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VOL. 105 


JULY 2003 

NO. 3 

(ISSN 0013-8797) 


of the 





ARAGON, AGUSTIN and MIGUEL-ANGEL MORON— Two new species of Phyllophaga Harris 

(Coleoptera: Scarabaeidae: Melolonthinae) from south central Mexico 559 

Diversity, abundance, and seasonality of adult and larval Symphyta (Hynienoptera) in the 
George Washington National Forest, Virginia, and the Monongahela National Forest, West 
Virginia 756 

BROWN. JOITN W. — Three new genera, two new species, and some rectifications in Neotropical 

Euliini (Lepidoptera: Tortricidae) 630 

BURKS, ROGER A. and RICHARD A. REDAK— The identity and reinstatement oi Homalodisca 

litumla Ball and Phera lacerta Fowler (Hemiptera: Cicadellidae) 674 

CARTER, MAUREEN E. and E. RICHARD HOEBEKE— Biology and seasonal history of 
Elasmostethiis atricornis (Van Duzee) (Hemiptera: Acanthosomatidae), with descriptions of 
the immature stages and notes on Pendergrast organs 525 

DZHANOK.MEN, K. A. and E. E. GRISSELL — Nomenclatural changes in Pteromalidae, with a 
description of the first New World species of Ormocerus Walker (Hymenoptera: 
Chalcidoidea) 535 

GORDON, ROBERT D. and SCOTT McCLEVE— Five new species of Euparixia Brown 

(Coleoptera: Aphodiidae: Eupariinae), with a revised key to species 685 

GREHAN, JOHN R. and JOHN E. RAWLINS— Larval description of a New Worid ghost moth, 
Phassus sp., and the evolutionary biogeography of wood-boring Hepialidae (Lepidoptera: 
E.xoporia: Hepialoidea) 733 

P. WRAIGHT — First report o{ Coenosia attenuata Stein (Diptera: Muscidae), an Old World 
"hunter fly" in North America 769 

MANLEY, DONALD G. — Dasymutilla Jalisco, a new species of velvet ant (Hymenoptera: 

Mutillidae), plus new synonymy for Dasymutilla canina ( Smith) 679 

MARSH, PAUL M. and SCOTT R. SHAW— Revision of North American Aleiodes Wesmael (Part 
7): The compressor Herrich-Schaeffer, ufei (Walley), gressitti (Muesebeck), and procerus 
Wesmael species-groups (Hymenoptera: Braconidae: Rogadinae) 698 

(Continued on back cover) 




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Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology. 
Smithsonian Institution. 10th and Constitution NW, Washington, D.C. 20560-0168. 

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105(3), 2003. pp. 525-5.34 





Maureen E. Carti-;r and E. Richard Hoebeke 

Department of Entomology, Comstock Hall. Cornell University. Ithaca. NY 14853, 
U.S.A. (e-mail: mec8@ cornel I. edii; 

Abstract. — Details of the seasonal history and habits, and duration of the immature 
stages of Elasmostethus athcornis (Van Duzee). a specialist acanthosomatid stink bug on 
spikenard (Arcilia raceniosa L.; Araliaceae) are presented and discussed. The bug's sea- 
sonal cycle is univoltine throughout its North American range, with peak adult emergence 
nearly synchronous with the first appearance of host plant inflorescences in early July. 
The egg and five nymphal instars are briefly described, with emphasis on color patterns, 
and all life stages are photographed. A brief review and illustrations of Pendergrast organs, 
imit|ue to the Acanthosomatidae. are also provided. 

Key Wonls: Hemiptera, Acanthosomatidae, Elasniostethus alrlconiis. seasonal history, 
immature stages, Pendererast organs 

The Acanthosomatidae, a group often 
considered by earlier workers as a subfam- 
ily or a tribe of an inclusive Pentatomidae, 
comprise three subfamilies, approximately 
47 genera, and 180 species worldwide (Ku- 
mar 1974. Schuh and Slater 1995). All are 
phytophagous, feeding on a variety of 
plants characteristic of mid- to late-succes- 
sional stages, with several genera feeding 
upon members of one of the oldest angio- 
sperm subclasses. Hamamelidae (orders 
Utricales. Fagales. and Hamamelidales) 
(Schaefer and Ahmad 1987). Six species, 
three each in the genera Elasmostethus Fie- 
ber and Elasimicha Stal. occur in North 
America (Thomas 1991). 

Elasmostethus atricornis (Van Duzee) 
is known from Montana to Quebec and 
south to South Carolina (Jones and Mc- 
Pherson 1980. McPherson 1982). Its pri- 
mary host plant. Aralia racemosa L. (Ar- 
aliaceae), is commonly referred to as 

American spikenard, which describes the 
elongated panicles of fragrant, greenish- 
white flowers. Leaves are divided and 
subdivided into 6 to 21 heart-shaped leaf- 
lets. It is a perennial shrub often found on 
ravine banks (Wiegand and Eames 1926), 
growing 1-2 m each summer and dying 
back completely in the fall. Maturing 
fruits (berries) become a deep crimson red 
to purple in the fall. American spikenard 
has a geographic distribution somewhat 
sympatric with that of E. atricornis. ex- 
cept the former is recorded farther south 
to Georgia and New Mexico. 

The objective of this study is to docu- 
ment the biology and seasonal history of 
this little-studied species through field ob- 
servations over multiple seasons: to photo- 
graph its unique habitat, host plant, and var- 
ious life stages: and to describe and illus- 
trate its immature stages. 



Table I. Seasonal history of Ekismostelhus alricomis on American spikenard, Aralia racemosa. in New 
York. (Note: Stippled bars indicate period when adults are overwintering in duff and leaf litter, and black bars 
indicate period when life stages arc on host plants.) 








Overwintering Adult$ 

Nevi Generator) 



2"' Instar 


4"' Instar 

S'" Instar 

Methods and Materials 

The seasonal history of E. atricornis was 
studied at several sites in Ithaca (Tompkins 
Co.), New York. Observations were made 
over four seasons, at irregular intervals (but 
usually every few days) from early to mid 
July through October, in 1987 and again 
from 1999 to 2001. The primary study sites 
included natural areas near Beebe Lake on 
the Cornell University campus and also 
along Fall Creek adjacent to the Cornell 
University Arboretum, and at two local 
state parks (Buttermilk Falls and Robert H. 

Developmental times were based on lab- 
oratory rearing in 2001, maintained at room 
temperature (ca. 20-22° C) under natural 

photoperiod (July-September). Nymphs 
were placed with spikenard inflorescences 
and fruits (in aquapics) in plastic boxes. 
Fruits were changed every 1-2 days, but 
developmental stages and molts were re- 
corded daily. 


Seasonal history and habits. — The gen- 
eralized field history (Table 1 ) of E. atri- 
cornis is based on populations found on 
American spikenard in the Ithaca area. New 
host plant shoots appear by mid May, and 
by mid July the plant produces large flow- 
ering stalks (Fig. 2). Overwintered adults 
(Fig. 1) began to appear on foliage of 
American spikenard (Fig. 3) after the plants 


Figs. 1-6. The acanthosomatid stink bug Elasmosielhus athcornis and its host plant. American spikenard. 
Alalia racemosa. 1. Adult feeding on host fruits. 2. Elongated flower panicle of host plant in early July. -^. 
Typical ravine bank habitat of host plant. 4. Cluster of fifth instars and two teneral adults on maturing fruits of 
host plant. ?. Aggregation of fifth instars on upper leaf surface of host plant, prior to molting lo new 
generation adults. 6. Adults clustered on host leaves, prior lo overwintering. 


produced inflorescences and small, devel- 
oping fruits. During all years studied, bugs 
generally were not observed until the first 
or second week of July. In 1987, adults 
were first noticed on July 10. In 1999, over- 
wintered adults were seen on July 14, with 
matings taking place on July 16. Adults 
were not seen on plants until July 20 in 
2001. Soon after they emerged from over- 
wintering sites, adults began to feed, prob- 
ing flowers and developing reproductive 
structures: they also sought mates. Soon af- 
ter mating, egg deposition began. When 
oviposition began before fruits were avail- 
able (i.e., late July to early August), eggs 
were deposited on the lower leaf surfaces, 
usually arranged (Fig. 7) in small tight clus- 
ters of 12-25 eggs, or often as individual 
eggs. After early to mid August, the major- 
ity of eggs were laid on the surface of in- 
dividual fruits or on the pedicels. Eggs were 
observed as early as July 22 in 1999 and 
July 30 in 2000, and females continued to 
lay eggs as late as September 15 in 2001. 
Eggs hatched approximately 2-3 days after 
deposition. First instars, which hatched syn- 
chronously in a clutch, were found in the 
field as early as July 25 in 2001. First in- 
stars do not stray from the egg shells (Fig. 
8), but remain motionless in place: they ap- 
parently did not feed on their host plant 
during this life stage. After molting to the 
second instar (Fig. 9), nymphs dispersed to 
host fruits. Second instars were observed by 
early August (ca. August 2) in 1999, closely 
followed by third instars (Fig. 10) and 
fourth instars (Fig. 1 1 ) (August 14 in 2000; 
August 11 in 2001), and then fifth instars 
(Fig. 12) (August 15 in 2001). Second 
through fifth instars fed on fruits after prob- 
ing with their stylets. Fourth and fifth in- 
stars, highly polymorphic in coloration 
(Figs. 4-5, 11-12), often clustered among 
the maturing berries (Fig. 4), where they 
were essentially camouflaged by the very 
similar fruit shape and coloration (Aldrich 
1988); this crypsis might protect the 
nymphs from predation. Fifth instars often 
aggregated on upper leaf surfaces, in fold- 

ed-over leaves, and in large numbers just 
prior to adult eclosion (Fig. 5). Adults also 
clustered on leaves in the fall (Fig. 6). 
Adults of the new generation began to ap- 
pear by late August (August 23 in 1987 and 
2001) and continued to appear until early 
to mid October (October 7 in 2001 and Oc- 
tober 1 1 in 1999). In 1999, new adults were 
present as late as October 19. By late Oc- 
tober, or after the first hard frosts, most 
adults disappeared from host plants, pre- 
sumably having moved to overwintering 
sites in and under the deep leaf litter below 
and around the senescing hosts, as observed 
for E. interstinctus (L.) in Alaska (Barnes 
et al. 1996). Only a single generation was 
produced annually in central New York, as 
is the case elsewhere in its range in North 
America (Jones and McPherson 1980). 

Duration of immature stages. — In the 
laboratory at 20-22° C (n = 6 egg clusters 
observed), the incubation time for eggs of 
E. atricornis ranged from 4—6 days after 
clutch deposition; by contrast, in the field 
the majority of eggs hatched 2-3 days after 
deposition. Nymphal development (from 
egg hatch to adult eclosion) ranged from 1 1 
to 39 days. The earliest appearance of new 
adults (from egg hatch) averaged 18.8 days 
(range 1 1-27) and the latest averaged 32.4 
days (range 28-39). Each of the first two 
nymphal stadia generally lasted 2-3 days, 
while the third and fourth nymphal stadia 
required 2-4 days (10-11 days for some 
individuals). The fifth nymphal stadium av- 
eraged 9 days (range 2-14). As in a natural 
setting, nymphs of all instars remained on 
the clusters of maturing host fruits through- 
out the period of nymphal development. 

Description of Immature Stages 

Egg (Fig. 7). — Length, 0.88-0.93 mm (n 
= 4); width, 0.64-0.73 mm (n = 7). Ovoid, 
slightly pointed apically, pale translucent 
green. Red eye spots visible through egg 
chorion just prior to egg hatch. 

First instar (Fig. 8).— Length, 0.97-1.18 
mm; width, 0.52-0.98 mm (n = 12). Head, 
thorax, and dorsal plates (= abdominal 


? ^- «. 


l-igs. 7-12, L.iK- Ma;jL-~ of EUisiii,istclhiis atrmnnis^ 1. Cluster of eggs on Iowlm IlmI sinUice ot ,4ri//)<( 
iiui'inosa. 8. Newly lialclied first instars. 9. Second inslars. II). Third instar. II. Fourth Instar. 12. Fifth mstar 

scent gland openings) uniformly brownish, 
except ecdysial lines of head and median 
line of thorax pale, subhyaline. Eye dark 
red. Antenna pale brown. Legs pale yellow: 
tarsi darker. Abdomen uniformly yellowish 

Anterior wing pads undeveloped. Ap- 
proximate ratio of lengths along median 
line of pro-: meso-: metanotum = 4.0: 3.0: 

1 . Antennomere ratio (approximate) of I: II: 
III: IV = 1: 1.2: 1.2: 2.2. 

Second instar (Fig. 9). — Length, 1.67- 
2.54 mm.; width, 1.26-1.67 mm (n = 12). 
Head, thorax, and dorsal plates mostly dark 
brown to black. Anterior dorsal plate with 
some reddish highlights. Abdomen pale 
yellowish. Eye dark red to black. Antenna 
pale yellow, subhyaline, except basal anten- 


nomere infuscated and apical antennomere 
darkened (reddish black) on apical half or 
more. Legs pale yellowish, except femora 
dark brown. Apical tarsal segment (bearing 
claws) slightly darkened. 

Anterior wing pads undeveloped. Ap- 
proximate ratio of lengths along median 
line of pro-: meso-: metanotum = 7.1: 4.9: 
1. Antennomere ratio (approximate) of I: II: 
III: IV = 1: 1.2: 1.3; 1.6. 

Second instar very similar to first instar. 
except head and thorax of latter darker; 
sternal plates of thorax also darkly pig- 
mented as well as femora and tip of ros- 
trum. Distal half or more of apical anten- 
nomere piceus or black. 

Third instar (Fig. 10). — Length. 2.39- 
4.75 mm; width, 1.32-3.61 mm (n =11). 
Head, thorax, and dorsal plates dark brown, 
except anterior portion of head (including 
juga and tylus), lateral margins and middle 
of pronotum and mesonotum, and central 
portions of dorsal plates (only narrow an- 
terior and lateral margins dark) pale yellow- 
ish. Abdomen mostly yellowish green. Nar- 
row darkened anterior margins of dorsal 
plates with some reddish highlights. Eye 
dark red or black. Antenna infuscated, basal 
antennomere mostly dark brown, anten- 
nomeres 2 and 3 centrally darkened, apical 
(4th) antennomere dark reddish brown. 
Legs blackish brown except bases of fem- 
ora and middle of tibiae pale yellowish. 

Anterior wing pads undeveloped. Ap- 
proximate ratio of lengths along median 
line of pro-: meso-: metanotum = 4.5: 4.0: 
1 . Antennomere ratio (approximate) of I: II: 
II: IV = 1: 1.5: 1.7: 2.2. 

Fourth Instar (Fig. 11). — Length, 3.45- 
5.16 mm; width, 2.73-4.01 mm (n = 14). 
Coloration of head and thorax highly vari- 
able, ranging from pale green to blackish. 
Dorsal plates also variable in color, from 
pale yellowish to dark reddish brown with 
reddish and black highlights. Abdominal 
segments pale yellowish green. Eye gener- 
ally reddish. Antennomere reddish brown to 
dark brown to blackish. Legs pale green to 

reddish brown; apical tarsal segment gen- 
erally darker. 

Anterior wing pads developed, attaining 
posterior margin of metanotum; posterior 
wing pads undeveloped. Approximate ratio 
of lengths along median line of pro-: meso-: 
metanotum = 4.0: 7.1: 1. Antennomere ra- 
tio (approximate) of I: II: III: IV = 1: 2.1: 
1.8: 2.3. 

Fifth Instar (Figs. 4-5, 12).— Length, 
6.61-8.55 mm; width. 4.54-5.49 mm (n = 
14). As in preceding instar, coloration of 
head, thorax, and dorsal plates highly var- 
iable. In non-maculated specimens, head, 
thorax, and abdomen pale yellowish green, 
except antennomeres reddish to reddish 
brown, tarsal segments reddish brown, and 
dorsal plates highlighted with some red and 
black. In maculated specimens, head and 
thorax (including well-developed wing 
pads) variously marked with black. Dorsal 
plates mostly dark reddish or black around 
perimeters, greenish yellow centrally. Ab- 
domen generally pale green. Ocelli and eye 
dark reddish. Antennomeres mostly dark 
reddish brown. Femora mostly pale green- 
ish; tibiae generally brownish; apical por- 
tion of terminal tarsal segment dark red to 

Scutellum well developed, attaining pos- 
terior margin of metanotum. Anterior wing 
pads highly developed, extending to third 
or fourth abdominal tergite. Posterior wing 
pads conspicuously developed. Approxi- 
mate ratio of lengths along median line of 
pro-: meso-: metanotum = 13.7: 20.7: 1. 
Antennomere ratio (approximate) of 1: II: 
III: IV = 1: 1.8: 1.9; 1.7. 

Note; Beginning with the fourth instar, 
which is nearly synchronous with fruit rip- 
ening (fruits become dark red to puiple), 
and continuing with the fifth instars, the 
dorsal coloration is generally highly vari- 
able, with pale morphs of mostly yellowish 
green with no dark markings to maculated 
morphs with red and black highlights on the 
head and thorax, including dark red, brown- 
ish or black wing pads. Late in the season 
(September into October), this variable dor- 



c v: 


-i. ^* 

Figs. 1,3-17. Scanning electron photomicrographs of abdominal venters of acanlhosomatid adults. \}>. Ab- 
dominal \enter and Pendergrast organs (arrows) of Elasmosrerlnis criiciatiis. 14. Abdominal venter of f/i/.v/zf^t/k; 
lateralis. 15. Close-up of Pendergrast organs on sterna 6 & 7 of E. cnicianis. 16-17. Magnification of setae 
( 16) and cuticular pores ( 17) of the Pendergrast organs of E. cnicialus. 

sal coloration makes nymphs difficult to see 
among the ripening fruits on which they 
feed (see Fig. 4). 

Pendergrast Organs 

Pendergrast organs, structures on the ab- 
dominal venter of females only, are unique 
to the Acanthosomatidae. They have been 

used as one of the characters defining 26 
genera (Kumar 1974). although a similar 
structure exists in at least one other hemip- 
teran family, the Lestoniidae (Fischer 
2000). In observing the oviposition behav- 
ior of E. atricornis. we found that the fe- 
male, after depositing an egg. brushes each 
hind tarsus alternately and repeatedly 


against the Pendergrast organs, followed by 
an apparent tapping of the egg. This se- 
quence of behavior was repeated 5-10 
times continuously for 30-60 seconds. Pen- 
dergrast (1952) noted similar behavior for 
the New Zealand species Rhopaliniorpha li- 
neolaris Pendergrast. 

In both E. atricornis and E. cruciatns 
(Say), the Pendergrast organs are circular, 
depressed areas located laterally on the 
sixth and seventh abdominal sterna, a pair 
of depressions on either side of the midline 
(Fig. 13). Under high magnification, these 
depressed cuticular areas are clothed with 
short setae (Figs. 15-16) interspersed with 
minute pores (Fig. 17). Microscopic exam- 
ination (Breddin 1903, Pendergrast 1953. 
Carayon 1981, Staddon 1990) and SEM 
and transmission electron microscopy (Fi- 
scher 1994) of cuticular preparations of oth- 
er acanthosomatid species indicate that the 
pores are connected by ducts to glandular 

Several hypotheses have been promoted 
that speculate on the function of the Pen- 
dergrast organs, including their involve- 
ment in copulation (Breddin 1903. Pender- 
grast 1953), in pheromone production 
(Staddon 1990), and in the secretion of 
compounds that deter egg predation (Al- 
drich 1988, Fischer 1994). Acanthosoma- 
tids in genera that lack Pendergrast organs 
instead brood or guard their eggs and 
young. For example, females of Elasmiicha 
lateralis (Say), a common North American 
acanthosomatid in which Pendergrast or- 
gans (Fig. 14) are absent, exhibits brooding 
behavior. Females of E. lateralis, and those 
of other species in this worldwide genus, 
guard their eggs, first instars and sometimes 
later instars (Frost and Haber 1944; Kudo 
1990, 2000; Kudo and Nakahira 1993; Kai- 
tala and Mappes 1997). 


The absence of apparent feeding in first 
instars and the late-season appearance of 
overwintered adults are not unique to Elas- 

mostethiis. These biological attributes are 
shared by other heteropteran taxa. 

Lack of feeding by first instars is not un- 
usual among pentatomomorphan Heterop- 
tera. In many instances, first-instar nymphs 
of seed-sucking species, such as the south- 
ern green stink bug (Nezara viridiila (L.)), 
the rice stink bug (Oebaliis pugna.x (F.)) 
(Bowling 1979, 1980), and the green stink 
bug (Acrostenmm liilare (Say)) (Simmons 
and Yeargan 1988). remain clustered on the 
eggs shells and do not feed on plant tissue; 
however, occasionally the nymphs have 
been observed probing the egg shells with 
their stylets. First instars of many predatory 
pentatomids (subfamily Asopinae) do not 
feed, or feed only on unhatched eggs of 
their own species (Wheeler 2001), such as 
Eiithyrhynchus floridaniis (L.) (Getting and 
Yonke 1975), Podisits niaciiliventris (Say) 
and P. placidiis Uhler (Mukerji and Le- 
Roux 1965, Getting and Yonke 1971), and 
Stiretnis fiinbriatits (Say) and S. anchorago 
(F) (Getting and Yonke 1971. Waddill and 
Shepard 1974). Some first-instar coreoids 
also do not feed (Pupedis et al. 1985). For 
example, first instars of Leptoglossus ful- 
vicornis (Westwood), a magnolia fruit-feed- 
ing specialist, do not feed or do so only 
minimally on foliage (Wheeler and Miller 
1990). while first instars of L. occidentalis 
Heidemann, the western conifer-seed bug, 
often are able to reach the second stadium 
without feeding (Koerber 1963). 

Several heteropterans also are character- 
ized by the late-season appearance of over- 
wintered adults. Wheeler and Miller (1990) 
documented that the coreid Leptoglossus 
fiilviconus overwinters in the adult stage in 
Pennsylvania and remains in hibernation 
sites until mid-June to early July at which 
time adults then colonize host trees when 
small, cone-like fruits begin to appear. Oth- 
er specialist Heteroptera that overwinter as 
adults and do not become active until late 
in the season when inflorescences or devel- 
oping fruits of their host plants are available 
include the lygaeid Ochriiwnis mimithis 
(Stal) (Palmer 1986) and the pentatomid 



Proxys punctulatus (Palisot de Beauvois) 
(Vangeison and McPherson 1975). 


