- 
bercle patterns. Eight had only the first 4 
anterior spot-pairs blue while the 7 poste- 
rior spot-pairs were red. Four individuals 
were missing one red spot (never the same 
one); three individuals were missing a pair 

of red spots (never the same pair), while one 
individual was missing 3 red spots. The final 
oddity was a caterpillar whose dorsal integ- 
ument was albino along its entire length on 
the left side, while the right side was a nor- 
mal mottled black. All dorsal tubercles were 
present, and of normal pigmentation, on this 
individual. All specimen were collected from 
6 locations in northern Baltimore County, 

All of the above caterpillars appeared 
normal except for their color patterns. The 
1 1 individuals not parasitized or diseased 
produced apparently normal pupae and 
adult females. If any of these patterns prove 
to be heritable, it could prove the basis of 
a marked strain of gypsy moths of potential 
utility in sterile-male release programs or 
for release-recapture ecological studies. 

Ralph E. Webb, Florist and Nursery Crops 
Laboratory, HSI, Agricultural Research 
Service, U.S.D.A., Belt svi lie, Maryland 

89(2), 1987, pp. 371-374 

Society Meetings 

925th Regular Meeting- October 2, 1986 

The 925th Regular Meeting of the Ento- 
mological Society of Washington was called 
to order by President E. M. Barrows in the 
Naturalist Center, National Museum of 
Natural History, at 8 p.m. on October 2, 
1 986. Twenty-one members and four guests 
were present. Minutes of the previous meet- 
ing were read. Membership Chairman, G. 
White, read the names of the following ap- 
plicants for membership: P. Adler, Clemson, 
South Carolina; C. Agnew, College Station, 
Texas; D. Haile, Reading, Pennsylvania; K. 
Hoffman, Clemson, South Carolina; E. Lip- 
pert, Guelph, Ontario, Canada; J. Mackley, 
Laredo, Texas; P. Newhouse, Silver Spring, 
Maryland; A. Zuccaro, Jr., Natchez, Mis- 
sissippi; A. Ritchie, Hamden, Connecticut; 
R. K. Robbins, Washington, D.C. 

Editor R. Gagne displayed the recently 
rediscovered Society seal embosser. This 
embosser, which may have been used in the 
past to emboss the seal on portfolios of 
greetings and other correspondence, will re- 
main with the Editor's office. 

R. Gagne showed slides of various ceci- 
domyiid galls on hickory including husk 
swellings caused by '"Cecidomyia" micicola. 
The discovery of this species, which has not 
been reported since its description in 1870, 
except for one additional record in 1906, 
allowed him to place the species in the genus 
to which it properly belongs. 

G. Steyskal showed a recent publication 
by Spencer and Steyskal on the agromyzid 
leaf miners of the United States. 

E. Barrows displayed an unusual day- 
flying trichopteran with dark wings and or- 
ange body. 

The speaker of the evening was F. Eugene 
Wood, University of Maryland who pre- 
sented a talk and video tape entitled "Den- 
izens of the Dark: Observations on Termite 
Biology and Management." 

Following the introduction of visitors, the 
meeting was adjourned at 9:10 p.m. 

Paul M. Marsh, Recording Secretary 

926th Regular Meeting— November 6, 1986 

The 926th Regular Meeting of the Ento- 
mological Society of Washington was called 
to order by President Edward M. Barrows 
in the Naturalist Center, National Museum 
of Natural History, at 8 p.m. on November 
6, 1986. Twenty-two members and thirteen 
guests were present. Corresponding Secre- 
tary R. G. Robbins read the names of the 
following applicants for membership: Paul 
E. Blom, Department of Entomology, Uni- 
versity of Idaho, Moscow; Paul Courneya, 
Harlingen, Texas; and Eugene G. Munroe, 
Ottawa, Ontario, Canada. 

President Barrows summarized the latest 
meeting of the Executive Committee, held 
on the 23rd of October. Among the topics 
covered were a proposed increase in dues 
for life memberships, an invitation to the 
Maryland Entomological Society to cospon- 
sor the annual banquet, use of name tags at 
all ESW meetings, revision and republica- 
tion of the Bylaws, and effiDrts to encourage 
young entomologists to become more active 
in the Society. 

The Nominating Committee Chairman, 
D. M. Anderson, presented the following 
nominations for 1987 officers: 
President-Elect- F. Eugene Wood 
Treasurer— Norman E. Woodley 
Corresponding Secretary — Robert G. 
Recording Secretary— Paul M. Marsh 
Editor— Raymond J. Gagne 
Associate Editor— no nominee 
Membership Chairman — Geoff'rey B. 
Program Chairman— Michael J. Raupp 
Custodian — Victor L. Blackburn 
Nominations will remain open until the 



December meeting when elections will take 

R. Gagne noted that there was no nom- 
inee for Associate Editor in the report of the 
Nominating Committee. He reminded the 
Nominating Committee that 1 987 would be 
his last year as Editor and that, according 
to a recently enacted bylaw (see Minutes for 
January 5, 1984, PESWS6: 974), the Nom- 
inating Committee is responsible for finding 
an Associate Editor, who would begin pro- 
cessing manuscripts about June in prepa- 
ration for the January 1 988 issue of the Pro- 

T. J. Spilman displayed a newly published 
reference. Insect and Spider Collections of 
the World, compiled by Ross H. Amett, Jr., 
and colleagues. 