We express our gratitude to Kent Loeffler 
(Plant Pathology, Cornell University) and 
Carole Daugherty (Plant Biology, Cornell 
University) for their assistance with macro- 
and scanning electron photomicroscopy, re- 
spectively, and to A. G. Wheeler, Jr. (Clem- 
son University), T. J. Henry (USDA-ARS. 
Systematic Entomology Laboratory, Wash- 
ington, D.C.), and D. A. Rider (North Da- 
kota State University, Fargo) for their crit- 
ical reviews of a draft of this paper. 

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Barnes, M. B., J. L. Barger, J. Seare.s, P. C. Tacquard, 
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Bowling. C. C. 1979. The stylet sheath as an indicator 
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. 1980. The stylet sheath as an indicator of 

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Carayon. J. 1981. Dimorphisme sexuel des glandes 
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zungsberichte der Gesellschaft Naturforschender 
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. 2000. The disc-like organ of the Lestoniidae 

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Frost. S. W. and V. R. Haber. 1944. A case of parental 
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105(3). 3()()3. pp. ?35-541 


DE:SCRIPTI0N of the first new world SPECIES OF ORMOCERUS 



(KAD) Institute of Zoology, Akademgorokok. Almaty, 480060. Republic of Kazakh- 
stan; (EEC) Systematic Entomology Laboratory, PSI, Agricultural Research Service. U.S. 
Department of Agriculture, c/o National Museum of Natural History. Washington. D.C. 
20560-0168. U.S.A. (e-mail: e2rissell@sel.barc.usda.20v) 

Abstract. — The new species Ormoceriis americaniis Dzhanokmen and Grissell is de- 
scribed based on specimens reared from cynipid galls on oak in Texas. The following 
new synonymies are proposed: Ahyrsoinele Dzhanokmen 1975a = Halticopterimi Erdos 
1946: Halticopterina peiithoconiie Dzhanokmen 1975a = H. luoczari Erdos 1954; Hom- 
oporiis longiventris Dzhanokmen 1999 = H. cupreus Erdos 1953; Phaenocytiis hepta- 
potamiciis Dzhanokmen 1990 = P. glechomue (Fcirster 1841); Pseudocatolacciis aine- 
golliis Dzhanokmen 1989 = P. iiitescens (Walker 1834); Pteromalus inaculatits Dzhan- 
okmen 1998 = P. vopisciis Walker 1839; Stenoselma luiplogcistra Dzhanokmen 1975a = 
S. nig mm Delucchi 1956; Stenoselma armeniaca Dzhanokmen and Herthevtzian 1990 = 
S. nigrum Delucchi 1956; Stirogeniiim Dzhanokmen 1985 = Paracarotomus Ashmead 
1894; Stirogeniiim asiaticiim Dzhanokmen 1985 = Paracarotomus cephalotes Ashmead 
1894. The following are new combinations: Chlorocytus arkansensis (Girault) from Ha- 
lirocytus Thomson; C. languriae (Ashmead) from Habrocytus Thomson; C. rhodobaeni 
(Ashmead) from Habrocytus: C. simillinuis (Gahan) from Habrocytus: C. vassiliefi (Ash- 
mead) from Homoporus Thomson; Halticopterina lauta (Dzhanokmen) from Abyrsomele 
Dzhanokmen; Mesopolobus elymi (Dzhanokmen) from Platneptis Boucek: Neocatolaccus 
carinatus (Howard) from Catolaccus Thomson. 

Key Words: Chalcidoidea. Pteromalidae. cynipid galls, oak 

The family Pteromalidae is worldwide in amination of the types and collection ma- 
distribution and is numerically the third terial from the Canadian National Collec- 
largest family within the superfamily Chal- tion. Ottawa (CNC); The Natural History 
cidoidea (Noyes 2001). Currently there is Museum, London (NHM. London); Nation- 
virtually no worldwide consensus of opin- al Museum of Natural History. Smithsonian 
ion on the ranks of taxa composing the fam- Institution. Washington D.C. (USNM); Nat- 
ily or of the definition of subfamilies that ura! History Museum, Budapest (NHM, 
should be recognized. The status of a great Budapest); Zoological Institute, St. Peters- 
many higher level taxa have not been eval- burg (ZI); and Institute of Zoology, Almaty, 
uated and most of the genera and species Kazakhstan. We also describe a new species 
remain unstudied. We take this opportunity of Ormocerus Walker, reared from cynipid 
to make some changes in the nomenclature galls on oak. which is the first reported spe- 
of Holarctic pteromalids based upon ex- cies of the genus in the New World. 


New Synonymies 
Hahicopterimi Erdos 
Hcilticopteriiui Erdos 1946; 160. 

Type species: Halticopterina trinumilatci 
Erdos. Original designation. (NHM, Buda- 
pest, examined.) 

Abyrsomele Dzhanokmen 1975a: 627. New 

Type species: Abyrsomele lauta Dzhan- 
okmen. Original designation. (ZI, exam- 

The type species of Aljyrsomele does not 
differ from Halticopterina in generic char- 
acters and is herein considered a subjective 
junior synonym of the latter genus. 

Halticopterina moczari Erdos 

Halticopterina moczari Erdos 1954: 153- 
154. (NHM. Budapest, examined.) 

Halticopterina penthocoryne Dzhanokmen 
1975a: 625-627. New synonymy. (ZI. 

Examination of the type specimens 
leaves no doubt that H. penthocoryne is a 
subjective junior synonym of H. moczari. 

Honioporus cupreus Erdos 

Honioporus cupreus Erdos 1953: 245. 
(NHM. Budapest, examined.) 

Honioporus longiventris Dzhanokmen 
1999: 183-185. New synonymy. (ZI, ex- 

Honioporus longiventris was distin- 
guished from H. cupreus chiefly by having 
lighter colored antennae and legs and by its 
narrower gaster. These differences, based 
upon an examination of several series of 
specimens, now appear to occur within the 
range of variation of a single species and 
we consider H. longiventris a subjective ju- 
nior synonym of H. cupreus. 

Phaenocytus glechomae (Forster) 

Pteromalus glechomae Forster 1 84 1 : 21. 
Phaenocytus glechomae: Graham 1969: 

Phaenocytus heptapotamicus Dzhanokmen 
1990: 65-67. New synonymy. (ZI. ex- 

The genus Phaenocytus Graham was cre- 
ated for Pteromalus glechomae Forster. Al- 
though we have not seen the type, we com- 
pared P. heptapotamicus with specimens of 
P. glechomae housed in the NHM, London. 
The two are identical and we consider the 
former a subjective junior synonym of P. 

Pseudocatolaccus nitesceiis (Walker) 

Amhlymerus nitescens Walker 1834: 347. 
(NHM, London, examined.) 

Pseudocatolaccus nitescens: Graham 1969: 

Pseudocatolaccus amegallus Dzhanokmen 
1989: 45-46. New synonymy. (ZI. ex- 

Examination of the type and additional 
series of Pseudocatolaccus nitescens 
(Walker) convinces us that it is the same as 
P. amegallus. Pseudocatolaccus nitescens 
is a rather variable species (Graham 1969), 
and it seems that P. amegallus is just a form 
of P. nitescens having the postmarginal 
vein shorter than normal. Thus we consider 
P. amegallus a subjective junior synonym 
of P. nitescens. 

Pteromcdus vopiscus Walker 

Pteromalus vopiscus Walker 1839: 274. 
(NHM, London, examined.) 

Pteromalus maculatus Dzhanokmen 1998: 
494-496. New synonymy. (ZI, exam- 

Examination of the type of Pteromalus 
vopiscus Walker showed that P. maculatus 
is almost certainly the same species. It dif- 
fers from the latter in small details that we 
consider to fall within the range of individ- 
ual variability of P. vopiscus. For this rea- 
son we place P. maculatus as a subjective 
junior synonym of P. vopiscus. 

Paracarotomus Ashmead 
Paracarotoimis Ashxnead 1894: 335-336. 


Type species: Paracarotomus cepludoles 
Ashmead. Monobasic. (USNM. examined.) 

Stiroi>eiuiiin Dzhanokmen 1985: 152. New 

Type species: Stirogenium asiaticiim 
Dzhanokmen. Original designation. (ZI, ex- 

Based upon an examination of the type 
species of Paracarotomus and Stirogeiuuin. 
the latter differs in no moiphological way 
and so is now considered a subjective junior 
synonym of Paracarotomus. 

Paracarotomus ceplialotes Ashmead 

Paracarotomus ceplialotes Ashmead 1894: 
335. (USNM. examined.) 

Stirogenium asiaticitm Dzhanokmen 1985: 
152-153. New synonymy. (ZI, exam- 

After comparing the type specimens of 
Stirogenium asiaticum and Paracarotomus 
ceplialotes we found that they differ in no 
way. Therefore we consider Stirogenium 
asiaticum a subjective junior synonym of 
Paracarotomus ceplialotes. 

Stenoselma nigrum Delucchi 

Stenoselma nigrum Delucchi 1956: 66-68. 

Stenoselma haplogastra Dzhanokmen 
1975b: 1096-1097. New synonymy. (ZI. 

Stenoselma armeniaca Dzhanokmen and 
Herthevtzian 1990: 139-142. New syn- 
onymy. (ZI, examined.) 

We examined all the determined speci- 
mens of S. nigrum in NHM, London. The 
differences between Stenoselma nigrum and 
S. haplogastra appear to fall within the 
range of variation of a single species, and 
we have no hesitation in placing S. haplo- 
gastra as a subjective junior synonym of S. 
nigrum. The differences given in the de- 
scription of armeniaca also fall within the 
range of variation of nigrum and we place 
■S'. armeniaca as a subjective junior syno- 
nym of .S'. nigrum. 

New Combinations 

As a result of the examination of the 
types of additional pteromalid species, we 
propose the following new combinations: 

Chloiocytus arkan.sensis (Girault 1917), n. 

comb, from Habrocytus Thtimson 
Chlorocytus Icmguriae (Ashmead 1896), n. 

comb, from Habrocytus Thomson 
Chlorocytus rhodobaeni (Ashmead 1896), 

n. comb, from Habrocytus Thomson 
Chlorocytus simillimus (Gahan 1919), n. 

comb, from Habrocytus Thomson 
Chlorocytus rassiliefi (Ashmead 1903), n. 

comb, from Homoporus Thomson 
Halticopterina lauta (Dzhanokmen 1975a), 

n. comb, from Abyrsomele Dzhanokmen 
Mesopolobus elymi (Dzhanokmen 1984). n. 

comb, from Platneptis Boucek 
Neocatolaccus carina tus (Howard 1897), n. 

comb, from Catolaccus Thomson 

The genus Chlorocytus. established by 
Graham (1956). was not formally reported 
in the Nearctic until 1997 (Boucek and 
Heydon 1997). No described species were 
included at that time, and the above new 
combinations are the first species reported 
for the Nearctic. These taxa were described 
much earlier when such important charac- 
teristics as the shape and structure of the 
prepectus and propodeum were not critical- 
ly analyzed. In Chlorocytus the prepectus is 
large and uniformly reticulate, the propo- 
deum is nearly always large (more than half 
as long as the scutellum) and with irregular 
carinae and shallow spiracular sulci. Addi- 
tionally, in Chlorocytus the antennal clava 
in profile is often asymmetrical, its upper 
edge strongly curved. These characters dis- 
tinguish Chlorocytus from the morphologi- 
cally similar genus Pteromalus Swederus 
(and its synonym Habrocytus, which some 
may regard as a subgenus). 

Ormocerus Walker 

Burks (1979) placed the genus Ormoce- 
rus Walker in the subfamily Miscogasteri- 
nae. tribe Ormocerini and included O. flora 


(Girault) and O. vulgaris (Ashmead) in the 
genus. Heydon and Boucek (1992) exam- 
ined the types of these two species and de- 
termined that neither belonged to Ornioce- 
riis. For O. flora they established the genus 
Ficicola Heydon and Boucek; they trans- 
ferred O. vulgaris to the genus Seladenna 
Walker. As a result of these actions, none 
of the described Nearctic species remained 
in the genus Onnoceriis. In their key to Ne- 
arctic genera of Pteromalidae, Boucek and 
Heydon (1997) cited Ormocerus as having 
"at least 4 spp." based apparently on their 
knowledge of undescribed species in the re- 
gion. Frankie et al. (1992; Table 17.6) list- 
ed, but did not discuss, '^ Ormocerus n. sp." 
as a parasitoid of the sexual generation of 
a cynipid gall on oak in Texas. It is this 
material, representing the first known spe- 
cies of the genus from the New World, that 
we describe herein. 

Ormocerus ainericaniis Dzhanoknien 
and Gri.ssell, new species 

(Figs. 1-4) 

Female holotype. — Body length 2.1 mm. 
Head, mesosoma, and coxae dark green 
with metallic gloss. Metasoma dark ful- 
vous. Antenna dark; scape dark with me- 
tallic tinge, pedicel dark brown, flagellum 
brown. Femora mainly brown to black with 
metallic green tinge, tibiae fuscous except 
at apices, remaining parts of legs testa- 
ceous, apices of tarsi fuscous. Forewing ap- 
pearing hyaline, but with slight fuscous 
cloud below stigmal vein extending below 
marginal to parastigmal vein; venation 
brownish testaceous. 

Head in dorsal view 2.3 times as broad 
as long. POL about 2.3 OOL. head trans- 
versely oval in frontal view. Gena relatively 
short (Fig. 4), very slightly curved, con- 
verging strongly towards mouth. Anterior 
margin of clypeus (Fig. 2) evenly curved 
forward. Eyes relatively large, height about 
2.5 X height of gena (Fig. 4). Antennae in- 
serted low on face, toruli at or slightly 
above level of ventral eye margin. Antennal 
flagellum clavate (Fig. 1); 2 transverse anel- 

li, anellus barely as wide as first funicle 
segment; first funicle segment shorter and 
narrower than second; second and third seg- 
ments slightly transverse; following seg- 
ments strongly transverse. Both mandibles 
with three teeth (Fig. 2). Maxillary palp 4- 
segmented. labial palp 3-segmented. 

Mesosoma about 1.4X as long as broad. 
Mesoscutum with reticulate sculpture, in- 
terspersed with numerous minute pits from 
which arise setae, which cover mesoscu- 
tum. Notaulus complete and deep, not in- 
terrupted by sculpture. Scutellum distinctly 
longer than broad, convex, sculptured like 
mesoscutum though with more conspicuous 
pits among reticulation on lateral sides and 
base. Frenum weakly delimited. Propodeum 
medially nearly !4 scutellum length, weakly 
alutaceous, almost smooth, with distinct 
median carina, without plicae. Callus with 
several hairs; spiracles subcircular, separat- 
ed by nearly % their greatest diameter from 
metanotum. Upper mesopleuron large, pol- 
ished and smooth. Prepectus finely reticu- 
late. Spur of midtibia longer than first tarsal 
segment. Forewing with speculum closed 
below, not extending farther than beginning 
of marginal vein; disc beyond speculum 
densely setose. Lower surface of costal cell 
with only a few setae in distal Vi. Basal cell 
with some setae proximally and distally 
along cubital vein; basal vein a conspicuous 
transparent tract, with only one or two setae 
at junction dorsally with parastigma. Apical 
margin of wing with fringe. Postmarginal 
vein slightly longer than marginal, latter 
I.2X longer than stigmal (Fig. 3). 

Metasoma about as long as mesosoma, 
I.8X as long as broad, acute apically, nar- 
rower than mesosoma. Basal tergite occu- 
pying Vi total length, its hind margin me- 
dially emarginate. Caster ventrally keeled, 
hypopygium extending nearly Vi distance 
from base of gaster to tips of ovipositor 
sheath, which project slightly at most. 

Male. — Body length 2.0 mm. Agreeing 
with female except as follows; forewing 
completely hyaline; stigmal vein elongately 
triangular (instead of linear), widening from 



Figs. 1-6. Ormocerus spp. 
margin of clypeus. 3, Forewing 
view. 6. Antenna. 

1 — 1. Ontiocerus americainis. female. 1, Antenna. 2. Mandibles and anterior 
venation. 4, Head, lateral view. 5-6, Onitoccnis lams, female. 3. Head, lateral 

marginal vein to junction with stigma 
where it is subequal in width; flagellum cy- 
lindrical, clothed with dense, semierect se- 
tae, funicular segments strongly transverse 
and nearly twice width of anellus. 

Variation. — Female specimens vary from 
1.8 to 2.1 mm in length. There is little var- 
iation in color except the general metallic 
green tinge of the femora may disappear, 
leaving them dark brown in color. The fore- 
wing infuscation is obscure at best; in some 
specimens it is little more than a slight 
darkening posterior to the stigma; in others 
it is barely visible but extends to the par- 
astigma. The eye height ranges from 2.5 to 
3 times the genal length. Only one male is 

Type material. — Holotype ?. Dallas. 
Dallas County, Texas, coll. 2-IV-1974, em. 
1-15-V-1974, coll. Gordon Frankie, ex bi- 
sexual generation Disholcaspis cinerosa 
(Bassett) [Hymenoptera: Cynipidae] on 
Qiierciis virginiana Mill [Fagaceae] 

(USNM); 15 9, 1 (5 paratypes same data 
as holotype; 3 $ paratypes same data ex- 
cept collected 29-III-1974 (no emergence 
date) (USNM; 1 9 paratype in ZI, 2 9 
paratypes in CNC). 

Host. — The species was reared from the 
bisexual generation of Disholcaspis cine- 
rosa on Quercus virginiana. 

Discussion. — Ormocerus americanus 
differs from the two European species. O. 
latiis Walker and O. venuilis Walker chiefly 
in its shorter gena and larger eye (Fig. 4). 
The eye is 2.5 to 3 times the genal length 
in O. americanus. whereas in O. latiis and 
O. vernalis it is about 2 times the genal 
length (Fig. 5). Both O. americanus and O. 
latus differ from O. vernalis in several fea- 
tures including: the presence of an apical 
wing fringe (absent from O. vernalis), the 
basal cell has many setae and the basal vein 
is relatively bare (basal cell bare but the 
basal vein with several rows of setae in O. 
vernalis), forewing with pale infuscation 


sometimes seen only with difficulty (darkly 
infuscated in O. venmlis). the stigmal vein 
is curved (straight in O. vernalis). and the 
propodeum is lightly reticulate (coarsely ru- 
gulose in O. vernalis). Ormocerus ameri- 
caniis differs from O. latus by the basal fla- 
gellomeres being wider than long (Fig. 1) 
(longer than wide in O. latus. Fig. 6); the 
notauli shaiply defined as nearly smooth 
grooves (ill-defined and interrupted by 
sculpture in O. latus): and the median area 
of the propodeum less reticulately sculp- 
tured (almost alutaceous) than the lateral ar- 
eas (nearly uniformly sculptured in O. la- 

The three known species of this genus 
attack cynipid gall-formers on oak. 


We acknowledge The Royal Society 
(London) and The Smithsonian Institution 
(Washington. DC) for financial support dur- 
ing much of this research. The help of John 
Noyes and Zdenek Boucek, The Natural 
History Museum, London, is greatly appre- 
ciated by the senior author. We thank Gary 
Gibson, Systematic Entomology Section. 
Ottawa, for donating the only known male 
specimen of Ormocerus americanus to our 
study. In addition we thank Michael Pogue, 
David Smith, Steve Heydon, and Gary Gib- 
son for reviewing the manuscript and mak- 
ing many helpful suggestions for its im- 

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Hungarica 7: 153-154. 


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333 pp. 


105(3), 2003, pp. 542-547 


Gary J. Steck, Bruce D. Sutton, and Don Defoe 

(GJS. BDS) Florida State Collection of Arthropods, Florida Department of Agriculture 
& Consumer Services, Division of Plant Industry, Gainesville FL 32614-7100, U.S.A. (e- 
mail: steckg@doacs. state. fl. us; suttonb@doacs. state. fl. us); (DD) Great Smoky Mountains 
National Park, Gatlinburg. TN 37738, U.S.A. (e-mail: 

Abstract. — Adults of Rhyiieiuina longirostris Johnson were successfully reared from 
larvae in the achenes of Sinallanthiis iivedaliiis (L.) Mackenzie (Asteraceae), the first 
confirmed host plant record for this fly. Adults of R. longirostris are univoltine with a 
flight-time corresponding to the flowering period of 5. uvedaliiis. which is July-September 
in the Great Smoky Mountains National Park. Heteroschema sp. (Hymenoptera: Pter- 
omalidae) parasitoids and Homoeosomo sp. (Lepidoptera: Pyralidae) feeding in capitula 
of the host are probably significant sources of mortality of/?, longirostris immature stages. 

Kex Words: 

Smallaiuhus uvedaliiis. Great Smoky Mountains National Park. All Taxa 
Biological Inventory, host plant 

In connection with the All Taxon Biolog- 
ical Inventory (ATBI) that was initiated in 
the Great Smoky Mountains National Park 
(GSMNP) in 1997. Gary Steck (GJS) and 
Bruce D. Sutton (BDS) joined a dipterists" 
"collecting blitz" in late May, 1999. During 
this visit, GSMNP collections curator, Don 
DeFoe (DD), kindly allowed us to peruse 
the collection in the Natural History Mu- 
seum (GSNP) at the park headquarters at 
Sugarlands. where we noted the presence of 
six specimens of a rarely collected tephritid 
fly, Rhynencina longirostris Johnson. These 
specimens are the first known record of R. 
longirostris in the state of Tennessee (Steck 
and Sutton 2000), with label data as fol- 
lows: '"Tennessee: Sevier Co., Park HQ, 
1520' elev., 5 Aug 1996.'" DD, as collector 
of the specimens, was able to take us di- 
rectly to the location and the plants upon 
which the adult flies had been taken; this 
was a small patch of Smallantlms uvedaliiis 
(L.) Mackenzie (Asteraceae) in a disturbed. 

shaded area next to a parking lot. The flies 
had been on the flower heads at the time of 
capture. That discovery prompted all of the 
following investigations reported here. 

Rhynencina is a small genus comprising 
only five New World species: three species 
are present in South America, one in Me- 
soamerica, and one, R. longirostris, in east- 
ern North America (Freidberg and Norrbom 
1999). The latter has been recorded from 
Pennsylvania, Maryland, North Carolina, 
and Georgia (Foote et al. 1993), and Ten- 
nessee (Steck and Sutton 2000). Limited 
host plant information is available for the 
genus. Rhynencina emphanes (Steyskal) 
(Colombia) has been reared from flowers of 
Espeletia sp.; adults of Rhynencina dyspha- 
nes (Steyskal) (Andean countries), Rhynen- 
cina spilogaster (Steyskal) (Mexico, Cen- 
tral America), and R. longirostris have all 
been collected on Polyinnia sp. and/or the 
closely related genus Sniallanthus; and R. 
spilogaster has been observed ovipositing 



into flowers of SwaUanilnis sp. (Fieidberg 
and Norrbom 1999) (host genera are all of 
Asteraceae, tribe Heliantheae); finally, R. 
longirostris was reared for the first time 
from Siuallantluts itvedolius (Steck and Sut- 
ton 2000) as more fully described below. 

Smallantluis iivedaliiis (formerly Polyin- 
iiia iivedalia). commonly known as leafciip 
or bearsfoot, is an herbaceous perennial that 
is widespread in woods and meadows of the 
eastern United States and flowers from 
summer to fall. Disk flowers are sterile, and 
thick achenes are produced from the ray 
flowers (Cronquist 1980). 


Most data were gathered from two patch- 
es of the host plant, Stnolhmthus iivedoUiis. 
in the GSMNP. one at Metcalf Bottoms and 
another at Sugarlands. The former stand of 
plants was spread along about 250 m of 
roadside and contained an estimated 1,300- 
1 .800 plants. The patch at Sugarlands was 
about 20 m X 3 m with an estimated 150- 
200 plants. 

Data on immature stages was obtained by 
dissections of flower and seed heads that 
were collected from 1999 to 2001. On 24 
and 27 September 1999, DD collected ma- 
ture seed heads from Metcalf Bottoms and 
Sugarlands, respectively, packaged them in 
brown paper bags, and sent them by over- 
night delivery to Gainesville, Florida; two 
additional collections of dried seed heads, 
from Sugarlands on 5 October, and from 
Metcalf Bottoms on 13 October 1999, were 
also sent to Gainesville in the same manner. 
In total about 50 dried seed heads were col- 
lected in 1999. Dissection of these heads 
revealed the presence of likely tephritid lar- 
vae inside achenes, as described more fully 
below. In an attempt to rear adults, several 
100s of unopened achenes were held dry 
and at room temperature in the laboratory 
until mid-November, 1999. At that time 
they were placed on damp sphagnum moss 
in a loose-lidded plastic container and kept 
in a refrigerator at about 4°C to break dia- 
pause. In mid-April, 2000, the achenes in 

their bed of damp sphagnum moss were 
transferred to a glass terrarium with 
screened top and bottom. At this time, some 
of the achenes were observed to be com- 
pletely softened and disintegrating, while 
others were still very hard as before. There- 
after the terrarium was kept in a shaded area 
outdoors under ambient Gainesville condi- 
tions (likely much warmer and drier than 
Smoky Mountains conditions), or alterna- 
tively under fluorescent lighting in a garage 
utility room with more moderate tempera- 
ture and humidity conditions. The terrarium 
was watered sporadically. 

Further dissections were made from seed 
heads collected from Metcalf Bottoms and 
Sugadands 4-8 September 2000 (Lionel 
Stange); from buds and flowering heads 
collected on 13 July 2001, Metcalf Bottoms 
(GJS, Victor M. Steck); and from buds, 
flower heads, and seed heads collected 2—4 
September 2001 (GJS, BDS), Metcalf Bot- 

Observation and collection of adult stag- 
es were made during 12-13 July 2000 
(GJS, BDS), 2 and 13 July 2001 (GJS, 
VMS), and 2-4 September 2001 (GJS, 

All life stages of R. longirostris are 
vouchered at the Museum of Entomology, 
Florida State Collection of Arthropods 
(FSCA) in Gainesville. A full description of 
all immature stages will be published at a 
later date. 


Rearing to adult stage. — Seeds held for 
overwintering in 1999 began germinating 
in mid-May, 2000. In mid-July the tenari- 
um was brought back into the laboratory in 
Gainesville for closer observation, and be- 
tween 24 and 27 July 2000 three adult R. 
longirostris flies emerged, thus proving 5. 
iivedaliiis to be a host of the immature stag- 
es. On 12 July 2000, GJS and BDS care- 
fully combed the litter below the host plants 
at Sugarlands looking for remains of the 
previous year's achenes. A few dozen intact 
achenes were located (along with numerous 



fragments) and held for rearing, but nnlhing 
came of them. 

Immature stages in flower and seed 
heads.— On 28 September 1999. GJS and 
BDS dissected and examined the contents 
of some of the seed heads collected by DD 
on -September 24 and 27 at Metcalf Bottoms 
and Sugarlands. respectively. There were 
no exposed tephritid immature stages pre- 
sent in the heads. Most seed heads retained 
a peripheral ring of black, dry, and well- 
hlled achenes (disk flowers are sterile, and 
a maximum of 11-12 seeds per head de- 
velop from the peripheral ray flowers). 
Achenes were very hard, and only with 
considerable difhculty did we manage to 
open them to examine the contents. Some 
achenes. often slightly lighter-colored and 
smaller, contained a bright yellow larva. In 
each case, a single larva filled the seed cav- 
ity, with only the thin, but hard, pericarp 
surrounding it. A total of seven 3rd instar 
larvae (based on mouthhook size; body 
lengths varied from about 1.6 to 3.7 mm) 
and five puparia was extracted in this fash- 
ion. Two t)f the puparia were held dry in a 
vial; in May 2001, it was noted that two 
adults of a Heleroscheina sp. (Hymenop- 
tera: Pteromalidae) had emerged. In addi- 
tion, of the three puparia preserved in al- 
cohol, two clearly contained hymenopteran 
larvae. Thus, the latter are likely larval-pu- 
pal parasitoids. There were no signs of exit 
holes or "windows" prepared by the R. lon- 
girostris larvae for later escape from the 
achene. The larvae were preserved in 70% 
alcohol after killing in hot water. Numerous 
seeds contained frass and tiny Lepidoptera 
larvae. There were also a few small, non- 
tephritid Diptera pupae in the dry seed 
heads, but not in the achenes. 

From the Sugarlands (5 October 1999) 
and Metcalf Bottoms (13 October 1999) 
seed head collections, we attempted to get 
a more quantitative estimate of their con- 
tents. To avoid destroying the achene con- 
tents, a small hand-held drill with a fine, 
bur bit was used to grind away enough of 
the pericarp to determine the contents. Of 

22 I achenes examined in this manner, a to- 
tal of 1 1 live R. longirostris 3rd instars. one 
dead 2nd instar. and 12 live pupae was ob- 
tained. The larvae had consumed the seed 
only. Two of the puparia were held dry for 
emergence — one yielded an adult Heteros- 
clieina sp., and the other an unemerged 
adult R. longirostris; five of the other ten 
puparia clearly contained hymenopteran 
larvae. Thus, this limited sample indicates 
a parasitism rate of 50%. Of the remaining 
achenes, 102 contained seemingly viable 
seeds (46%) and 89 appeared non-viable 
(empty or shriveled) (40%); another six 
achenes were full of insect frass (3%). The 
221 achenes examined were not selected 
randomly, as we were trying to maximize 
the number R. longirostris specimens 
found, and so chose smaller, discolored 
achenes: thus, these numbers do not accu- 
rately reflect actual infestation rates. See 
Table 1 for a summary of achene contents. 
Because of the time, effort, and difficulty 
encountered in dissecting dried achenes. the 
seed heads collected in year 2000 were 
treated differently: achenes were separated 
from their capitula. placed into a beaker of 
water, and gradually heated to about 70°C; 
after cooling they were transferred to 70% 
isopropanol. After a few months of storage 
in alcohol, the achenes were much more 
easily dissected with forceps without dam- 
aging their contents. One hundred achenes 
from each site were randomly selected and 
dissected. From the Metcalf Bottoms col- 
lection, these yielded 24 immature speci- 
mens of R. longirostris (three 1st instars, 
seven 2nd instars. four 3rd instars, ten pu- 
pae; nine of the ten puparia clearly con- 
tained a hymenopteran larva) (Table 1). As 
noted above, only a single R. longirostris 
larva or puparium was present in a given 
achene. Each of the immature specimens, 
excepting one of the 1st instars, was inside 
the seed coat. In the case of the 1st and 2nd 
instars, the surrounding seed coat was 
merely a shrivelled bag. (These could not 
be distinguished from the seed coat of nu- 
merous other inviable but uninfested seeds.) 


Table 1. Immature stages iil /?/n7(cm //ii; /i>ni;ini.\n-i.\ in seed heads of Sniullaiuhiis iivcilaliiis. 

1 fills 11,, ri'i'i scpi :iHiii luK :i)ni Si-pi :ni)i 

Nil. achenes e.xami 





ca. 1 1 7 

No. eggs 


No. 1st instars 



No. 2nd instars 




No. 3rd instars 



No. puparia 



% parasitized puparia 





Lepidoptera trass 



ca. 70 

+ Indicates present, but not quantitied. 

* Does not represent total damage from Lepidoptera leeding. because only intact achenes were selected for 

There wa.s nothing resembUng an obvious 
oviposition scar on the achene exterior, sug- 
gesting that oviposition occurs directly into 
the flower ovary at a very early stage of 
development. Of the 100 achenes dissected, 
only 32 contained what appeared to be vi- 
able seeds. Of the remainder. 1 1 had been 
destroyed by Lepidoptera larvae as evi- 
denced by their frass: other seed coats were 
empty or only partially filled for no appar- 
ent reason. Infested achenes were indeed 
smaller than those with viable seeds. Av- 
erage diameter of infested achenes was 4.7 
mm (range 3.3-5.4 mm. N = 24), while 
that of achenes with viable seed was 5.1 
mm (range 4.2-6.0 mm, N = 32). Infesta- 
tion at the Sugarlands site was very low: of 
100 achenes, only two contained R. longi- 
rostris (one 1st instar and one 2nd instar), 
while 71 contained apparently viable seeds. 

Plant collections made in year 2001 were 
treated somewhat differently from before. 
Entire capitula (rather than culled, intact 
achenes) were subjected to hot water treat- 
ment, then preserved in alcohol. This pro- 
vided data on R. loiigirostris egg deposition 
and the full impact of Lepidoptera larvae 
on seed production. 

On 2 July 2001 there were only a very 
few buds of S. iivedaliiis to be found at 
Metcalf Bottoms, and none was larger than 
about 5 mm in diameter. By 13 July, a small 
number of buds had developed to the stage 
where capitula disks were exposed and pet- 
als of ray flowers partially developed. All 

of the capitula (88 total) in the most ad- 
vanced stage of development were collect- 
ed, preserved in alcohol, and examined un- 
der a microscope (see Table 1). Oviposition 
and probe punctures, where an aculeus had 
penetrated the outer phyllaries and young 
achenes, were easily visible. Dissected 
heads ranged in size from unopened buds 
as small as 4 mm diameter to the most de- 
veloped capitulum in the population, which 
had a 12 mm-diameter receptacle (mea- 
sured with outer phyllaries removed) and 
petals of 12 mm length. Six capitula (7%) 
were found with eggs oviposited into 
achenes as follows: ( 1 ) two of ten achenes 
with one egg each in a 12 mm-diameter ca- 
pitulum with petals expanded, which was 
the most developed capitulum present at 
Metcalf Bottoms; (2) nine of nine achenes 
with a range of two to five eggs per achene 
in a 10 mm-diameter capitulum with outer 
phyllaries reflexed and one petal just begin- 
ning to unfurl; (3) five of 1 1 achenes with 
one egg each in a 10 mm-diameter capitu- 
lum with none of the petals unfurled; (4) 
four of nine achenes with one to two eggs 
each in a 10 mm-diameter capitulum with 
bracts partly reflexed and no petals; (5) four 
of 11 achenes with one egg each in a 10 
mm-diameter capitulum with none of the 
petals unfurled; and. (6) one of 12 achenes 
with one egg in a 9 mm-diameter capitulum 
with disk exposed and no petals. The 
achenes containing eggs ranged from 1.2 to 
2.8 mm long. Two capitula of 9 mm di- 


ameter had punctures that indicated probing 
by an aculeus but no eggs. There were no 
probing punctures found on achenes of un- 
opened buds or heads with a receptacle di- 
ameter of less than 9 mm. although a few 
probing punctures were found in the phyl- 
laries of smaller capitula (two 8-mm and 
one 6-mm capitula). None of the eggs had 
yet hatched. Thus, the few days before 13 
July at the time when capitula of 5. iivedali- 
iis first began to open and attained about 9 
mm diameter, marked the very beginning of 
the reproductive period of R. longirostris at 
Metcalf Bottoms for the year 2001. 

By 2 September 2001. the S. medal ins 
population at Metcalf Bottoms was found 
to be highly senescent, i.e.. fewer than 1% 
of all flower heads remained in bud or the 
flowering stage. No adult flies were ob- 
served at this time, but oviposition had oc- 
curred probably not long before, as 1st in- 
star larvae were still to be found in devel- 
oping achenes (Table 1). A notable obser- 
vation at this time was the very extensive 
feeding damage by larvae of a Homoeoso- 
ina sp. (Lepidoptera: Pyralidae). Of the 14 
capitula examined, 1 1 had been fed upon 
by Hoinoeosoimr, seven were 90—100% 
consumed, and three were 33-50% con- 
sumed. Thus, it is likely that some, perhaps 
many, late season R. longirostris immature 
stages were eaten by larvae of Homoeoso- 
ma sp. 

Observations on adult flies. — During 12- 
13 July 2000. GJS and BDS found male 
and female adult R. longirostris in abun- 
dance on 5. uvedaliiis foliage and flower 
heads at the Metcalf Bottoms site. Plants 
were well developed at this site, many to 1 
m or more in height, and some were in 
fiower A smaller number of adult R. lon- 
girostris was observed at Abram's Creek 
campground, also on S. uvedaliiis. but no 
adults were found at the Sugarlands site, 
where S. iivedalius was not yet in flower 

On 2 July 2001, very few or no flower 
buds were present at Metcalf Bottoms and 
Sugarlands, respectively, and no adults 
were observed. On 13 July 2001. one fe- 

male and three males were observed at 
Metcalf Bottoms. One male was observed 
mounted on the female on the most devel- 
oped flower head at this site, and a second 
male was facing them on the periphery of 
the same flower head. Therefore, courtship 
and mating apparently occur on the host 
plant, as is typical for many tephritid flies. 
By 2 September 2001. no adults were to be 
found at Metcalf Bottoms, where flowering 
of host plants was nearly finished. However, 
in other areas, such as Cades Cove and 
Abrams Creek campground, many plants 
were still in full flower, and adult R. lon- 
girostris were collected at Abrams Creek 
campground on 4 September 2001. 


Based on our rearings and collections 
from 1996 to 2001, we conclude that R. 
longirostris is univoltine with a flight-time 
corresponding to the flowering period of S. 
uvedaliiis, i.e., July-September in the 
GSMNP. Local populations of 5. iivedalius 
flower at somewhat different times. It re- 
mains to be determined whether the emer- 
gence and disappearance of the associated 
fly populations are locally synchronized 
with their hosts, or whether flies disperse 
among local host populations according to 
the presence/absence of capitula at an ap- 
propriate stage for oviposition. Courtship 
and mating occur on the host plant. Ovi- 
position probably begins as soon as achenes 
of S. iivedalius develop to the minimum 
size needed to support larval development. 
At the beginning of the flowering season, 
when buds are scarce, a single achene may 
have several eggs oviposited; thereafter, fe- 
males lay only a single egg into an achene. 
Only one larva develops per achene. Para- 
sitism by hymenopteran larval-pupal para- 
sites may be very high. Given the extreme 
hardness of the mature, dry achenes, hy- 
menopteran parasites may be able to attack 
only during the early stages of achene de- 
velopment. Late in the flowering season. 
Lepidoptera larvae feed voraciously on ca- 
pitula and probably also inflict significant 



mortality on immature stages of R. longi- 
rostris. Many achenes from the preceding 
year's seed crop that were found under host 
plants at Sugariands appeared to have been 
cracked open leaving only very hard frag- 
ments. It is possible that vertebrate seed 
predation is another significant cause of 
mortality, particularly if rodents, such as 
squirrels or mice, or seed-feeding birds, are 
targeting the slightly more easily opened, 
protein-rich, infested achenes. By what 
means the adult flies escape the achene re- 
mains unknown. Dried achenes are very 
hard and it is difficult to imagine an adult 
extricating itself without a pre-thinned 
"window' in place. Perhaps they rely on 
simple weathering from the elements to de- 
compose the pericarp to a point where 
adults can eclose and escape successfully. 


We thank Keith Langdon, Inventory and 
Monitoring Coordinator for GSMNP. for fa- 
cilitating our studies in the park. Lionel 
Stange (FSCA) for a sample of 5. iivedalius 
seed heads in year 2000. Steve Heydon 
(Bohart Museum. University of California. 
Davis, CA) for identification of parasitoids. 
John Heppner (FSCA) for identification of 
Lepidoptera larvae, and Al Norrbom 

(USDA, ARS. Systematic Entomology 
Laboratory, Washington, DC) for reviewing 
the manuscript. We also thank Ms. Jeanie 
Hilton and the others at Discover Life In 
America for providing accommodations to 
support the field work for this project. This 
research was conducted under National 
Park Service permits GRSM-00-103 and 
GRSM-200I-SCI-0153. All specimens are 
vouchered in the Natural History Museum 
(GSNP) and/or the Museum of Entomolo- 
gy. Florida State Collection of Arthropods. 
Gainesville. Florida. Contribution No. 929. 
Bureau of Entomology. Nematology and 
Plant Pathology — Entomology Section. 

Literature Cited 

Cronquist, A. 1980. Vascular flora of the southeastern 
United States, Vol. 1. Asteraceae. University of 
North Carolina Press. Chapel Hill, North Carolina. 
261 pp. 

Foote, R. H., F L. Blanc, and A. L. Norrbom. 1993. 
Handbook of the fruit flies (Diptera: Tephritidae) 
of America north of Mexico. Cornell University 
Press, Ithaca, New York. 571 pp. 

Freidberg, A. and A. L. Norrbom. 1999. A generic 
reclassification and phytogeny of the tribe My- 
opitini (Tephritidae), pp. 581-637. In Aluja. M. 
and A. L. Norrbom. eds. Fruit Flies (Tephritidae): 
Phylogeny and Evolution of Behavior. CRC Press, 
Boca Raton, Florida. 944 pp. 

Steck. G. J. and B. D. Sutton. 2000. New records for 
Tephritidae (Diptera: Tephritidae) in Great Smoky 
Mountains National Park. Insecta Mundi 14: 256. 


105(3), 2003, pp. 548-5.54 


Di;HR A I.. Ml'RUA> 

Departnient of Entomology, Oregon Slate Llniversiiy, Corvallis, OR 97.^.^1. U.S.A. (e- 
niail: nuinaydefe'bcc.orst.edii) 

Ahstnnt. — FoMtciyi^ctix pciiclca Cramer lar\ae are common in hamlioo luulersiory, pre- 
sent year round, and can be found in a wide variety ot habitats. Larvae specialize on 
woody bamboos within the genus Giuuliui (Poaceao). and adults are highly associated 
with bamboo groves. The presence of the host plant is the only important habitat require- 
ment for this species, in that they can be found within bamboo groves in forest habitats, 
forest edges, and pastures. The immature stages of P. iwiwleci are described and head 
capsules and chaetotaxy are illustrated. Both laivae and pupae possess distinct features 
which separate this monotypic genus from the closely lelatetl genera Tcixiictis lliibncrand 
Pscuthnlchis Forster. 

Ki\ Wonls: 

luiptychiina. Satyrinae. TdXiiclis 

l\scu(h)(liliis. bamboo. Iar\ae. s|-)ecialist. 

Posttaxiii'tis Forster (1964) is a monotyp- 
ic butterlly genus within the diverse satyr- 
ine subtribc Euptychiina (Nymphalidae: .Sa- 
tyrinae). Posttaxfii'tis peneleu (Cramer 
1777) occurs throughout the Amazon basin 
to southern Brazil and was recently col- 
lected as far north as Nicaragua (Maes 
1995). Individuals are normally enciiun- 
tered within lowland Gnadtia bamboo 
groves (Poaceae), although 1 collected a 
worn specimen from premontane forest 
(Wilson Botanical Garden, Costa Rica, el- 
evation 1,1 10 m). 

Posttaxi>ctis /H'lU'U'd is easy to recognize 
by the combination of somber satyrine 
markings and a bright yellov\ band travers- 
ing the ventral hind wing (Fig. I). This yel- 
low band is greatly reduced or absent in the 
one named form. P. pcnelea van peneliiui 
(Staudinger IS8S). found in southern Brazil 
(Fig. 2). Forster (1964) provided no infor- 
mation on delineation of the Posttaxgetis 
other than commenting; that it is distin- 

guished by structures of the male genitalia. 
However, larvae possess several distinctive 
features which separate this species from 
the closely related genera Taxiii'tis Hiibner 
and Pseitdoili'his Forster. These diagnostic 
characters, along with the immature stages, 
are described in this paper. 

Mai'i;ri.'\ls and Methods 

Larvae and eggs were located in tiic tield 
by searching known host plants and by ob- 
serving female oviposition. Specimens 
were reared individually in plastic contain- 
ers, and fresh host plant was added every 
two to three days. Appearance, behavior, 
and development times were noted for all 
stages. Larvae were preser\ed in 95'/^ eth- 
anol after being immersed in boiling water 
for se\eral seconds. Vouchers are dept>sited 
in the Natit>nal Museum of Natural History, 
Smithsonian Institution, Washington DC. 
Much of the work lV>r this study was con- 
ilucled in Ecuador at Jatuu Sacha Biological 

V()[AIME 105, NUMBER 3 

1-igs. 1-2. Adulls, dorsal view on Icll. sciiliiil \ lew mi riiilu, I. l'(niniYt;fti\ /itiic/ca. mnW, Ixuador, Napii 
lov.. Puerto Misahualli. 2. /'. pciicUa var pciwIiiKi. iiiale. Bra/il. Mato Grosso, Cliapada |BNHM|. 