The speaker for the evening was Jonathan 
A. Coddington, Associate Curator, Depart- 
ment of Entomology, Smithsonian Institu- 
tion. His talk was entitled "Evolution in 
Orb-Weaving Spiders." Refinements in web 
construction and prey capture strategies were 
discussed and illustrated. 

Visitors were introduced and the meeting 
was adjourned at 9:30 p.m., after which re- 
freshments were served. 

Richard G. Robbins, Corresponding Sec- 

927th Regular Meeting- December 4, 1 986 

The 927th Regular Meeting of The En- 
tomological Society of Washington was 
called to order by President E. M. Barrows 
in the Naturalist Center, National Museum 
of Natural History, at 8 p.m. on December 
4, 1 986. Twenty-two members and six guests 
were present. Minutes of the previous two 
meetings were read. Membership Chair- 
man, G. White, read the names of the fol- 
lowing applicants for membership: G. T. 
Baker, Mississippi State, Mississippi; J. 
Johnson, Moscow, Idaho; J. Rosenheim, 
Berkeley, California; M. Sharkey, Ottawa, 
Canada; D. Wahl, Gainesville, Florida; D. 

Wood, Ottawa, Canada; S. Larcher, Wash- 
ington, D.C. 

Annual reports of officers were given by 
the Treasurer, Editor, Membership Chair- 
man, Corresponding Secretary, and Custo- 
dian. Program Chairman, M. Raupp, an- 
nounced that the next regular meeting would 
be held on the second Thursday of next 
month, January 8, 1987. 

Nominating Committee Chairman, D. 
Anderson, presented the slate of nominees 
for 1987 officers. Wayne N. Mathis was 
nominated as Associate Editor. President 
Barrows called for further nominations of 
which there were none. A motion was made 
and seconded that the slate be accepted as 
presented. The motion was unanimously 

President Barrows discussed the Execu- 
tive Committee recommendation that life 
membership dues be raised from $150 to 
$200. After some discussion of procedure, 
a motion to amend the bylaws to raise life 
membership dues was made, seconded, and 

President Barrows also discussed the Ex- 
ecutive Committee recommendation to in- 
vite the Maryland Entomological Society to 
cosponsor the annual June banquet. It was 
decided to table any action until the Wash- 
ington Pest Science Society was contacted. 

E. Bickley presented two notes in the area 
of medical entomology. The first concerned 
Aedes albopictus and its movement north 
and east; it is now established in 12 states. 
The second concerned Lyme disease of 
which there were 74 human cases in Cali- 
fornia during 1985 and more expected in 
1986. Ixodes pacificus is thought to be the 
most important vector. 

R. Robbins displayed a brochure an- 
nouncing a new book on insect dormancy 
from the Biological Survey of Canada. 

D. Nickle exhibited several live and un- 
usual katydids collected during his recent 
Earthwatch trip to Peru. 

The speaker of the evening was Douglas 
W. Tallamy, University of Delaware, who 



presented a talk entitled "The Ecology of 
Maternal Behavior in Insects." 

Following the introduction of visitors, 
President Barrows thanked the Society of- 
ficers individually for their help to him and 
their services to the Society. He thanked 
also the Hospitality Committee composed 
of Margaret Collins, Mignon Davis, and 
David Nickle for preparing the fine refresh- 

ments that are served following each meet- 
ing. The gavel is usually passed to the new 
President at this time, but T. Wallenmaier 
did not attend the meeting because of a con- 
flict with his honeymoon. The meeting was 
adjourned at 9:50 p.m. 

Paul M. Marsh, Recording Secretary 

Reports of Officers 

Treasurer's Report 



Assets: November 1, 1985 
Total Receipts for 1986 
Total Disbursements for 1986 
Assets: October 31, 1986 










Net Changes in Funds $ 5,684.05 

Norman E. Woodley, Treasurer 



Corresponding Secretary's Summary of 

Major Activities for Calendar 

Year 1986 

Letters of welcome were sent to 38 new 
members. Ten members who had earlier 
been dropped from our rolls for nonpay- 
ment of dues were readmitted. The mem- 
bership list (now computerized) tonight 
stands at 592. Two of our members were 
welcomed to Emeritus status: Clyde F. Smith 
of Raleigh, North Carolina and John V. 
Thompson of Linwood, New Jersey. Eight 
(after this evening, 9) letters were written 
thanking our guest speakers.' Some two doz- 
en thank-you letters were written to mem- 
bers who contributed to our Special Publi- 
cation Fund when paying their dues. Fifteen 
letters (about the same number as in pre- 
vious years) were written to members who 
persist in paying the institutional subscrip- 
tion rate ($35/U.S., $40/foreign) rather than 

regular member's dues. The postage costs 
of this Office amounted to $25.00. 

Richard G. Robbins, Corresponding Sec- 

Editor's Report 

Beginning with the January, 1987 issue, 
the Proceedings will begin a two-column 
format. The Publications and Executive 
Committees agree that this measure will save 
some expense without sacrificing utility or 
beauty. Arly Allen of Allen Press, our print- 
er, suggested the change to us as a cost- 
effective measure and to reduce the size of 
future volumes. The main space-saving fea- 
ture comes from reducing the width of some 
tables and plates to one column instead of 
spreading them across the width of a whole 
one-column page. 



Next year the Society will publish a mem- 
oir, an identification manual to the North 
American genera of Braconidae. It will be 
about 100 pages long, half of them filled 
with drawings and SEM photographs. We 
think it will be a good seller. The memoir 
will be spiral bound. This will allow the 
book to lie perfectly flat while in use, a good 
feature when both hands are occupied turn- 
ing a specimen or focussing a dissecting mi-