Slalion. Napo PiDvincc, during 1990 to 
1993. Some comments on adult natural his- 
tory, in particular observations on hahilat 
associations, are taken from data gathered 
tiuring a year long trapping study conducted 
at that site (DeVries et al. 1997). 

.Setal names for the head capsule follow 
Heinrich (1916), with modifications as in- 
coiporated by Stehr (1987). Body chaeto- 
ta.xy follows Hinton (1946). More detailed 
information on euptychiine morphology can 
be found in Murray (2001). Head capsule 
width is measured by the distance between 
the third stemmata. Instar durations are giv- 
en as averages of five individuals. 

Posttaygetis peiwlea Cramer 

Diagnosis. — FirM instar: No unique fea- 
lures separate species from other euptychi- 
ines other than combination of larger si/e. 
head capsule primary setae terminating in a 
line point, and extremely dense pattern of 
ridges on head capsule surface, which ex- 
tend medially to A2 (Fig. 3). 

Second and third instar: White band bi- 
secting dorsal aspect of head capsule, ex- 
panding medially; four large apical setae on 
head horns, P 1 , P2, and two secondary se- 
tae (Fig. 4-.'i); slender green body with 
white, broad dorsolateral stripe: caudal hl- 
aments projecting dorsolaterally. 

Foiirlli instar: White band bisecting dor- 
sal aspect of head capsule; head horns and 
lateral lobe setal bases elongated; tour large 
apical setae on head horns, PI, P2, and two 
secondary setae with large setal bases and 
long, recurved setae (Fig. 6); specialized se- 
tae bordering epicranial suture terminating 
at apex of adfrontal area; body with green 
and brown dorsolateral slashes forming 
broad inverted "v" in dorsal view. 

Pupa: Four prominent pairs of tubercles 
dorsally on abdcimen (Fig. 10). 

Description. — Egg: .Shiny, round, semi- 
transparent, pale white to pale green (n = 
28). Duration 6.0 days. 

First instar: Head capsule 0.8 mm; 
black; lobe setae (PI, P2, P.^, LI, and A3) 
long, narrow, terminating in a Hne point; 
lobe setae basally thick but not flattened; PI 
located more basal than P2; dense dendritic 
pattern of ridges on the head capsule integ- 
ument (Fig. 3). Body shiny dark green; Dl 
and D2 shorter than XDl and XD2 on pro- 
thoracic segment (Fig. 7); SDl on Tl and 
A9 sensory with long, filamentous seta; Dl 
and D2 approximately equal on all abdom- 
inal segments except A8 and A9, where Dl 
larger than D2 (Fig. 8); caudal filaments 
short, rounded (n = 17). Duration 4.8 days. 

Second instar: Head capsule 1 .2 mm; 
light brown darkening to brown one to two 
days after nu)lting; head horns and lateral 


Figs. 3-6. Head capsules of Posttaygetis pene/cii. frontal view (scale 
instar. 5. Third inslar 6. Fourth instar. 

2.0 mm). 3. First instar. 4, Second 

head capsule dark brown; apical portion of 
frons outlined in dark brown; lateral setal 
lobes white; distinctive white band extend- 
ing between P3 setae, widening medially 
(Fig. 4); head horns large, projecting slight- 
ly anteriorly, with two large apical second- 
ary setae in addition to PI and P2; lobe 
setae long, flattened, and twisted, at least as 
long as sclerotized base; P3 thinner and 
shorter than other setae; lateral lobe setal 
bases white; S2 setae long, flattened, and 
twisted. Body bright green with bright 
white dorsolateral stripe; caudal rtlaments 
pink and projecting dorsolaterally; numer- 

ous secondary setae on body; primary setae 
indistinguishable from surrounding second- 
ary setae except for SDI sensory setae on 
TI-T3, A4, A5, and A9; integument cov- 
ered with fine microgranulations (n = 14). 
Duration 4.9 days. 

Third instar: Head capsule 1.9 mm; 
same as in previous instar except dark 
brown, basally reddish brown; head horns 
with white basal stripe; P3 considerably 
smaller than other lateral lobe setae (Fig. 5). 
Body dark green; median dorsal stripe faint 
reddish green, outlined by undulating white 
lines on abdomen only; dorsolateral band 



Figs. 7-8. Body chaetmaxy of first instar P,:snayfiL'ti\ peiwlca. 1. Tl-Al. X, A6-AI(). 

broad, bright white to yellow, and extend- 
ing laterally on caudal filaments: thin white 
line laterally; caudal filaments pink apically 
and projecting dorsolaterally: body chaeto- 
taxy same as previous instar (n = 18). Du- 
ration 5.0 days. 

Fourth instar: Head capsule 3. 1 mm; 
dark brown; reddish brown basally and lat- 
erally; white basal stripe; white band as in 
previous instar; PI and P2 flattened, re- 
curved, and one and a half times length of 
setal base; two secondary apical setae on 

head horns as large as PI and P2; special- 
ized setae bordering epicranial suture from 
apex of adfrontal area to head capsule base, 
curving over suture and criss-crossing setae 
on opposing side (Fig. 6). Body pale brown 
(Fig. 9); prothoracic shield green; dorsal 
median stripe on thorax rust brown outlined 
thinly in dark brown; dorsal median stripe 
on abdomen green, often fading to greenish 
brown as instar progresses, outlined with 
thin white undulating lines; green colora- 
tion extends dorsolaterallv on each seg- 


9-10. Posriaygelis penelea. 9, Mature larva, dorsal view. 10. Pupa, lateral view. 

ment, forming a broad "v" pattern in dorsal 
view; fine lines of brown and white later- 
ally; black spots dorsolaterally. two per seg- 
ment, on T2. T3, Al. and A9; spots variable 
in size and may not be present; large prom- 
inent dark brown spot laterally on A3 to 
A4, tapering smoothly to form a line on A5 
through A6 and bordered ventrally by 
bright cream line; caudal filaments green; 
chaetotaxy of SDl sensory setae same as 
previous instar; secondary setal bases on 
body covered in fine microgranulations: 
specialized setae within thoracic dorsal 
midline stripe, setae similar in morphology 
to specialized setae on head capsule, not 
found elsewhere on body (n = 23). Dura- 
tion 8.8 days. 

Pupa: Dark brown with green and pale 
brown markings (Fig. 10); compressed; 
dorsal apical point on thorax knobbed, pro- 
jecting anteriorly; four prominent tubercles 
dorsally on abdomen (n = 12). Pupates flat 
against substrate. Duration 9.8 days. 

Biology. — Posttaygetis penelea adults 
are encountered in and around Guadiia 
bamboo groves. At Jatun Sacha, Ecuador, 
adults were present in all months of the 
year, but were never abundant (DeVries et 
al. 1997). All individuals trapped were from 
sites in forest habitats containing bamboo 
(n = 25). Within the bamboo groves males 
defend small sunlit patches but do not ex- 
hibit lekking behavior that occurs in many 
Taygetis species (Mun^ay 2001). Females 
oviposit in late afternoon, usually from 4:00 

until dusk, and often remain in small, lo- 
calized areas to select oviposition sites. 

Larvae specialize on bamboo (Giiadua 
sp.). Mature larvae provided with two her- 
baceous bamboos (Olyra lotifolia L. and 
Pciricina sp.) and one grass species (Ortho- 
clacla la.xa (Rich) P. Beau v.) did not feed. 
Eggs are laid singly on small bamboo 
shoots in the understory. Although bamboo 
groves are comprised of a substantial 
amount of plant biomass, most is in the can- 
opy, with only isolated resprouts, young 
plants, or new growth in the understory. For 
butterflies restricted to understory sites, the 
number of oviposition sites can be limiting. 
There can be up to a half dozen P. penelea 
eggs on the underside of some leaf blades. 

Although adults are not commonly en- 
countered in the field, the immature stages 
of P. penelea are often easily located in 
suitable bamboo habitat. This species does 
not appear to go into reproductive diapause 
in the upper Napo region of Ecuador, as lar- 
vae are present year round. Systematic 
searches for larvae were conducted at Jatun 
Sacha, including bamboo groves located in 
forests, forest edges, and open pastures. 
Posttaygetis penelea larvae were found in 
all habitats and were the only euptychiine 
bamboo feeders found in isolated bamboo 
groves in pastures. 


Despite the paucity of information pro- 
vided by Forster ( 1964), several unique lar- 



val characters were found that support Posi- 
laygeiis as a vaHd genus. Posttaygctis pe- 
nelea was originally placed within Taygetis. 
and the larvae share many moiphological 
traits. Most notable of these traits are the 
retention of both PI and P2 setae at the 
apex of the head horns and the enlarged pri- 
mary setae on the head capsule. However, 
P. peuelea differs from closely related spe- 
cies by the presence of two large apical sec- 
ondary setae between PI and P2 on the 
head horns, the distinct white or dark brown 
band transversing the head capsule, and the 
widely spaced caudal Hlaments, which pro- 
ject dorsolaterally, albeit weakly, in later in- 
stars. In the fourth instar, the distribution of 
specialized setae bordering the epicranial 
suture is also an important diagnostic char- 
acter, with the setae terminating at the apex 
of the adfrontal suture. Other euptychiines 
either do not possess these setae, or the se- 
tae are found on the adfrontal area and/or 
fions. Posttaygetis penelea pupae differ 
from Taygetis and Pseiidodehis pupae by 
their compressed shape and large tubercles 
on the abdominal dorsal surface. There are 
differences in ecological traits as well. Most 
Taygetis species are crepuscular and many 
males exhibit lekking behavior (Young 
1984. Munay 2001 ). In contrast, P. penelea 
is diurnal and males do not form leks. Also, 
females of Taygetis bamboo specialists ovi- 
posit at dusk in the canopy, not in the un- 
derstory as P. peuelea. 

There is only slight color change within 
each instar as the larvae develop, a trait 
commonly found in Taygetis larvae (Mur- 
ray 2001). Unlike many satyrines. P. pe- 
nelea larvae and pupae are not polymor- 
phic. Other than the previously noted 
named form, P. penelea van penelina. there 
is also little variation in wing pattern, which 
can be highly variable in related Taygetis 
species. This form is reported to occur near 
the southern edge of the species' distribu- 
tion, but I have collected individuals from 
the Ecuadorian rainforest that also lack the 
distinctive band, although I have never 
reared an individual. Most likely this form 

is present in many P. penelea populations, 
but in low numbers. 

The fact that the immature stages of P. 
penelea have not been described yet is 
somewhat surprising. As noted earlier, lar- 
vae are common in the bamboo understory, 
present year round, and can be found in a 
wide variety of habitats, including pasture, 
as long as bamboo is present. Indeed, the 
presence of bamboo is the only important 
habitat requirement, suggesting that this 
species is not heavily impacted by habitat 
disturbance. This result was also found by 
Singer and Ehrlich ( 1991 ) for several other 
specialist euptychiines. 


Alejandro Suarez and Gabriel Tapuy pro- 
vided valuable assistance in collecting and 
rearing larvae. I also thank Gerardo Lamas, 
Giovanni Onore, the Organization for Trop- 
ical Studies, and Jatun Sacha Biological 
Station for assistance with logistics and 
fieldwork. David Neil identified host plants 
and Jon Seals translated sections of manu- 
scripts, and these services were much ap- 
preciated. Helpful suggestions to this man- 
uscript came from Chris Carlton, Gerardo 
Lamas. Lee Miller, and Dorothy Prowell. 
The following organizations provided in- 
valuable support for this research project: 
Fundacfon Jatun Sacha, the United States 
Peace Corps, the Organization for Tropical 
Studies, and American Women in Science. 

Literature Cited 

DeViies. P. J.. D. L. Murray, und R. Landc. I9')7. Spe- 
cies diversity in verticaL hori/ontaL and lempcMal 
dimensions of a fruit-feeding butterfly community 
in an Ecuadorian rainforest. Biological Journal of 
the Linnean Society 62(3): 343-364. 

Forster. V. W. 1964. Beitrage zur Kenntnis der Insek- 
tenfauna Boliviens XIX. Veroffentlichungen der 
Zoologischen Staatssammlung. Miinchen 8; 31- 

Heinrich, C. 1916. t)n the taxonomic value of some 
larval characters ni the Lepidoptera. Proceedings 
of the Entomological Society of Washington 18: 

Hinton, H. 1946. On the homology and nomenclature 
of the setae of lepidopterous larvae, with some 


notes on the phylogeny of the Lepidoptera. Trans- 
actions of the Entomological Society of London 
97: 1-37. 

Maes, J.-M. 1995. Fauna entomologica de la Reserva 
Natural Bosawas, Nicaragua. IV. Taygetis penelea 
(Cramer) nuevo para la fauna de Nicaragua (Lep- 
idoptera: Nymphalidae: Satyrinae). Revista Nica- 
ragua Entomologico 35(5): 5-8. 

Murray, D. L. 2001. Immature stages and biology of 
Taygetis HCibner (Lepidoptera: Nymphalidae). 
Proceedings of the Entomological Society of 
Washington 103(4): 932-945. 

Singer. M. C. and R R. Ehrlich. 1991. speciali- 
zation of satyrine butterHies. and their responses 
to habitat fragmentation in Trinidad. Journal of 
Research on the Lepidoptera 30(3-4): 248-256. 

Stehr, F W. 1987. Order Lepidoptera, pp. 288-305. In 
Stehr. F. W., ed. Immature Insects. Kendall/Hunt 
Publishing Co., Dubuque, Iowa. 754 pp. 

Young, A. M. 1984. Natural history notes for Taygetis 
cmdromeda (Cramer) (Satyridae) in eastern Costa 
Rica. Journal of the Lepidopterists' Society 38(2): 
102-1 13. 


103(31. 2003. pp. 555-558 


David R. Smith and Malkiat S. Saini 

(DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- 
tution. Washington, DC 20560-0168, U.S.A. (e-mail:; (MSS) 
Department of Zoology, Punjabi University, Patiala 147002, India (e-mail: saini20@ 
glide. net. in) 

Abstract. — Coiiaspidia maculosa, new species, is described from Sikkim, India. It is 
the fourth species of this Asian genus known from India, the other species being C. 
clubiosa Forsius, C. fasciatipennis Turner, and C. sikkimensis Konow. The new species 
belongs in the scuteUaris group of Coiiaspidia and is separated from related species in 
that group and the other Indian species. 

Key Words: India. Coiiaspidia. sawfly. Tenthredininae 

Coiiaspidia Konow is an exclusively 
eastern and southeastern Asian genus of the 
subfamily Tenthredininae. It is known from 
eastern Russia and Japan, through China to 
northeastern India. Myanmar, and Vietnam. 
The Hrst compilation of species with a key 
was that by Malaise (1945). who treated 
eight species. In the most recent treatment. 
Wie and Nie (1997) recognized five groups 
and keyed 18 species. Here we describe a 
fourth species from India, the others being 
C. sikkimensis Konow 1898 (type species), 
C. fasciatipennis Turner 1919, and C. dii- 
hiosa Forsius 1931, all recorded from Sik- 
kim. The genus has been characterized by 
Malaise (1945) and Wie and Nie (1997). 

Coiiaspidia maculosa Smith and Saini, 
new species 

(Figs. 1-4) 

Female. — Unknown. 

Male. — Length, 10.0 mm. Antenna yel- 
low with apical 3 segments black dorsally. 
Head and mouthparts reddish brown with 
black markings as follows: postocellar area. 

large spot lateral to postocellar area; small 
spot lateral to lateral ocellus: stripes from 
clypeus to just above antennae between an- 
tenna and eye and between antennae, con- 
nected by a transverse stripe dorsal to an- 
tennal insertions. Thorax black with cervi- 
cal sclerites laterally, dorsal half of prono- 
tum, tegula, and parapterum reddish brown: 
small spots on mesonotal lateral lobe 
opposite mesoprescutum, on lateral margin, 
and on metanotum posterior to each cen- 
chrus. Metanotum lateral to cenchri reddish 
brown. Abdomen orange with anterior mar- 
gin of first segment, anterior margin of 2nd 
segment and segments 5 to apex black; api- 
cal tergum reddish brown at center. Legs 
orange yellow with hind tibia blackish. 
Forewing yellow orange with black macula 
at base of wing, black stripe from stigma to 
posterior margin at center of wing, and 
large black macula at apex (Fig. 3); base of 
costa reddish brown, all other veins and 
stigma black, with veins orange yellow in 
orange-yellow portion; hind wing uniform- 
ly clear 


PROCi:i;niN(;s op iiii' I'Ntomoi ocikai. socihty oI' Washington 

I 3. (\tiiii\/'iiiiii 

■ili'ul. I, Dois;il 

AiUcniui. 3, I'DTVwiii" 

Anicnna (I'it:. 2) tililomi. Iciiylh I.7X 
head width; scape slightly less than 2x lon- 
ger than hioad anil longer than pedicel; ped- 
icel slightly longer than brt)ad; 3rd segment 
1.3X longer than 4lh segment. Head im- 
punclate, shining; and broad behind eyes in 
dorsal view (hig. I); eyes slightly converg- 
iiiu below, with lower inlerociilar ilislance 

I.IX eye length. Clypeiis vsilh deep emar- 
gination about half its medial length, lateral 
lobes narrow and rounded; labrum about 
2X broader than long, anterior margin 
rounded; malar space nearly linear, less than 
half tliameter oi t'ronl ocellus; postocellar 
area about l.2x longer than broad; genal 
carina jircscnt. extentliug to top ol head but 


Fig. 4. Male genitalia of Coruispidia inacuhisu: 
ventral view of left half of genital capsule on left, lat- 
eral view of penis valve on right. 

iwt including postoceilar area. Mesopres- 
cLitum and mesonotal lateral lobes shining, 
with fine, widely spaced punctures; meso- 
scutellum in lateral view produced as a high 
conical projection, its height equal to length 
of posttergite, with deep, large punctures 
laterally and separating scutellum from 
posttergite; posttergite uniformly punctate. 
Punctures on mesepisternum large, close. 
iiTegular. extending to mesosternum; me- 
sosternum not punctate, shining; mesoster- 
nal area rounded, without mesosternal pos- 
lero-lateral thorn or projection. Hind basi- 
tarsus about 0.8 x length of following tarsal 
segments combined, about equal to follow- 
ing three segments. Inner tooth of tarsal 
claw longer than outer tooth and placed lat- 
eral to it on inner side. Abdomen shining, 
without surface sculpture. Hind wing with- 
out peripheral vein; cells M and Rs present; 
anal cell petiolate. with petiole length about 
equal to widest width of anal cell. Genitalia 
in Fig. 4. 

Holotype. — S. Chungthang. Sikkim. hi- 
dia. May 1995. 1800 m. M.S. Saini. De- 

posited in the National Pusa Collections, 
Division of Entomology. India Agricultural 
Research Institute. New Delhi. 

Etymology. — The species name is from 
the Latin maculosiis. spotted, referring not 
only to the three maculae of the wings but 
extensive black markings on the head and 

Discussion. — This species belongs to the 
scutelloris group of Wei and Nie (1997). 
The sciitellaris group includes C. liiiiuitci 
(Matsumura 1912) from Japan, C. kuhipan- 
acis Xiao and Huang 1983 from China. C. 
liiii Wei 1997 from China, C. miirotai To- 
gashi 1984 from Japan, and C. scitteUahs 
Malaise 1945 from China and Myanmar. 
The sciitellaris group is separated from oth- 
er groups of Cojuispidia by the strongly in- 
fuscate cross bands of the forewing with 
black maculae at the base and apex, the bas- 
al tooth of the left mandible small and ob- 
tuse, the scutellum strongly elevated, and 
the lance without a spine near the aiinuli 
(Wie and Nie 1997). 

This new species will key to C. liui Wei 
from Hunan. China, in the key to species 
by Wie and Nie (1997), as it shares the fol- 
lowing characters: short hind basitarsus. 
shorter than the following tarsal segments 
combined; third antennal segment longer 
than the fourth; hind wing of the male with- 
out a peripheral vein; head subparallel be- 
hind the eyes; maculation of the forewing; 
elevated mesoscutellum; small, obtuse basal 
tooth of the left mandible; punctures later- 
ally on the mesoscutellum; dense, large 
punctures of the mesepisternum; impunc- 
tate mesosternum; and inner tooth of the 
tarsal claws longer than the outer tooth. The 
genitalia are also very similar to C. liui (see 
Wie and Nie 1997. fig. 37) though they are 
difficult to compare since they might not be 
drawn from exactly the same aspect. The 
new species differs from C //(// by the 
black apical segments of the antenna, black 
mesonotum, black mesopleuron, black an- 
terior margins of the first and second ab- 
dominal terga, black apical segments of the 
abdomen, lower interocular distance slight- 


ly longer than eye length, postocellar area 
longer than broad, and more slender, and 
curved penis valve. 

Very few Coiuispidia have the inner 
tooth longer than the outer tooth. This oc- 
curs only in C. liiii and C. latifasciata Wei 
1997. the latter assigned to the biciispis 

The coloration of C. maculosa is very 
similar to C. guttata, but C. guttata has 
large, isolated punctures on the mesepister- 
num. mesosternum with dense small punc- 
tures, the inner tooth of the tarsal claws 
shorter than the outer tooth, and mesoscu- 
tellum lower and more convex and smooth- 
er without the large lateral and posterior 

Species in India 

In India. Conaspidia is known only from 
Sikkim. Conaspidia maculosa is the only 
species with the basal, central, and apical 
maculae of the forewing. Conaspidia sik- 
kinwnsis has only the apex of the forewing 
infuscated. C. fasciatipennis has only the 
central band and apical macula, and the 
forewing of C dubiosa is uniformly yel- 
lowish. The species are included in the keys 
by Malaise (1945) and Wie and Nie (1997). 

Conaspidia dubiosa. — Both sexes of this 
species were described from "Sikkim"" by 
Forsius (1931). The forewing is uniformly 
yellowish, the third antennal segment is 
slightly shorter than the fourth segment, and 
the hind basitarsus is longer than the re- 
maining tarsal segments combined. We 
have not seen additional specimens. 

Conaspidia fasciatipennis. — This species 
was described from "Tong Lap. Haul Me- 
kong" by Turner (1919). Only the female 
was treated by Turner (1919) and Malaise 
(1945): however. Wie and Nie (1997) in- 
dicated that both sexes are known. It was 

recorded from "Sikkim" by Wie and Nie 
(1997). The forewing has the central dark 
band and apical macula but lacks the basal 
macula present in C. maculosa. We have 
not seen specimens from India. 

Conaspidia sikkimensis. — Konow (1898) 
described this species from "Sikkim." He 
described only the female, and only the fe- 
male was included by Malaise (1945), but 
Wie and Nie (1997) stated that both sexes 
are known. Only the apex of the forewing 
is black. The species is Hgured by Malaise 
(1945, pi. lib). We have seen the following 
specimens from India: Sikkim, Gangtok, 
2200 m, 12.V.1983 (2 $, 6 cJ). 


We thank US PL-480, Project No. FG- 
In-753, and ICAR. New Delhi, for financial 
assistance. Cathy Apgar. Systematic Ento- 
mology Laboratory, USDA (SEL), took the 
photos and arranged the plates. We also 
thank the following for review of the man- 
uscript: H. Goulet. Agriculture and Agri- 
Food Canada. Ottawa: and N. J. Vanden- 
berg and M. G. Pogue. SEL. 

Literature Cited 

Forsius. R. 1931. Uber einige neue oder wenig be- 
kuiinte orientalische Tenthredinoiden (Hymen- 
opt.). Annalen des Naturhistori.schen Museum in 
Wien 66: 29-48. 

Konow. F W. 1898. Neue Chalastogastra-Gattungen 
und Arten. Entomologische Nachrichten 24: 268- 

Malaise. R. 1945. Tenthredinoidea of south-eastern 
Asia. Opuscula Entomologica. Supplementum IV, 
288 pp. + XX plates. 

Turner. R. E. 1919. XLVIIL— On Indo-Chinese Hy- 
menoptera collected by R. Vitalis de Salvaza. — II. 
Annals and Magazine of Natural History (9)3: 

Wei. M.-C. and H.-Y. Nie. 1997. Studies on the genus 
Conaspidia Konow (Hymenoplera: Tenthredini- 
dae) from China with a key to known species of 
the world. Entomota.xononiia 19: 95-117. 

105(3), 2003. pp. 559-567 


Agustin Aragon and Miguel-Angel Moron 

(AA) Departamento de Investigacion en Ciencias Agn'colas, Institute) de Ciencias. Be- 
nemerita Universidad Autononia de Puebla, Apdo. Postal 1292, Ciudad Universitaria. 
Puebla. Puebla 72370, Mexico; (MAM) Departamento de Entomologia, Instituto de Ecol- 
ogia, A.C. (SEP-CONACYT). Apdo. Postal 63. Xalapa, Veracruz 91()()(). Mexico (e-mail: ) 

Abstract. — Phyllophaga {Chlaenohia) rzedowskiana, new species, is described from 
remnants of deciduous tropical forests at 940-1.145 m altitude at seven localities of 
southwestern Puebla and eastern and southeastern Morelos, Mexico: it is included in the 
species group "ve.xcita." Phyllophaga (IJstrochelus) harrerana, new species, is described 
from remnants of deciduous tropical forests at 1,000- 1. 2 10 m altitude at five localities of 
southern State of Mexico, southeastern Morelos, and southwestern Puebla, Mexico; it is 
included in the species group "cavata." Drawings of the head, pronotum, abdomen, tarsal 
claws, and genital structures are provided. 

Resiimen. — Se describe Phyllophaga (Chlaenohia) rzedowskiana nueva especie, pro- 
cedente de los remanentes de bosques tropicales caducifolios localizados entre los 940 y 
1.145 m de altitud en siete localidades del suroeste de Puebla, del este y sureste de 
Morelos. Mexico; la que se agrega al grupo de especies ^^vexata." Tambien se describe 
Phyllophaga (Listrochelus) harrerana nueva especie. procedente de los remanentes de 
bosques tropicales caducifolios ubicados entre los 1.000 y 1.210 m de altitud en cinco 
localidades al sur del Estado de Mexico, el sureste de Morelos y el suroeste de Puebla, 
Mexico; la cual se incorpora al grupo de especies "cavata." Se incluyen ilustraciones de 
la cabeza, pronoto. abdomen, uiias tarsales y estructuras genitales. 

Key Wonl.\: May beetles, Phyllopluii^a, taxonomy, deciduous tropical forests, Mexico 

A large area between the Mexican states dre del Sur, and species from the xeric 
of Morelos, Puebla, Guerrero, and Oaxaca scrubs extended across the Tehuacan, Acat- 
(approximately 17°30'-19°N 98°-99°45'W) Ian, and Huajuapan valleys, 
is known as the upper basin of the Rio Bal- Thirty species of the genus Phyllophaga 
sas. This area is especially interesting be- Hanis (sensn lata) have been listed from 
cause it represents a transition between spe- southwestern Puebla (Moron and Aragon 
cies from the tropical deciduous forests, 1997; Moron 1992, 1998; Aragon et al. 
widely extended along the lower Pacihc 1998; Moron et al. 2000), and twenty spe- 
slopes and lower basin of the Balsas River, cies have been found in southeastern Mo- 
species from the pine-oak forests located relos (Deloya and Moron 1994. 1998; 
along the Neovolcanic Axis and Sierra Ma- Perez-Garcia 1999). but the adjacent areas 


of Guerrero and Oaxaca included in the up- 
per basin of the Balsas River remain poorly 

Field work during 1999 at Puebla local- 
ities and curatorial work of some Mexican 
collections during 2000-2002 led to the 
discovery of two undescribed species of 
Phyllopha^u. apparently restricted to the 
upper basin of the Balsas River. Below, we 
give descriptions of males and females of 
one new species of P. {Chlaetwbia Blan- 
chard). and males and females of one new 
species of P. (Listrochehis Blanchard). 

The characters and terms used in the de- 
scriptions are those of Chapin (1935), Say- 
lor (1940), and Moron ( 1986, 1992). Draw- 
ings were made with the aid of a camera 
lucida and Zeiss stereomicroscope: mea- 
surements were obtained with an ocular mi- 
crometer or caliper 

Acronyms used in the text are as follows; 
CAS, California Academy of Sciences, San 
Francisco: CNC, Canadian National Collec- 
tion, Ottawa; HAHC, Henry and Anne 
Howden, Canadian Museum of Nature, Ot- 
tawa; IBUNAM, Institute de Biologi'a, 
UNAM. Mexico City; lEXA, Instituto de 
Ecologia, Xalapa, Mexico; MHNM, Museo 
de Historia Natural, Mexico City; MXAL, 
private collection M.A. Moron, Xalapa, 
Mexico; UNSM, University of Nebraska 
State Museum, Lincoln. 

Phyllophaga (Chlaenobia) rzedowskiana 
Aragon and Moron, new .species 

(Figs. 1-11) 

Descriptit)n. — Holotype male: Clypeus 
and frons yellowish brown; pronotum 
shiny yellowish; elytron straw yellowish 
without macroscopic vestiture, glabrous; 
mouthparts, sterna, pygidium and legs 
shiny straw yellowish. Clypeus 2.5X wid- 
er than long, anterior border slightly sin- 
uate, with elevated margin (Fig. 1), disk 
surface glabrous, slightly concave at 
sides, with many uniformly distributed, 
deep, rounded punctures. Frontoclypeal 
suture slightly sinuate and deeply im- 
pressed. Frons 2. IX wider than long, gla- 

brous, convex, regularly and deeply punc- 
tate. Vertex without punctures. Antenna 9- 
segmented (Fig. 1), with 3-segmented 
club, lamellae 1.5x longer than length of 
preceeding five segments combined. 
Frons 3.1 X wider than dorsal diameter of 
each eye. Eye canthus long and narrow, 
with 1 1-12 setae. Labrum bilobed, deeply 
sinuate, with scattered long, slender setae 
on borders. Mentum slightly concave, im- 
punctate, with scarce lateral setae, ante- 
rior border slightly sinuate. Pronotum 
1 .6X wider than long and 2.5 X wider than 
frons. Pronotal disk shiny, with round, 
moderately deep punctures, irregularly 
distributed; lateral borders widely angu- 
lated, lateral marginal bead with scattered, 
short, slender setae; basal bead strongly 
indicated, inclusive in front of scutellum 
where indicated by deep punctures; ante- 
rior angles obtuse, not prominent; poste- 
rior angles not directed downward, ob- 
tuse. Scutellum 1.2X wider than long, 
with small punctures irregularly distrib- 
uted. Elytron 2.7 X longer than wide, 
shiny, glabrous, densely rugo-punctate; 
punctures small, shallow, separated by 1- 
2 diameters; epipleural border progres- 
sively narrowed along complete margin, 
with a nearly homogeneous fringe of slen- 
der setae; humeral callus rounded, prom- 
inent; apical callus rounded. Metathoracic 
wings completely developed. Propygi- 
dium slightly shiny, yellowish with shal- 
low punctuation and scattered short setae. 
Pygidium convex, mainly towards basal 
border (Fig. 9), shiny, densely punctate, 
with many short, erect setae on disk; api- 
cal margin with 8 long, slender setae; bas- 
al margin incomplete at middle. Pteroster- 
num with moderately dense, long yellow- 
ish setae. Visible abdominal sternites II to 

IV of similar length, slightly flattened, 
with dense, short setae at middle; sternite 

V flattened, slightly longer than precee- 
ding sternite, with short setae at middle 
and one row of long setae toward poste- 
rior border. Anal plate nearly as long as 
preceeding sternite, convex, slightly fur- 


Figs. 1-11. PliyllophaMii (Chlaenohia) rzedowskiana. 1, Male head and pronotum, dorsal view. 2, Male 
protarsal claw, lateral view. 3, Female protarsal claw, lateral view. 4. Parameres. distal view. 5, Apex of aedeagus. 
ventral view. 6, Genital capsule, right lateral view. 7. Genital capsule, dorsal view. 8, Apex of aedeagus, left 
lateral view. 9, Male abdomen, lateral view. 10, Female abdomen, lateral view. 1 1, Female genital plates, ventral 
view. Scale lines = I mm, except Figs. 2-3 = 0.3 mm. 

rowed at midline, with scattered, long se- 
tae, posterior margin poorly indicated at 
sides. Protibia shorter than protarsus 
(0.53:1), with external border tridentate, 
proximal tooth much shortened, preapical 
spur narrow, nearly straight, apex acute, 
half as long as 2nd protarsomere. Meso- 
tibia with one oblic|ue, sharp, setiferous 

carina on external side; upper apical spur 
straight, narrow, and l.lx longer than 
lower spur. Metatibia shorter than meta- 
tarsus (0.75:1), with one oblique, sharp, 
setiferous carina on external side; upper 
apical spur lanceolate, slightly curved, 
apex acute, nearly as long as basal meta- 
tarsomere, and 1.3x lc)nger than lower 


spur; lower apical spur articulated with 
tibial border, with acute apex. Tarsomeres 
nearly cylindrical, elongate, with enlarged 
apices, with some setae apically and two 
lines of thick setae on ventral side. Pro- 
tarsal claws narrowly cleft (Fig. 2), with 
lower tooth shorter than upper tooth; 
meso- and metatarsal claws with lower 
tooth slightly shorter than upper tooth. 
Genital capsule with short parameres 
(Figs. 4, 6-7), dorsally and ventrally 
fused, symmetrical, apex of each para- 
mere widely rounded and directed down- 
ward. Aedeagus with sclerotized, tubelike 
support, and three, angled, apical projec- 
tions, one of them notably largest, with 
apex directed downward (Figs. 5-8). Tec- 
tum enlarged, widely convex. Length of 
genital capsule from apex of parameres to 
border of basal piece: 3.7 mm. Total body 
length: 12.0 mm. Humeral width: 5.0 mm. 
Allotype female: Similar to male except 
as follows: antenna with lamella of seg- 
ments 7-9 slightly shorter than the length 
of five preceeding segments combined. Vis- 
ible abdominal sternites II to V convex, 
with scattered setiferous punctures near 
middle; anal plate convex, with scattered 
setiferous punctures, and 6 slender setae at 
posterior border. Complete surface of py- 
gidium with many erect setae: basal half 
convex, distal half with deep concavity at 
middle, with 8 slender setae along apical 
border (Fig. 10). Protibia with teeth of ex- 
ternal border slightly longer than in male. 
Both apical spurs of metatibia articulated, 
wide, lanceolate and curved. Protarsus 
slightly longer than protibia (1.12:1). Tarsal 
claws similar on all legs; lower tooth slight- 
ly shorter than upper (Fig. 3). Ventral gen- 
ital plates convex, poorly sclerotized, nearly 
symmetrical, without setae, apical borders 
briefly sinuated; dorsal genital plates small, 
with long setae on distal border (Fig. 11). 
Total body length: 12.1 mm. Humeral 
width: 5.0 mm. 

Variation. — Male: Similar to holotype 
except as follows: sclerotized preapical pro- 
jections of aedeagus more or less narrower 

or longer. Total body length: 11.8-12.3 
mm; humeral width: 4.9-5.2 mm. Female: 
similar to allotype except head and prono- 
tum light reddish brown, disc and apical 
border of anal plate with more or less setae. 
Total body length: 12.0-12.5 mm; humeral 
width: 5.0-5.3 mm. 

Type material. — Described from 51 S. 
54 $ . Holotype 6 MXAL: Mexico: Puebla, 
Atencingo, Chietla, 1,145 m, 30-V-1999, A. 
Aragon. Allotype 9 MXAL: same data as 
holotype. Paratypes CAS, CNC, HAHC, 
UNSM: Mexico: Morelos, Emiliano Zapa- 
ta, IO-VI-78. E Cervantes (3 cJ ); Huautla, 
2.5 km N and 4 km W Estacion CEAMISH, 
940 m, 8/12-VI-I996, A. Perez (3 6 2 9); 
Huahutla, Estacion CEMISH, 2.5 km N 4 
km W, 940 m, 8-12-VI-1996, luz, A. Perez 
(11 (5, 5 9); Zacatepec, Galeana, 28-VI- 
1983, C. Deloya (6 (5 8 9); same data ex- 
cept 27-VI-1983 (3 cJ 6 9); 3-VII-1983 (1 
6 1 9); 4-VII-1983 (1 d ); Acamilpa, Tlal- 
tizapan, 27-VI-1983, C. Deloya (1 cJ); Jo- 
jutla, Cerro del Higueron, 4-VII-1982, C. 
Deloya (1 c?); Jojutla, 9-Vin-1983, C. De- 
loya (1 S 1 9); 26-VL1983 (Id); Jojutla, 
Unidad Morelos, VI- 1988, C. and G. De- 
loya (18 (5 30 9). 

Type locality. — Chietla, Atencingo mu- 
nicipality. State of Puebla, Mexico (approx. 
18°3rN; 98°35'W). 

Distribution. — Southwestern Puebla, 
eastern and southeastern Morelos, Mexico 
(Fig. 12). 

Biological Data. — Males and females of 
P. rzedowskiana were collected at white 
fluorescent lights and Hg lights near rem- 
nants of tropical deciduous forests, that in- 
clude species of Bursera. PseiidosmoiUn- 
giuDi. Amphipteiygium. Lysiloma, Ceiba, 
Acacia. Ipomoea. Lemaireocereus, Cepha- 
locereiis. and Pachycereiis (Rzedowski 
1978), surrounded by sugarcane planta- 
tions, at 940-1,145 m altitude. Phenology: 
May (2), June (97), July (4), August (2). 
Other species of Phyllophaga flying at the 
same time and place were P. (Chlaenohia) 
howdcniaiia Moron, P. (Pliyllopluiga) hre- 





Fig. 12. Distribution of Phyl/opluii^a (C.) i:ed(m\kiaiui (•) and P. {L) harrciaini (A) in the Upper Basin 
of Balsas River. Mexico. Doited lines represent the approximate flow of the Balsas River and its main tributaries. 
Map adapted from Garcia and Falcon ( 1974). 

videus (Bates). P. (Phyllopluiiici) illuiicci- 
iiiinai Moron, and P. {Listroclwlus) barre- 
raiui, n.sp. 

Remarks. — Phyllophaga ( Chlaeiwbia ) 
rzedowskiatui belongs to the species group 
"vexatci" {sensit Moron 1986). By body 
shape, size, color, general vestitiire, punc- 
tuation on the clypeus. pronotum and elytra. 
as well as the general shape of the male 
genital capsule, it is similar to P. (Chlae- 
nobia) howdeniana Moron. But the short 
parameres with rounded apex, the three ir- 
regular projections at the apex of the scler- 
otized support of aedeagus (see Moron 
1992. figs. 8-10) and the shallow, preapical 
depression in the pigydium of the female, 
with small, rounded prominences at each 
side separate P. rz.edowskiaiia from P. how- 
den iana. 

Etymology. — This new species is dedi- 
cated to Dr. Jerzy Rzedowski. a well-known 
Mexican botanist, whose pioneering studies 

on the xeric vegetation and deciduous trop- 
ical forests from Mexico are continuous ref- 
erences to scientists and students. 

Phyllophaga {Listrochelus) barrerana 
Aragon and Moron, new species 

(Figs. 13-22) 

Description. — Holotype imile: Clypeus 
and frons reddish brown; pronotum shiny 
reddish yellow; elytron straw yellowish 
without macroscopic vestiture, glabrous; 
mouthparts, sterna, pygidium and legs 
shiny reddish yellow. Clypeus 2.8 X wider 
than long, anterior border brief and widely 
sinuate, with elevated margin (Fig. 13), disk 
surface glabrous, slightly concave at sides, 
with uniformly distributed, deep, rounded 
punctures. Frontoclypeal suture slightly sin- 
uate and finely impressed. Frons 2.7 X wid- 
er than long, glabrous, nearly flattened, ir- 
regularly and deeply punctate; transverse 
carina of vertex strongly indicated; surface 


3-— ~J*i*MJ^ 


,_i ,-»AM»A^sJ^ 




[ /^^ 


Figs. 13-22. Phyllophcif-a (Listrochelus) barrenma. L3. Male head and pronotum, dorsal view. 14, Male 
pmtarsal claw, lateral view. 15. Female protarsal claw, lateral view. 16, Parameres. distal view. 17. Genital 
capsule, right lateral view. 18. Genital capsule, dorsal view. 1'). .^pex of aedeagus. ventral view. 20. Male 
abdomen, lateral view. 21, Female abdomen, lateral \icw. 22. Female jicnital plates, \entral view. Scale lines 
= 1 mm, except fig. 14 = 0.5 mm. 

behind carina with scattered, small punc- 
tures. Antenna 10-segmented (Fig. 13). 
with 3-segmented club. lamellae 1.4X lon- 
ger than length of preceeding six segments 
combined. Frons 2.5 X wider than dorsal di- 
ameter of each eye. Eye canthus long and 
naiTow. with 12 setae. Labrum bilobed. 

widely sinuate, with scattered long, slender 
setae on borders. Mentum slightly concave, 
impunctate. with scarce lateral setae, ante- 
rior border briefly sinuate. Pronotum 1.5X 
wider than long and 2.6X wider than frons. 
Pronotal disk shiny, with round, shallow 
punctures, iiregularly distributed, leaving 


an impunctate mesial strip on anterior half; 
lateral borders widely angulated, lateral 
marginal bead crenulate. with slender setae; 
basal bead strongly indicated, inclusive in 
front of scutellum where it is indicated by 
deep punctures; anterior angles obtuse, 
slightly prominent; posterior angles obtuse, 
widely rounded. Scutellum 1.2X wider than 
long, with small punctures apically. Elytron 
2.4 X longer than wide, shiny, glabrous, 
rugo-punctate; punctures small, shallow, 
separated by 2-4 diameters; epipleural bor- 
der progressively narrowed along complete 
margin, with a fringe of slender setae; hu- 
meral callus rounded, prominent; apical cal- 
lus rounded. Metathoracic wings complete- 
ly developed. Propygidium slightly shiny, 
yellowish with shallow and scattered seti- 
ferous punctures. Pygidium uniformly con- 
vex (Fig. 16), shiny, with scattered, shallow 
punctures and short setae on disk; apical 
margin with 10 slender setae; basal margin 
incomplete at middle. Pterosternum with 
dense, long yellowish setae. Visible abdom- 
inal sternites II to IV of similar length, 
slightly flattened, with scattered, short setae 
near middle; sternite V flattened, slightly 
longer than preceeding sternite. with scarce, 
short setae, shallowly funowed at midline 
of posterior half, and with posterior border 
briefly notched. Anal plate much narrowed 
(Fig. 16), with scattered, shallow setiferous 
punctures, briefly furrowed at midline. Pro- 
tibia shorter than protarsus (0.5 1:1). with 
external border tridentate. proximal tooth 
much shortened, preapical spur narrow, 
nearly straight, apex acute, 0.6 X as long as 
2nd protarsomere. Mesotibia with one 
oblique, sharp, setiferous carina on external 
side; upper apical spur straight, narrow, as 
long as lower spur Metatibia shorter than 
metatarsus (0.8:1). with one oblique, sharp, 
setiferous carina on external side; upper 
apical spur sinuate, apex acute, 1.4X longer 
than basal metatarsomere, and 1.2X longer 
than lower spur; lower apical spur articu- 
lated with tibial border, with acute apex. 
Tarsomeres nearly cylindrical, elongate, 
with enlarged apices, with some setae sub- 

apically. and scattered setae on ventral side. 
All tarsal claws irregularly unipectinate 
(Fig. 14). Genital capsule with short para- 
meres (Figs. 15, 18-19), dorsally fused, 
symmetrical, with strong carinae along dor- 
so-distal border and lateral border, apex of 
each paramere widely rounded and directed 
downward. Tectum with shallow, wide sul- 
cus along midline and rounded prominen- 
ces basally (Fig. 18). Aedeagus with scler- 
otized, tubelike support, with dorsal pre- 
apical projection, and one subapical, slight- 
ly asymmetrical sclerite (Figs. 17-19). 
Length of genital capsule from apex of pa- 
rameres to border of basal piece: 3.8 mm. 
Total body length: 11.2 mm. Humeral 
width: 5.0 mm. 

Allotype female: Similar to male except 
as follows: antenna with lamella of seg- 
ments 8-10 0.8X longer than length of pre- 
ceeding seven segments combined. Anal 
plate large and convex, with short setae. Py- 
gidium moderately convex with scattered 
setiferous punctures (Fig. 21). Tarsal claws 
with acute tooth before the middle of ven- 
tral border, and posterior border unipecti- 
nate (Fig. 15). Ventral genital plates with 
posterior border narrowed, with rounded 
apex, glabrous. Dorsal genital plates with 
apical border widely curved, briefly 
notched, with erect setae at apex (Fig. 22). 
Total body length: 13.2 mm. Humeral 
width: 5.8 mm. 

Variation. — Male specimen from Ixtapan 
de la Sal similar to holotype except as fol- 
lows: pronotai and elytral punctures deeper; 
carinae of the parameres more elevated. To- 
tal body length in both sexes: 13.0-13.5 
mm; humeral width in both sexes: 5.6-5.9 

Type material. — Described from 24 (5 3 
9. Holotype 6 MXAL: Mexico: Puebla, 
Tilapa, Casa Blanca. 1,210 m, 30-V-1999, 
A. Aragon. Allotype 9 MXAL: Mexico: 
Morelos, Acamilpa, Tlaltizapan, 27-VII- 
1983, C. Deloya. Paratypes CAS, CNC, 
MHNM, UNSM: same data as holotype (2 
6 ); Mexico: Puebla: Chietla. Atencingo, 



1.145 m, 30-V-1999, A. Aragon (2 6): Es- 
tado de Mexico, Ixtapan de la Sal. 29- V- 
1971. A. Bairera (1 6): Mexico: Morelos: 
Jojutla. Unidad Morelos. VM988. C. and 
G. Deloya (1 6); Jojutla. 9-VII1-1983. C. 
Deloya (1 6): Zacatepec. Galeana. 28-VI- 
1983. C. Deloya (5 6): same data except 
27-VI-1983 (Id): Acamilpa, Tlaltizapan. 
27-VII-1983, C. Deloya (10 d 2 9). 

Type locality. — Tilapa. Atencingo mu- 
nicipality, state of Puebla. Mexico (aprox. 
18°34'N: 98°32'W). 

Distribution. — Southwestern Puebla, 
southern Mexico and southeastern Morelos. 
Mexico (Fig. 12). 

Biological Data. — Males of P. banenma 
were collected at white fluorescent lights 
near remnants of tropical deciduous forests, 
that include species of Biirsera, Pseiidos- 
modingiiim. Ampbipterygium. Lysi Ionia, 
Ceiha, Acacia. Ipowoea, Lemaireocereiis. 
Cephalocereus. and Pachycereus (Rze- 
dowski 1978). suiTounded by sugarcane 
plantations, or avocado and mango planta- 
tions, at 1.145-1,210 m altitude. Phenolo- 
gy: May (6). June (7). July (13), August { 1 ). 
Other species of PbyUopbaga flying at the 
same time and place were P. (Cblaenobia) 
bowdeniana Moron. P. (Cblaenobia) rze- 
dowskiana Aragon and Moron. P. (PbyUop- 
baga) brevidens (Bates), and P. (PbyUop- 
baga) illniicaminai Moron. 

Remarks. — PbyUopbaga (Litrocbelus) 
harrenma belongs to the species group 
"cavata" (sensu Moron 1986). By body 
shape and size, punctuation on the clypeus. 
pronotum and elytra, structure of pygidium. 
as well as the general shape of the male 
genital capsule, it is similar to P. (Listro- 
cbeliis) valia Saylor and P. (L.) cochisa 
Saylor. But the pruinose elytra with basal, 
erect setae; pruinose and setiferous pygidi- 
um; parameres with acute apices curved in- 
ward; and wide sclerotized support of the 
aedeagus with the apex abruptly narrowed, 
separate P. valia from P. barrerana. The 
male antennal club as long as the preceed- 
ing six segments combined; non-furrowed 
anal plate; parameres with enlarged, nearly 

parallel, acute apices; and narrow sclero- 
tized support of aedeagus without apical re- 
duction, separate P. cocbisa from P. barre- 

Etymology. — This new species is dedi- 
cated to the late Dr. Alfredo Barrera (1926- 
1980). well known Mexican entomologist, 
ethnobiologist. teacher and promoter of the 
biological sciences, whose publications on 
the taxonomy of Siphonaptera. scientific 
collections, natural history museums, ecol- 
ogy of tropical vegetation, and Mayan eth- 
nobotanical nomenclature, are classical ref- 


We are indebted to Alejandro Perez 
Garcia and Maria Eugenia Di'az (Mexico 
City) for the loan or donation of some spec- 
imens here described. Field trips in the state 
of Puebla were conduced with the support 
of project FB626/R024/98. CONABIO, 
Mexico-BUAP. This paper is a contribution 
to the project "'Sistematica y Biologi'a del 
genero PbyUopbaga en America Latina" 
partially supported by Instituto de Ecologia, 
A.C. Xalapa (account 902-02). 

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103(3). 2003. pp. 568-577 




Gu-STAVO R. Spinelli and William L. Grogan, Jr. 

(GRS) Departamento Cienti'fico de Entomologi'a. Miiseo de La Plata. Paseo del Bosque 
s/n. 1900 La Plata. Argentina (email:; (WLG) De- 
partment of Biological Science.s, Salisbury University, Salisbury. MD 21801, U.S.A. 

Abstract. — This revision of the Neotropical biting midges of the genus Purodasyhelea 
Ingram and Macfie recognizes three species of this rare genus in southern South America 
and includes a key for the recognition of both sexes of all Neotropical species. In addition 
to the previously described P. hrevipalpis (Ingram and Macfie), which is redescribed and 
ilkistratcd. two new .species are described and illustrated: P. ingrami. from Argentina, and 
P. inacfiei, from Argentina and Chile. 

Resiiinen. — En esta revision de las especies patagonicas del curioso genero Paradasy- 
lu'lea Ingram & Macfie. se reconocen tres especies para America del Sur, incluyendose 
asimismo una clave identificatoria de todas las especies Neotropicales, para ambos sexos. 
Ademas de la especie previamente descripta P. hrevipalpis (Ingram y Macfie), la cual es 
redescripta e ilustrada, se describen e ilustran dos especies nuevas: P. ingrami de la 
Argentina, y P. macfiei de la Argentina y Chile. 

Ki'\ Words: 

Diptera, Ceratopogonidae. Paradasyhclcii. biting midges, new species. Neo- 

A single, very unusual biting midge that 
was collected by Frank W. Edwards during 
an expedition to Patagonia in 1926 was 
subsequently described as Dasyhelea hre- 
vipalpis by Ingram and Macfie (19.^1). 
However, Ingram and Macfie noted that 
"This species falls into that group of insects 
still included in the genus Dasyhelea Kief- 
fer in which the antennal segments are not 
sculptured, and segments 12-14 in the male 
are not binodose. It differs moreover from 
a typical species in the genus in having 
small humeral pits, a rather long petiole to 
the median fork, a fringe on the alula, a T. 
R. (tarsal ratio of hind leg) less than 2. and 
in the peculiar form oi the palp."" Perhaps 

because this species was otherwise so dif- 
ferent from other species of Dasyhelea. 
Macfie ( 1940) designated it as the type spe- 
cies of the new genus Paradasyhelea, 
which he included in his ''Dasyhelea 
Group" in his classic paper '"The genera of 
Ceratopogon idae. " ' 

Subsequently, Wirth and Lee (1959) as- 
signed Dasyhelea egre^ia Macfie (1932) 
from New Zealand to Paradasyhelea Mac- 
fie when they described two new species in 
this genus from eastern Australia. They not- 
ed that species of Paradasyhelea were 
unique among three apparently related gen- 
era in having both radial cells obsolete, a 
short costa extendine less than (15 of wing 



length and otherwise resembling these three 
genera as follows: 1 ) Forcipomyia Meigen. 
because of its broadly separated eyes, 
oblique r-m crossvein, hind tarsal ratio less 
ilian 2.0. the presence of slender hyaline 
sensilla on the flagellum. and the shape of 
the male gonostylus and parameres; 2) 
Dasyhelea. because of its 4-seginented pal- 
pus, pubescent eyes, short proboscis with 
vestigial mandibles, and the shape of the 
apicolateral processes of male genitalia; and 
3) Ciilicoides Latreille, by the presence of 
humeral pits and sensory pits (sensilla coe- 
loconica) on some flagellomeres and ter- 
minal fiagellomere without a terminal pa- 
pilla. Wirth and Blanton (1969) essentially 
reiterated what Wirth and Lee had said 
about the relationships of these four genera 
but noted that "Panulasyhelea seems to be 
a relict ceratopogonid of an extremely an- 
nectant type with characters cutting across 
three subfamily lines." 

Wirth et al. (1974) were the first workers 
to include Paradasyhelea in the tribe Cu- 
licoidini of the subfamily Ceratopogoninae 
in their list and key of world genera. They 
noted that "The genera placed in the tribe 
Culicoidini appear to be as primitive and 
non-specialized as any in the family . . . We 
believe it is no accident that the annectant 
genera Paradasyhelea and Austrocoiiops 
Wirth and Lee from Australia, New Zea- 
land, and Patagonia fall here." Wirth and 
Lee ( 1959) described the pupae of two spe- 
cies of Paradasyhelea. however. Kettle and 
Elson (1975) first noted the nearly identical 
nature of pupae in that genus to those of 
Ciilicoides. Furthermore, when Kettle and 
Elson described and illustrated the larvae of 
P. minuta they noted ". . . it is clear that 
larvae of P. minuta cannot be separated 
from those of Ciilicoides. with which they 
share (1) a head ratio of about 1.5: (2) in- 
conspicuous antennae; (3) an absence of 
multiple setae on the head; (4) a poorly de- 
veloped ventral suture on the head: and (5) 
with most Culicoides. an absence of long 
perianal bristles." Subsequently, when El- 
son-HaiTis and Kettle (1985) described the 

adults and immature stages of their new 
Australian species P. reyei. they noted 
"The immatures of P. reyei confirm this 
close similarity between Paradasyhelea and 
Ciilicoides.'' Elson-Harris and Kettle also 
listed several features of the immature stag- 
es of Parada.syhelea that differed from 
those in Culicoides. Perhaps a study in pro- 
gress by Borkent (personal communication) 
on all available immature stages of Cera- 
topogonidae will help resolve questions of 
intergeneric relationships between these 
two genera. 

Using modern phylogenetic methods in- 
volving detailed outgroup comparisons to 
determine polarity of character states, Bor- 
kent (1995) implied a sister group relation- 
ship between Culicoides and Paradasyhe- 
lea when he noted "The presence of a num- 
ber of flagellomeres with sensilla coeloco- 
nica in at least some members of this genus 
may indicate that the derived condition is 
actually a synapomorphy of Culicoides + 
Parada.syhelea." Using the same phyloge- 
netic methods. Borkent and Grogan (1995) 
determined, based on the absence of pali- 
sade setae on tarsomere 1 of the hind leg in 
adults, that "the earliest lineage of the Ce- 
ratopogoninae are Culicoides. Parada.syhe- 
lea Macfie, and Washingtonhelea Wirth and 
Grogan." Finally, while the monophyly of 
Paradasyhelea has not yet been firmly es- 
tablished, it is the only genus in the tribe 
Culicoidini in which the adult females of 
all species have reduced, vestigial mandi- 
bles, thereby, at least making them unique 
within the tribe. While some species of Cu- 
licoides are autogenous (non-feeding) with 
reduced, vestigial mandibles, this is almost 
certainly a derived condition for these spe- 
cies in that genus and is not evidence of a 
sister group relationship of these two gen- 

Currently, there are nine known species 
of Paradasyhelea, eight of which are from 
the southern hemisphere (Borkent and 
Wirth 1997): P. albi/niiictata Wirth and Lee 
(1959), from New South Wales, Australia; 
P. houcheti Clastrier (1989). from New 


Caledonia; P. brevipalpis Ingram and Mac- 
fie (1931), from Argentina; P. egregia 
(Macfie) (1932). from New Zealand; P. 
horrisoni Wirth (1981). from New Zealand; 
P. miniiia Wirth and Lee ( 1959). from New 
South Wales and Queensland. Australia; P. 
iieocaledouiensis Clastrier (1989), from 
New Caledonia; and P. reyei Elson-Harris 
and Kettle (1985), from Queensland. Aus- 
tralia. Only one species, P. olympiae Wirth 
and Blanton (1969), is known from the 
northern hemisphere, but only from the 
original specimens that were collected on 
the Olympic Peninsula in Olympic National 
Park, Washington. USA. by Willis Wirth in 
1968. However. Wirth and Blanton were 
not entirely confident of the generic assign- 
ment of P. olympiae and noted when they 
compared it with the then four other known 
species of Panuiasyhelea that it "differs 
from all of them in the shape of the api- 
colateral processes of the ninth tergum. the 
aedeagus. and the median sclerite of the pa- 

A recent collecting trip to Patagonia in 
Argentina and Chile by GRS produced 
some specimens of Paradasyhelea that dif- 
fered from those of P. brevipalpis. Art Bor- 
kent kindly lent us additional specimens of 
Paradasyhelea in the Canadian National 
Collection of Insects (CNCI) in Ottawa. Ca- 
nada. In addition. GRS recently re-exam- 
ined the holotype of P. brevipalpis in The 
Natural History Museum (BMNH) in Lon- 
don. United Kingdom. A careful examina- 
tion of these specimens revealed two un- 
described species that we describe and il- 
lustrate herein, we also redescribe and il- 
lustrate P. brevipalpis and provide a key for 
the recognition of all Neotropical species. 
Clastrier ( 1989) provided a key to the world 
species of Paradasyhelea. but. inexplicably, 
failed to include P. egregia. 

Unless otherwise noted, all recently ac- 
quired specimens were mounted on micro- 
scope slides in Canada balsam in the man- 
ner of Wirth and Marston ( 1968). Types are 
deposited in the Museo de La Plata. Argen- 
tina (MLPA) or in the CNCI. as indicated. 

General terms follow those in Downes and 
Wirth (1981). We dedicate this article to 
our recently departed colleague. Dr Larry 
Quate, in recognition of his important con- 
tributions to the study of primitive Diptera. 

Key to Neotropical Species 
OF Paradasyhelea 

1 . Female 2 

- Male 4 

2. Palpus 5-segmented. third segment with capi- 
tate sensilla enclosed in a rounded pit (Fig. 14); 
spermatheca small, ovoid 

Paradasyhelea macfiei, n. sp. 

- Palpus 4 or 5-seamented. capitate sensilla not 
enclosed in a pit (Figs. 2, 8): spermatheca 
large, retort-shaped 3 

.^. Palpus 5-segmented; flagellomere 9 of female 
unusually short and narrow, shorter than 10 
(Fig. 7) Paradasyhelea ingraini. n. sp. 

- Palpus 4-segmented: flagellomere 9 of female 
longer than 10 (Figs. L 13) 

Paradasyhelea brevipalpis (Ingram and Macfie) 
4. Gonostylus abruptly narrowed at midlength 
(Fig. 18). nearly straight to blunt tip; distal por- 
tion of aedeagus tapering to pointed tip (Fig. 
18): parameres with inverted U-shaped basal 
sclerite and a pair of posteriorly projecting pro- 
cesses (Fig. 19) Paradasyhelea macfiei. n. sp. 

- Gonostylus stout, flask-shaped (Fig. 6) or pro- 
gressively tapering to almost pointed tip (Fig. 
12); distal portion of aedeagus with knob-like 
tip (Figs. 6. 12); parameres reduced to a stout 
m\erted U-shaped sclerite (Figs. 6. 12) 5 

3. Gonostylus progressively tapering to almost 
pointed tip; apicolateral process of male geni- 
talia stout (Fig. 12) 

Paradasyhelea ingrami. n. sp. 

- Gonostylus flask-shaped; apicolateral process 
of male genitalia long and slender (Fig. 6) . . 

Paradasyhelea brevipalpis (Ingram and Macfie) 

Paradasyhelea brevipalpis (Ingram and 


(Figs. 1-6) 

Dasyhelea brevipalpis Ingram and Macfie 
193 1: 178 (6\ Argentina, Lake Nahuel 

Paradasyhelea brevipalpis: Macfie 1940: 
17 (comb.); Wirth 1981: 386 (in key); 
Spinelli 1987: 667 (9; Argentina. Neu- 
quen and Rio Negro provinces); Spinelli 
and Grogan 1999: 709 (Argentina. Tierra 
del Fueso). 


Figs. 1-6. Faradasyhelca hrevipulpis. I, FlagelUim of lemaic. 2, Palpus of lenialc. 3. Wing iif female. 4, 
Spermatheca. 5, Flagellomeres 1 1-13 of male. 6, Genitalia of male. Scales = O.O.I mm. lev = Intercalary vein; 
M, = Vein M,; M, = Vein M,; r, = Cell r,. 

Diagnoses. — Only Neotropical species of 
Paradasyheleu with a 4-segmented palpus. 
Female with flagellomere 9 longer than 10; 
capitate sensilla of palpus not enclosed in a 
pit; scutellum with 9 stout setae; vein M, 
present or absent; costal ratio 0.41-0.43; 
spermatheca large, retort-shaped; halter 
brownish. Male with apicolaleral process 

long and slender; gonostylus flask-shaped; 
distal portion of aedeagus with knoblike tip; 
parameres reduced to a stout inverted U- 
shaped sclerite. 

Female. — Head: Dark brown. Eyes pu- 
bescent, separated by breadth of 2 omma- 
tidia. Antennal flagellum (Fig. 1 ) brown; 
flagellomere 9 longer than 10; sensilla coe- 


loconica on flagellomeres 1, 6—8, or rarely 
9; scape pale, with 9-1 1 setae; antennal ra- 
tio 0.70 (0.65-0.75, n = 6). Palpus (Fig. 2) 
4-segmented, pale; segment 2 (fused prim- 
itive 2nd & 3rd) with scattered capitate sen- 
silla on surface of distal third. Thorax: Uni- 
formly brown. Scutum with sparse vestiture 
of setae; scutellum with 9 stout setae in row 
plus 20 smaller setae. Legs brownish in- 
cluding tarsi, femorotibial joints darker; 
hind tibial comb with 4 bristles; hind tarsal 
ratio 1.65 (1.52-1.82, n = 6); claws short, 
slightly curved. Wing (Fig. 3) densely cov- 
ered with mactrotrichia (as determined by 
their insertions, not illustrated); radial cells 
obliterated; vein M, nearly imperceptible or 
absent; a conspicuous, forked intercalary 
vein present in cell r,; wing length 1.20 
(1.10-1.32, n = 6) mm, breadth 0.50 (0.47- 
0.55, n = 6) mm; costal ratio 0.42 (0.41- 
0.43). Halter brownish. Abdomen: Pale 
brown. One retort-shaped, very heavily 
sclerotized spermatheca (Fig. 4) with slen- 
der oblique neck, measuring 0.070 by 0.061 
mm, neck 0.008 mm. 

Male. — Similar to female with usual sex- 
ual differences. Lengths of distal three fla- 
gellomeres (Fig. 5) in |xm 97-56-52. Hind 
tarsal ratio 1.47 (1.42-1.60, n = 5). Wing 
length 1.10(1.08-1.17, n = 5) mm; breadth 
0.36 (0.34-0.37, n = 5) mm; costal ratio 
0.45 (0.44-0.46, n = 5). Genitalia (Fig. 6): 
Sternite 9 with moderately deep, broad cau- 
domedian excavation; tergite 9 moderately 
long, distal margin nearly straight; apico- 
lateral process very slender, each with a mi- 
nute apical seta; cercus elongated, tip trun- 
cate. Gonocoxite short, twice as long as 
broad with poorly developed mesobasal tu- 
bercle; gonostylus stout, flask-shaped, as 
long as gonocoxite, tapering abruptly on 
distal half, tip pointed, beaklike. Aedeagus 
Y-shaped; basal arms heavily sclerotized, 
subparallel; distal portion nearly straight 
with slightly expanded, knoblike tip. Para- 
meres reduced to stout, inverted U-shaped 
basal sclerite; basal apodemes curved, 
heavily sclerotized. 

Distribution. — Argentina, in subantarctic 

Nothofagiis forests, from 40°S south to Ti- 
erra del Fuego. 

Type. — Holotype male, Argentina, Rio 
Negro Province, eastern end of Lake Na- 
huel Huapi (BMNH). Examined during the 
present study. 

Other specimens examined. — Argentina, 
Neuquen, 10 km N San Martin de Los An- 
des, 24-XI-1984, G. Spinelli, 4 9, 1 cJ; Ne- 
uquen. 6-7 km S San Martin de Los Andes 
(900 m). 26-XM986. G. Spinelli. 19,3 
6 ; Neuquen, Lanin National Park, Lake 
Queni. 6/8-n-1999, P Marino. 1 6: Nahuel 
Huapi National Park. Chall-huaco. 6-XII- 
1992, G. Spinelli, 1 9, CDC light trap; Na- 
huel Huapi National Park, Lake Escondido, 
29-XI- 1984, G. Spinelli, 1 9 ; same data ex- 
cept 3-XII-1988, D. Afion Suarez, 2 6 
CDC light trap; Chubut, Los Alerces Na- 
tional Park, lake Futalaufquen. 22-11-1994, 
G. Spinelli, 1 9, sweep net; Chubut. Los 
Alerces National Park. Puerto Mermoud, 
23-n-1994. G. Spinelli. 1 9. 1 cJ, sweep 
net; Santa Cruz, Ri'o Turbio, mina I, 14-L 
1992. A. Estevez-J. Muzon, 2 9, 4 c5, 
sweep net; Tierra del Fuego, lake Escon- 
dido (140 m), 2-III-1993, G. Spinelli, 1 9; 
Tierra del Fuego National Park, Lapataia, 
9/10-1-1995. G. Spinelli. 6 9. CDC light 

Paradasyhelea ingrami Spinelli and 
Grogan, new species 

(Figs. 7-12) 

Diagnosis. — Only species of Paradasy- 
helea with female flagellomere 9 unusually 
short; palpus 5-segmented. capitate sensilla 
on segment 3 not enclosed in a pit; scutel- 
lum with 9 stout setae; vein M, absent; cos- 
tal ratio 0.35; halter brownish; spermatheca 
large, retort-shaped; apicolateral process of 
male genitalia stout; gonostylus progres- 
sively tapering; distal portion of aedeagus 
with knoblike tip; parameres reduced to a 
stout inverted U-shaped basal sclerite. 

Female. — Head: Dark brown. Eyes pu- 
bescent, separated by breadth of 2 omma- 
tidia. Antennal flagellum (Fig. 7) brown; 
flagellomere 9 unusually shorter, nanower 


Figs. 7-12. Parcidasxheleci ingniiiii. 7. Flagellum of female. 8. Palpus of female. 9. Wing of female. 10, 
Spermatheca. 1 1. Flagellomeres 1 1-1,^ of male. 12. Genitalia of male. Scales = 0.03 mm. 

than those immediately proximad, distad; 
.sensilla coeloconica on flagellomeres 1. 6- 
8; scape with 7 setae; antennal ratio 0.74. 
Palpus (Fig. 8) 5-segmented, slightly paler 
than flagellum; segment 3 with few scat- 
tered capitate sensilla on mesal surface, not 
enclosed in a pit. Thorax: Uniformly 
brown. Scutum with sparse vestiture of se- 
tae; scutellum with 9 stout setae in a row. 
and 9 slender anterior setae. Legs brownish 
including tarsi, femorotibial joints darker; 
hind tibial comb with 4 bristles; hind tarsal 
ratio 1 .60; claws short, slightly curved. 
Wing (Fig. 9) densely covered with mactro- 
trichia (as determined by their insertions, 
not illustrated); radial cells obliterated; vein 
M, absent; a conspicuous, forked interca- 
lary vein present in cell r^; wing length 0.85 
mm. breadth 0.34 mm; costa very short. 

costal ratio 0.35. Halter brownish. Abdo- 
men: Pale brown. One retort-shaped, very 
heavily sclerotized spermatheca (Fig. 10) 
with slender, curved, oblique neck, measur- 
ing 0.070 by 0.050 mm, neck 0.015 mm. 

Male. — Similar to female with usual sex- 
ual differences. Lengths of distal three fla- 
gellomeres (Fig. 11) in |xm 68-34-44. Hind 
tarsal ratio 1.60 (1.50-1.75. n = 5). Wing 
length 0.82 (0.74-0.88. n = 5) mm; breadth 
0.28 (0.26-0.30. n = 5) mm; costal ratio 
0.37 (0.36-0.38. n = 5). Genitalia (Fig. 12): 
Sternite 9 with deep, broad caudomedian 
excavation; tergite 9 large, distal margin 
nearly straight; apicolateral process stout, 
each with a minute apical seta; cercus con- 
spicuous, tip bluntly rounded. Gonocoxite 
short, twice as long as broad with large 
blunt mesobasal tubercle; gonostylus stout. 


as long as gonocoxite. distal portion mod- 
erately curved, tapering distally to slender 
weakly pointed tip. Aedeagus Y-shaped; 
basal arch heavily sclerotized; distal portion 
nearly straight with slightly expanded, 
knob-like tip. Paranieres reduced to a stout, 
inverted U-shaped basal sclerite. basal apo- 
demes curved, heavily sclerotized. 

Distribution. — Argentina (Chubut and 
Rio Negro provinces). 

Types. — Hoiotype <J, Argentina, Rio Ne- 
gro, Somuncura plateau, Chipauquil, 27- 
XI-1995, G. Spinelli, sweep net (MLPA). 
Paratypes, I 9. 7 cJ (MLPA), as follows: 
Argentina, Rio Negro, Somuncura plateau, 
Estancia El Rincon, 29-XI-1995, G. Spi- 
nelli, 1 9 (allotype), sweep net; same data, 
1 6. CDC light trap; same data, 1 6. Mal- 
aise trap; Chubut, Sierra Cuadrada plateau, 
Estancia Don Eduardo, 1-2-XII-1996, G. 
Spinelli, 5 6. Malaise trap. 

Etymology. — We are pleased to name 
this new species in honor of the late Alex- 
ander Ingram in recognition of his pioneer- 
ing publication on Patagonian biting midg- 
es that he co-authored with J. W. S. Macfie 
(Ingram and Macfie 1931). 

Discussion. — This is the only Patagonian 
species inhabiting steppe areas, which are 
located as far as 350 km from the Notho- 
fagus forests. 

Paradasyhelea ingranii resembles P. hre- 
vipalpis by virtue of the large, retort-shaped 
spermatheca, the distal portion of aedeagus 
with knoblike tip. and parameres reduced to 
a stout inverted U-shaped sclerite. Howev- 
er, it clearly differs from P. brevipalpis by 
its 5-segmented palpus, much shorter costa, 
flagellomere 9 unusually shorter and nar- 
rower than those immediately proximad and 
distad, the gonostylus progressively taper- 
ing to weakly pointed tip, and the stouter 
apicolateral process of the male genitalia. 
This new species keys to near couplets 5a 
(P. boucheti) and 5b (P. brevipalpis) in the 
world key by Clastrier (1989), but both of 
these species differ from P. ingraini in hav- 
ing 4-segmented palpi. 

Two females collected from a forested 

area in the western portion of the Argenti- 
nean province of Neuquen (15 km E Ca- 
viahue, 13-X1-1994, G. Spinelli, 2 females, 
sweep net. (MLPA) are very similar to this 
species, apparently differing only by their 
more broadly separated eyes (by width of 
4-5 ommatidia). Because of this difference 
in these two specimens and also because no 
males were found associated with them, we 
do not consider them as members of P. in- 
graini and, therefore, have not designated 
them as paratypes. 

Paradasyhelea macfiei Spinelli and 
Grogan, new species 

(Figs. 13-19) 

Diagnosis. — Only species of Paradasy- 
helea with palpus 5-segmented, segment 3 
with capitate sensilla enclosed in a rounded 
pit; vein M2 absent; halter whitish; sper- 
matheca small, ovoid; gonostylus abruptly 
narrowed at midlength. nearly straight to 
blunt tip; distal portion of aedeagus taper- 
ing to pointed tip; parameres with inverted 
U-shaped basal sclerite and pair of poste- 
riorly projecting processes. 

Female. — Head: Dark brown. Eyes pu- 
bescent, separated by breadth of 2 omma- 
tidia. Antennal flagellum (Fig. 13) brown; 
flagellomere 9 subequal to or slightly short- 
er than those immediately proximad. distad; 
sensilla coeloconica on flagellomere 1 very 
inconspicuous, these sensilla may be pre- 
sent or absent on flagellomeres 6-9; scape 
with 7 setae; antennal ratio 0.71 (0.68-0.75, 
n = 5). Palpus (Fig. 14) 5-segmented, 
slightly paler than flagellum; segment 3 
with a rounded, shallow sensory pit bearing 
capitate sensilla. Thorax: Uniformly brown. 
Scutum with sparse vestiture of setae; scu- 
tellum not in position to count number of 
setae. Legs brownish including tarsi, fe- 
morotibial joints darker; hind tibial comb 
with 4 bristles; hind tarsal ratio 1.72 ( 1.68- 
1.82, n = 5); claws short, slightly curved. 
Wing (Fig. 15) densely covered with mac- 
trotrichia (as determined by their insertions, 
not illustrated); radial cells obliterated; vein 
M, absent; a conspicuous forked intercalary 




Figs. 13-19. Panulcisyheleci macfiei. 13. Flagellum of female. 14, Palpus of female. 15, Wing of female. 
16. Spermatheca. 17, Flagellomeres 11-13 of male. 18. Genitalia of male, parameres removed. 19, parameres. 
Scales = 0.05 mm. 

vein present in cell r,; wing length 0.80 
(0.70-0.92, n = 5) mm, breadth 0.37 (0.32- 
0.42, n = 5) mm; costa .short, costal ratio 
0.36 (0.35-0.38, n = 5). Halter whitish. Ab- 
domen: Pale brown. One small ovoid sper- 

matheca (Fig. 16) with oblique neck, mea- 
suring 0.036 by 0.029 mm, neck 0.007 mm 
(n = 2). 

Male. — Similar to female with usual se,\- 
ual differences. Lengths of distal three fia- 



gellomeres (Fig. 17) in jjim 54-35-46. Scu- 
tellum with 7 stout setae in row. 7 slender, 
more anterior setae. Hind tarsal ratio 1.70 
(1.65-1.75, n = 5). \V"mg length 0.82 
(0.77-0.86, n = 5) mm; breadth 0.32 (0.30- 
0.33. n = 5) mm; costal ratio 0.37 (0.36— 
0.39. n = 5). Genitalia (Figs. 1S-19»: Ster- 
nite 9 with very deep, broad caudomedian 
excavation; tergite 9 moderately long, distal 
margin ne;irly straight; apicolateral process 
long, slender, with minute apical seta; cer- 
cus conspicuous, conical, somewhat p>oint- 
ed at tip. Gonixoxite twice as long as broad 
with weakly developed mesobasal tubercle; 
gonostylus nearly as long as gonocoxite. 
stout at base, abruptly narrowed at mid- 
length, nearly straight to bluntly rounded tip 
with ventral point. .Aedeagus Y-shaped; 
basal arch heavily sclerotized, distal portion 
tapering to slender pointed tip. Parameres 
(Fig. 19) with in\erted U-shaped basal apo- 
demes hea\ ily sclerotized and pair of pos- 
teriorly projecting processes arising basally 
at basal sclerite, distal half of each process 
bent \entrally. expanded apicalh with lat- 
eral tooth-like projections, tip pointed. 

Distribution. — In subantarcdc forests of 
Argentina (west of Chubut and Rio Negrot 
and Chile. 

Types. — Holotype o. Chile. Cautin 
(1 150 m). Conguillio National Park, 4/5-11- 
1988. L. Masner (CNCI). Paratypes. 12 i, 
8 d, as follows; same data as holot>pe, 12 
9,3 d (CNCI); Chile. Llanquihue. 3 km N 
Ensenada. 4-.X1I-1994. L. Quate. 2 c. Mal- 
aise trap (MLP.A); .Argentina. Nahuel Huapi 
National Park. Chall-huaco. 6-.\U-1992. G. 
Spinelli. 1 c. sweep net (MLP.A); .Argen- 
tina. Chubut. Los .Alerces National Park. 9/ 
12-.XII-1994. L. Quate. 1 d. Malaise trap 
(MLP.A); Neuquen. Laguna Epulafquen. 21/ 
23-11-2001. G. SpineUi. 1 f . Malaise crap. 

Etymology. — We are pleased to nanie 
this new species in honor of John William 
Scoli Macfie in recognition of his pioneer- 
ing paper on Patagonian biting midges that 
he co-authored w ith .A. Mactie (Ingram and 
Mactie 1931 ) and his proposal of the genus 
Paradasyhelea (Macfie 1940). 

Discussion. — This is the only species in 
the genus Paradasyhelea in which the 
males have a pair of posteriorl\ projecting 
processes arising from the basal sclerites of 
the parameres. This new species keys to 
near couplets 5a (P. boiicheri) and 5b (P. 
hrevipalpis) in the world ke\ by Clastrier 
( 1 989). but both of these species differ from 
P. macfiei in ha\ing 4-segmented palpi. 


We gratefully acknowledge grants to 
GRS from the National Geographic Society 
and the Eppley Foundation for Research. 
GRS is grateful to Hari Bhat and the late 
Larry Quate for their companionship during 
the 1994 field trip to Patagonia. Special 
thanks are extended to Zoe Adams 
(BMNH) for information on the first names 
of Drs. Ingram and Macfie. W"e also thank 
Art Borkent and Steve Murphree for their 
helpful re\iews of an e:irlier draft of the 

Liter-\tlre Cited 

Borkent. .A. i995. Biting Midges in the Cretaceous 
.Amber of North .America (Diptera: Ceratopogo- 
nidaeV Backuys Publishers. Leiden. The Nether- 
lands. 23" pp. 

Borkent. .A. and W. L. Grogan. Jr. 1995. .A re\ision of 
the genus Ceraiopogon Meigen with a discussion 
of phylogenetic relationships, zoogeography, and 
bionomic divergence (Diptera: Ceratopogonidae). 
Memoirs of the Entomological Socier> of Wash- 
ington 15: I-I9S. 

Borkent. .A. and \V. \V. VMrth. 1997. World species of 
biting midges (Diptera: Ceratopogonidae). Bulle- 
tin of the .American Museum of Natural History. 
No. 233. 257 pp. 

Clastrier. J. I9S9- Ceratopogonidae de Nouvell-Cale- 
donie. \ U- Genre Paradasyhelea (Diptera. Ne- 
matoceraK Re\Tie Francaise d'EntomoIogique II: 

Di>wnes. J. .A. and W. W. Wlrth. 1 98 1. Chapter 28. 
Ceratopogonidae. pp. 393 — %Z\. In Mc.AIpine. J. F, 
B- Peterson. G. E. Shewell. H. J. Teske>. J. R. 
Vockeroth. and D. M. Wood. eds. Manual of Ne- 
arctic Diptera. \'oI. I. .Agricultiut Canada Mono- 
graph, 27. 674 pp. 

Elson-Harris. M. M. and D. S. Kenle. 1985. .A new 
species of Paradasyhelea Mactie (Diptera: Cera- 
topogonidae). with descriptions and keys to the 
inunature stases of .Australian Paradasyhelea. 



Journal of the Australian Entomological Society 
24: 233-240. 

Ingram, A. and J. W. S. Macfie. 1931. Ceratopogoni- 
dae. pp. 155-232. In Diptera of Patagonia and 
South Chile. Part II. Ease. 4. 

Kettle. D. S. and M. M. Elson. 1975. The immature 
stages of Paradasyhelea minuta Wirth and Lee 
with a note on adult antennal sensilla and a dis- 
cussion on the relationships of the genus Para- 
dasyhelea Macfie. Journal of the Australian En- 
tomological Society 14: 255-261. 

Macfie. J. W. S. 1932. New Zealand biting midges 
(Diptera. Ceratopogonidae). Annals of Tropical 
Medicine and Parasitology 26: 23-53. 

. 1940. The genera of Ceratopogonidae. .Annals 

of Tropical Medicine and Parasitology 34: 13-30. 

Spinelli, G. R. 1987. Notas sobre Ceratopogonidae 
(Diptera Nematocera) de la Reptiblica Argentina. 
VI. Las hembras de Paradasyhelea brevipalpis y 
de Macrurohelea paracaudata. Limnobios 2: 

Spinelli. G. R. and W. L. Grogan. Jr. 1999. A new 
species of Macrurohelea Ingram & Macfie. and 
new records of bitina midses of the tribes Culi- 

coidini and Ceratopogonini from Tierra del Fuego 
and the Magallanes (Diptera: Ceratopogonidae). 
Proceedings of the Entomological Society of 
Washington 101: 708-713. 

Wirth. W. W. 1981. Paradasyhelea harrisoni n. sp. 
from the Auckland Islands, and additional records 
of subantarctic Ceratopogonidae. New Zealand 
Journal of Zoology 8: 383-386. 

Wirth. W. W. and E S. Blanton. 1969. A new Nearctic 
species of the genus Paradasyhelea Macfie (Dip- 
tera: Ceratopogonidae). Pan-Pacific Entomologist 
45: 97-100. 

Wirth. W. W. and D. J. Lee. 1959. The genus Para- 
dasyhelea Macfie. with descriptions of two new 
species from eastern Australia (Diptera: Cerato- 
pogonidae). Bulletin of the Brooklyn Entomolog- 
ical Society 54: 114-121. 

Winh. W. W. and N. Marston. 1968. A method for 
mounting small insects on microscope slides in 
Canada balsam. Annals of the Entomological So- 
ciety of America 61: 783-784. 

Wirth. W. W., N. C. Ratanaworabhan. and E S. Blan- 
ton. 1974. Synopsis of the genera of Ceratopo- 
gonidae (Diptera). Annales de Parasitologic Hu- 
maine et Comparee 49: 595-613. 


105(3). 2003. pp. 578-591 


M.AlTthew S. Wall.ace. Lewis L. Deitz. and 
Mark J. Rothschild 

(MSW. LLD) Depariment of Entomology. North Carolina State University. Raleigh. 
NC 27695-7613. U.S.A. (e-mail:; 
(MJR) Miu-yland Department of Agriculture. Plant Protection and Weed Management 
Section, 27722 Nanticoke Road. Unit 2. Salisbury. MD 21801. U.S.A. (e-mail: 

Ahsmict. — This checklist of treehoppers (Hemiptera: Membracidae) from Great Smoky 
Mountains National Park documents a total of 53 species in 25 genera, based on exami- 
nation of museum specimens and 3 collecting trips in association with the All Taxa Bio- 
diversity Inventory (ATBI). The park's treehopper fauna represents roughly half of the 
treehopper species reported from North Carolina. Twenty-four new county records are 
given for North Carolina species. Moreover. 28 species are new state records (= first 
published records) for Tennessee, increasing its known treehopper fauna to 51 species (all 
listed herein). Fifty-two species are apparently new records for GSMNP. Cades Cove, an 
expansive meadow region in the western portion of the park, is especially species rich 
w ith 35 species, many of which feed on oaks. 

Key Words: Membracidae, treehoppers. Great Smoky Mountains National Park. Smo- 
kies, ATBI. Tennessee. North Carolina, host plants 

Great Smok> Mountams National Park 2000. Sharkey 2001). The All Taxa Biodi- 
(GSMNP). located in the Southern Appa- versity Inventory (ATBI) was created to ad- 
lachian Mountains, is among the most spe- dress this lack of knowledge of native fau- 
cies-rich temperate en\ ironments in the na. Its major goal is to document every liv- 
world (Pedersen 1999. Brown 2000. Shar- ing organism in the park to provide a 
key 2001). The park straddles the North benchmark of the biodiversity prior to in- 
Carolina-Tennessee border, occupying por- vasion by non-indigenous organisms (Shar- 
tions of five counties: Haywood and Swain key 2001). The following checklist repre- 
in North Carolina and Blount. Cocke, and sents the first attempt to document the spe- 
Sevier in Tennessee. It ranges in elevation cies richness of treehoppers within Great 
from 256 m to 2.025 m (Clingmans Dome). Smoky Mountains National Park. 
Although certainly diverse, the precise Treehoppers. known for their interesting 
numbers and kinds of organisms within the shapes (Figs. 1-6) and beha\ iors. are her- 
park. especially invertebrates, are largely bivorous insects common in the eastern de- 
undocumented (Pedersen 1999. Sharkey ciduous forest (Dietrich et al. 1999) and 
2001). Additionally, the diversity of the other biomes worldwide (Wood 1993). Ex- 
park is threatened by invasive animals, tensive research has been published on the 
plants, and diseases (Pederson 1999. Brown systematics. taxonomy, and biology of the 


Figs. 1-6. Representative treehoppers ot Great Smoky Mountains National Park. 1. Cyriolobus fenestratus. 
2. Aryinna c/iierci. 3, Telainona collina. 4. Enlylia carinata. 5, Archasia belfiagei. 6, Thelia bimaculata. With 
the exception of T. bimaculata. all the above species are new state records for Tennessee. C. fenestratus ( 9 
shown). .4. cjuerci ( 6 ). and T. bimaculata (3) are sexually dimorphic in coloration. Photographs by M. S. Wallace 
and L. L. Deitz. 

New World treehoppers ( Kopp and Yonke 
1973a-c, 1974; Deitz 1975; Deitz and 
Kopp 1987; Deitz 1989; Deitz and Dietrich 
1993; Wood 1993; McKamey 1998; Die- 
trich et ul. 2001). Numerous eastern North 

American treehoppers feed on various spe- 
cies in the Family Fagaceae, especially oaks 
[Que reus spp.). 

North Carolina's treehopper fauna is rel- 
atively rich and well known, with at least 


Tiible 1 . Collecting sites within Great Smoky Mountains National Park. Some locality records are incomplete 

because data labels were imprecise. 

Sites H-1 to H-1 1. North Carolina. Haywood Counts. GSMNP 

H-1. Cataloochee. grass field 

H-2. Cataloochee. Beach Grove School 

H-3. Cataloochee. Palmer Chapel 

H-4. Cataloochee. head of Rough Fork Ridge Trail 

H-5. Cataloochee. head of Big Fork Ridge Trail 

H-6. Chestnut Branch, I mi. upstream Ranger Station 

H-7. Cataloochee Creek, at USGS Gag'ng Station 

H-8. Cataloochee. elevation 853 m 

H-9. Harmon den. Pigeon River 

H-10. Sterling l = Mt. Sterling?] 

H-1 1. Cataloochee, Cataloochee Watershed, N35°35' 10.50" W83"04'53.61". elevation 1382 m 

SW-1 to SW-18. North Carolina. Swain County. GSMNP 

SW-1. Hwy-441 at Smokemont Campground. N35°33.14' W83°18.50'. elevation 701 m 

SW-2. Beetree Ridge. Thomas Divide Tr.. 0.1 mi. S of Newfound Gap Rd. 

SW-3. Hwy-441 at Tow String Road, N35°32.37' W83°17.73'. elevation 701 m 

SW-4. Hwy-441. 3.1 mi SE of NC/TN state line 

SW-5. Hwy-441. 7.3 mi. SE of NC/TN state line 

SW-6. Hwy-441 at Collins Creek Picnic Area, elevation 732 m 

SW-7. Clingmans Creek 

SW-8. nr. Shuckstack 

SW-9. nr. bunkhouse at Hazel Creek 

SW-10. Clingmans Dome 

SW- 1 1 . Bone Valley Creek 

SW-1 2. Mingus Creek (approx. 0.5 mi. N of Oconaluftee Visitors Center) 

SW-13. Big Cove Road. Malaise Trap Site #1. N35°31.15' W83°17.77' 

SW-14. Big Cove Road. Malaise Trap Site #2. N35°31.12' W83°17.45' 

SW-I5. Big Cove Road, Malaise Trap Site #3, N35°30.76' WSB'H.Sg' 

SW-I6. Big Cove Road, Malaise Trap Site #4. N35°30.64' W83°I7.87' 

SW-17. Andrews Bald. Noland Watershed. N35°32'19.77" W83°29'39.02", elevation 1757 m 

SW-18. Clingmans Dome, Noland Watershed, N35°33'37.14" W83°29'43.40", elevation 1944 m 

Sites B-1 to B-22, Tennessee, Blount County, GSMNP 

B-1. Cades Cove, N old field 

B-2. Cades Cove. Methodist Church. N35°36.44' W83°48.99'. elevation 549 m 

B-3. Cades Cove. Primitive Baptist Church. N35°36.14' W83°48.82'. elevation 549 m 

B-4. Cades Cove. Hyatt Road. N35'36.28' W83°49.36'. elevation 533 m 

B-5. Cades Cove, Missionary Baptist Church. N35°36.43' W83°49.63'. elevation 573 m 

B-6. Cades Cove, Loop Road, near entry donation box, N35°36.45' W83°47.22'. elevation 561 m 

B-7. Cades Cove. Sparks Lane 

B-8. Cades Cove. E of Cooper Road Trail. N35°36.30' W83°50.07'. elevation 564 m 

B-9. Cades Cove Campground. N35-86.17' W83°46.44'. elevation 573 m 

BIO. Cades Cove 

B-II. Cades Cove. Gregory Place 

B-I2. Cades Cove. W of John Oliver Place. N35°36.34' W83°47.88'. elevation 549 m 

B-I3. Cades Cove. Abrams Creek 

B-14. Cades Cove. 0.3 km E of Primitive Baptist Church 

B-15. Cades Cove West 

B-16. Cades Cove. Cable Mill Visitor Center. N35=35.28' W83°50.52'. elevation 536 m 

B-17. Foothills Parkway. N Look Rock 

B-18. along Lynn Camp Prong, elevation 649 m 

B-I9, Rich Mountain Trail/Road 

B-20. Cades Cove. Abrams Watershed. N35"35'31.05" W83°50' 16.94". elevation 520 m 

B-21. Cades Cove. N35.60'' W83.78°. elevation 5^6 m 

B-22. Tremont Env[ironmental| Sta[tion| 

VOLUME 105. NUMBER 3 58 

Tabic I Conlmucd 

Sites C- 1 to C-5. Tennessee. Cocke County. GSMNP 

C-1. Albright Grove. Indian Camp Watershed. N35=43'?y.76" WS3 16'30.23". elevation 1034 m 
C-2. Snakeden Ridge. Cosby Watershed. N35°44'36.23" W,S3 13'l 1.99". elevation 993 m 
C-3. Cosby Ranger Station. N35.78" W83.21°. elevation 533 m 
C-4. Sutton Ridge Overlook. Mt. Camnierer 
C-5. Cosby 

Sites SE-1 to SE-16. Tennessee. Sevier County. GSMNP 

SE-1. Sugarlands Visitor Center, elevation 488 m 

SE-2. Park Headquarters 

SE-3. Mount LeConte 

SE-4. Twin Creeks Natural Resources Center. N35'4I '05.97" W83=29'56.61". elevation 594 m 

SE-5. Greenbrier ( = Greenbriar Pinnacle?) 

SE-6. Mid. Prong Little Pigeon River, nr. Park Boundary, elevation 419 ni 

SE-7. Grotto Falls traiihead. N35°40' W83"28'. elevation 687 m 

SE-8. Little River Road. 3.1 nii. SW of Visitors Center 

SE-9. Baskins Creek Trail 
SE-10. Hwy-441 
SE-1 1. Brushy Mountain. Middle Prong Lillle Pigeon Watershed. N35 40'35.64" W83'25'50.89". elevii 

tion 1467 ni 
SE-12. Goshen Prong. East Prong Little Pigeon Watershed. N35°36'38.22" W83"32'33.77". elevation 

S95 m 
SE-1 3. Indian Gap. West Prong Little Pigeon Watershed. N35 36'39.()()" W83°26'37.44". elc\ation 

1672 m 
SE-1 4. Foothills Parkway. Wears Valley 
SE-1 5. Elkmont 
SE-1 6. nr. Gallinburti 

S9 confirmed species and another 13 spe- 
cies reported but not confirmed by voucher 
material (Dietrich et al. 1999). By compar- 
ison, Tennessee's fauna is poorly docu- 
mented with only 23 membracid species re- 
ported in the literature (Lawson 1922: 
Funkhouser 1927: Ball 1931: Meyer 1937: 
Metcalf and Wade 1965: Kopp and Yonke 
1973a-c. 1974 [distribution mapsj) — of 
which only two species were listed with 
specific locality data (Meyer 1937). 

With the exception of Helonica excelsci 
(Fairmaire 1846) (Wallace and Deitz 1999), 
we have found no published records of tree- 
hoppers from GSMNP, although numerous 
identified specimens were present in the 
park's collection. Whittaker (1952) listed 
the number of specimens (but not species) 
he collected from the family Membracidae 
in his study of foliar insects of GSMNP. 
The objectives of the present project were 
to investigate the species richness of tree- 

hiippers in the park and to provide an au- 
thoritative checklist of the known species. 

Materials and Methods 

The treehopper species records listed in 
the Results and Discussion section are 
based on specimens in the following col- 
lections: the North Carolina State Univer- 
sity Insect Collection (NCSU), Raleigh: 
Museum Collection of Great Smoky Moun- 
tains National Park (GSNP |= GSMNP 
Collection)). Gatlinburg. Tennessee: De- 
partment of Zoology and Entomology Col- 
lection, University of Tennessee (ECUT), 
Knoxville; and the National Museum of 
Natural Hi.story (USNM), Smithsonian In- 
stitution, Washington, D.C. (collection co- 
dons follow Arnett et al. 1993). Many spec- 
imens deposited at GSNP were collected by 
Dmitri Novikov who surveyed the park's 
treehoppers in 1995. Also, some specimens 
were collected at UV-lights. Malaise traps. 


Rich MounteinS Road 

■ Methodist Church 

Cooper Road Trail* 

■ Primitive 
Church Baptist Church 


Sparks! Lane 

Cades Cove 


Blount Co., TN 

.«, Road — Creek 

Cable Mill 
Visitor Center/ 

Cades Cove Loop Road 


/Laurel: Creek Road 



\ Cocke . . 


_>S^st,}C ^S>i 

^ *^0. 


Twin Creeks 
Nat. Res, Center 

y \.. Haywood\ Co 


■■•»..' /Cataloochee -/v\. 

"^v.> '••'■ 

'Xove '^k^ 


Creetif ^ 

Cades Cove 


Fontana Lake _ 

Graham Co. 

Clingmans /Q^^^,^ 
,.....°°"LV" Ridge 


-"-state Line County Line 

5 10 mi 

Fig. 7. Diagrainniatic map of Great Smoky Mountains National Park showing state and county borders, witli 
selected details of Cades Cove. Delineated by K. H. Spieler. 

or insecticidal fogging of Quercus rubra L. 
Most of the specimens deposited at NCSU 
are from recent collecting by the authors in 
May 1999, June 2000, and October 2001 in 
association with the ATBI. A majority of 
our specimens were collected by sweeping 
and beating foliage and branches using 
sweep nets and by hand picking. No at- 
tempt has been made to compare the effec- 
tiveness of various collecting methods. 

Species records are based on adult and 
nymphal specimens and are organized al- 
phabetically by genus and species. For each 
species, localities from which the speci- 
mens were collected are listed, organized 
by state, county, and the site codes listed in 
Table 1. Site descriptions were compiled 
from specimen labels and major sites are 
indicated in Fig. 7 (sites from some early 
specimens were too imprecise to map). In 
a few instances, data not explicitly stated 

on the labels, such as "Malaise trap," is 
indicated in square brackets by the authors 
if known. Data on the date(s) of collection, 
sex, host plant(s). and site of deposition 
(collection) are listed for each site record 
when available. Host plant nomenclature is 
based on Dietrich et al. (1999: see host in- 
dex, pp. 258-261) who listed the common 
and scientific names for hosts of North Car- 
olina treehoppers and on Radford et al. 
( 1968). Hosts from which both nymphs and 
adults are reported are likely true hosts: 
hosts from which only adults are known are 
less certain. The sex could not be deter- 
mined for a number of specimens with bro- 
ken or parasitoid-damaged abdomens. Met- 
calf and Wade (1965), Kopp and Yonke 
(1973a-c, 1974), McKamey (1998), and 
Dietrich et al. (1999) were used to verify 
nomenclature, check synonymies, and iden- 
tify new state and county records. The pho- 


tographs of live treehoppers (Figs. 1-6) 
were made using an MTI-' DC330E digital 
video camera. Image Pro Plus® version 3.0, 
and Adobe Photoshop* version 5.0. The 
term "new record" is used here in the sense 
of "the first published record" rather than 
the "first occunence" or "first collection" 
of a species. Undoubtedly, most of the spe- 
cies listed here have flourished in the park 
for a very long time. 

Results and Discussion 

Our examination of museum specimens 
and three seasons of collecting in associa- 
tion with the ATBI produced a total of 53 
species in 25 genera from the park. These 
records represent 60% of the treehopper 
species known (at least 89) in North Caro- 
lina (Dietrich et al. 1999). None represented 
new species records for the state, but 24 
species are noted as new county records for 
North Carolina. Twenty-eight species are 
noted as new state records for Tennessee. A 
total of 5 1 treehoppers are now recorded for 
Tennessee, 47 given in the checklist below 
and 4 others not yet found in the park: Cyr- 
lolobiis fuUginosiis (Emmons 1854). Heli- 
ria niexicana Stal 1869, Stictocephala taitr- 
iiui (Fitch 1856), and Tortislilus triliueatus 
(Funkhouser 1918) (Funkhouser 1927, Ball 
1931, Kopp and Yonke 1973a-c). All re- 
cords from Blount, Cocke, and Sevier 
Counties, Tennessee, appear to be the first 
published records for these counties (not in- 
dividually noted as new county records in 
the checklist). Of the 53 treehoppers listed 
below, 52 are apparently the first published 
records for GSMNP. Helanica e.xcelsa. pre- 
viously recorded from the park (Wallace 
and Deitz 1999), is not yet known from 
North Carolina. 

Treehoppers are known from 72 sites in 
the park (Table 1, Fig. 7), notably Cades 
Cove (TN: Blount County), Cataloochee 
(NC: Haywood Co.), Cosby (TN; Cocke 
Co.), and several sites off Hwy-441 (NC: 
Swain and TN: Sevier counties), including 
Smokemont Campground (NC: Swain Co.). 
Thirty-five species were collected from 

sites within Cades Cove, toward the park's 
western border ( 1 2 of these species have yet 
to be collected elsewhere in the park). 

Although the genera Eiinlia German Mi- 
crutalis Fowler, and Pithlilia Stiil were col- 
lected primarily from herbaceous plants, 13 
treehopper genera were collected from var- 
ious oak species. Nearly half of the North 
Carolina treehopper fauna are known to 
feed on plants within the Fagaceae, espe- 
cially oaks (Dietrich et al. 1999). Based on 
our collecting in the eastern United States. 
Cyrtolahiis Coding (represented by 9 spe- 
cies in this checklist), Ophidenna Fair- 
maire. Sinilia German and Xantholobiis Van 
Duzee feed exclusively on oaks (Dietrich et 
al. 1999, MJR unpublished observations). 
Although Wood (1993) reported that Pla- 
Tycotis vittata (Fabricius) is also restricted 
to oaks, MJR has collected populations, in- 
cluding nymphal aggregations, on Betiila. 
Castcmea. Fagiis, and Platoinis. 

Cades Cove (Fig. 7) is an ideal habitat 
for treehoppers because much of its area is 
open grassland with oaks occurring singly 
in open fields, along roadsides, in associa- 
tion with restored historic structures, and at 
the edge of open fields and forests. Tree- 
hoppers are thought to show positive pho- 
totaxis on their host plants (Johnson and 
Freytag 1997) and thus are often collected 
towards the tips of branches or on other ar- 
eas of the plant exposed to sunlight. 

Most treehoppers were collected from 
branches and foliage well within the reach 
of ordinary collecting nets. Attempts to col- 
lect from areas of continuous forest were 
limited due to the height of the foliage in 
these areas. Thus, the relative distribution 
and numbers of treehoppers listed in this 
study, especially the many species collected 
in Cades Cove, are likely proportional to 
the intensity of our collecting efforts in par- 
ticular areas. Indeed, in an early study of 
foliar insects of the Smokies. Whittaker 
(1952) collected 21 membracid specimens 
(unidentified) from various habitats in the 
park including "cove forest, oak-chestnut 
forest, heath bald, pine forest, spruce forest. 


and pine heath." Johnson and Freytag 
(1997), using sticky traps, found treehop- 
pers to be most commonly associated with 
the top and middle levels of pin oak cano- 
pies in Kentucky. Therefore, the potential 
for discovering more species in various 
habitats within the park and at higher levels 
within the various host plants is apprecia- 
ble. Notably, species such as Archasia ciii- 
riculata (Fitch 1851), Cyrtolohiis tuherosits 
(Fairmaire 1846), Ophidenna griseo Wood- 
ruff 1919, and Telamona e.xtrema Ball 1903 
are not reported in Tennessee or GSMNP. 
yet they occur commonly in neighboring 
states (Kopp and Yonke 1973a-c, 1974; 
Dietrich et al. 1999) and thus are likely to 
occur in the park. 

Checklist of the Treehoppers of 
Smoky Mountains National Park 

Aciitcilis taruireci (Say 1830). TN: Blount 
County.: Site B-1 (see Table 1): 11 July 
1995, 3 9, 2 d (GSNF). Sevier Co.: Site 
SE-1: 27 Sept. 1982, 1 9 (GSNP). Site 
SE-16: 25 June 1947, 1 9. on Erigenm 
sp. (NCSU). 

Archasia belfnigei Stal 1869. Fig. 5. TN 
(NEW STATE RECORD): Blount Co.: 
Site B-2: 28 May 1999, 3 9, 1 cJ, on 
QuercHS alba L. and Q. falcata Michaux 
(NCSU). Site B-3: 28 May 1999. 2 9,1 
S.onQ. alba (NCSU). Site B-4: 15 June 

2000. 2 (? , 1 9 , on Platanus occidentalis 
L. and Robinia pseiidoacacia L. (GSNP. 
NCSU). Site B-21: 20 May 2001. 1 9 

Atymna castaiieae (Fitch 1851). NC: Swain 
Co.: Site SW-10: 21 June 1941, 1 6 
Blount Co.: Site B-19: 15 May 1995, 3 
nymphs, on Casianea dentata (Marshall) 
Borkh.; 8 June 1995, 7 9, 16 d, on C. 
dentata: 7 July 1995, 1 9 (GSNP). 

Atymna qiierci (Fitch 1851). Fig. 2. NC: 
Site SW-1: 14 June 2000. 1 9. on Qiier- 
cus sp. (NCSU). Site SW-14: 19-26 May 

2001, 2 9, [Malaise trap] (GSNP). Site 
SW-17: 3-17 July 2001. 1 9. [Malaise 

trapl (GSNP). TN (NEW STATE RE- 
CORD): Blount Co.: Site B-5: 15 June 
2000, 1 9, on 2- stellata Wangenh. 
(NCSU). Site B-8: 15 June 2000, 1 9, on 
Q. alba (NCSU). Site B-10: 14 June 
1995, 4 9, on 2. alba (GSNP). Cocke 
Co.: Site C-3: 16 May 2001, 1 9 
(GSNP): 17 May 2001, 1 S (GSNP). 

Campylenchia latipes (Say 1824). NC: 
Haywood Co.: Site H-9: 24 June 1995, 1 
9, 1 (J, sweeping; 13 July 1995, 19,1 
i. sweeping (GSNP). Swain Co.: Site 
SW-11: 21-24 July 1964, 2 9. 7 d 
(NCSU). Site SW-16: 21-28 July 2001, 
1 d. [Malaise trap] (GSNP). TN: Blount 
Co.: Site B-20: 30 July-Aug. 2001, 1 i. 
[Malaise trap] (GSNP). Sevier Co.: Site 
SE-1: 21 May 1943, 1 nymph (GSNP). 
Site SE-3: July 1959, 1 9 (GSNP). 

Cannota mera (Say 1830). TN: Blount 
Co.: Site B-2: 28 May 1999. 1 nymph, 
on Carya sp. (NCSU). 

Cyrtolobiis arcnatiis (Emmons 1854). TN: 
Blount Co.: Site B-20: 21 May 2001, 1 
9. [Malaise trap] (GSNP). Cocke Co.: 
Site C-3: 16 May 2001. 1 9 (GSNP). 

Cyrtolohiis aiiroreus Woodruff 1924. TN 
(NEW STATE RECORD): Blount Co.: 
Site B-19: June 1995, I 9. on C?. rubra. 
insecticidal fogging (ECUT); 8 June 
1995. 1 d. on Qiiercits sp. (GSNP). Se- 
vier Co.: Site SE-4: 25 May 2000. 1 d. 
( trap] (GSNP). 

Cyrtolohiis fenestratiis (,F\U-h 1851). Fig. 1. 
NC: Haywood Co. (NEW COUNTY RE- 
CORD): Site H-10: 17 June 1994, 1 9, 
1 (5, on Qiierciis rubra, insecticidal fog- 
ging (ECUT). Swain Co. (NEW COUN- 
TY RECORD): Site SW-2: 10 June 1994, 
4 9. 1 cJ. 1 [sex?], on Q. rubra, insec- 
ticidal fogging (ECUT. GSNP). Site SW- 
17: 3-177uly 2001. 1 9, [Malaise trap] 
(GSNP). Site SW-18: 3-17 July 2001, 1 
9, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
9: 14 June 2000. 1 9, on (?. sp. (NCSU). 
Site B-19: 31 May 1994. 1 S.onQ. rub- 
ra, insecticidal fogging (ECUT). 

Cyrtolobus flavolatiis Woodruff 1924. TN 


(NEW STATE RECORD): Blount Co.: 
Site B-10: 10 June 1995. 1 9, on Qiier- 
cus alba (GSNP), 

Cyrtolobus fuscipeimis Van Duzee 19()X. 
NC: Swain Co. (NEW COUNTY RE- 
CORD): Site SW-1: 29 May 1999, 1 9, 
on Que reus alba (NCSU). Site SW-1 3: 
30 June to 7 July 2001. 1 9. [Malaise 
trap] (GSNP). Site SW-1 4: 19-26 May 
2001, 1 9, [Malaise trap]: 7 July 2001. 
1 9. [Malaise trap]; 21-28 July 2001. 1 
9, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
9: 14 June 2000. 1 9 . 2 6 . on Q. sp., 
blacklighting: 15 June 2000. 1 6. black- 
lighting (NCSU). Site B-10: 14 June 
1995, 1 9, on Q. stellata (GSNP). Site 
B-11: 8 June 1995. 1 9, on Q. alba 
(GSNP). Cocke Co.: Site C-2; 9-22 May 
2001. 1 (5, [Malaise trap] (GSNP). Site 
C-3: 16 May 2001, 1 9 (GSNP). 

Cyrtolobus maculifrontis (Emmons 1854). 
NC: Swain Co. (NEW COUNTY RE- 
CORD): Site SW-1 7: 3-17 July 2001. 1 
9, [ trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
2: 28 May 1999, 3 9, 1 i . on Quercus 
alba and Q. stellata; 15 June 2000, 3 9. 
1 6, on Q. sp. (NCSU). Site B-5: 15 June 
2000, 3 9 , 7 (5 . on 2- stellata and Q. sp. 
(GSNP NCSU). Site B-6: 28 May 1999, 
1 9, on 2- (ilb'i (NCSU). Site B-9: 14 
June 2000, 1 i. blacklighting (NCSU). 
Site B-19: 31 May 1994, 1 9, on Q. rub- 
ra, insecticidal fogging (ECUT). 

Cyrtolobus pallidijrontis (Emmons 1854). 
NC: Swain Co. (NEW COUNTY RE- 
CORD): Site SW-1 8: 6-25 June 2001. 2 
9, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD); Bk)unt Co.; Site B- 
3: 15 June 2000. 2 9. on Quercus alba 
(NCSU). Site B-5: 15 June 2000, 1 9, on 
Q. alba (NCSU). Site B-7; 28 May 1999, 
1 S. on Q. alba (NCSU). Site B-8; 15 
June 2000, 1 9. on Q. alba (NCSU). Site 
B-9; 14 June 2000, 2 9. 1 d, on Q- sp- 
(NCSU). Site B-10; 10 June 1995. 1 9. 
on Q. alba: 14 June 1995, 4 9, 1 d, on 
Q. alba (GSNP). Site B-19; 31 May 

1994, 1 9, 1 6,on Q. rubra, insecticidal 
fogging; 8 June 1995. \ 6. \ [sex.'j. on 
Q. rubra, insecticidal fogging (ECUT). 
Sevier Co.: Site SE-4: 8-15 Oct. 2001, 1 
9, [Malaise trap] (GSNP). Site SE-12: 22 
Oct.-5 Nov. 2001, 1 9, [Malaise trap] 

Cyrtolobus toi^atus Woodruff 1924. NC; 
Site SW-3; 14 June 2000, 1 9. on Quer- 
cus sp. (NCSU). 

Cyrtolobus vau (Say 1830). NC: Haywood 
Co.: Site H-11: 20 Aug- 11 Sept. 2001, 3 
9, [ trap] (GSNP). Swain Co. 

14 June 2000, 1 [sex ?|, on Quercus sp. 
(NCSU). Site SW-17: 3-17 July 2001, 1 
6, [Malaise trap[ (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
8: 15 June 2000, 1 9, on Q. alba 
(NCSU). Site B-9: 14 June 2000, 19.1 
6. on Q. sp., blacklighting; 15 June 2000, 
1 9, on Q. alba (NCSU). Site B-10: 10 
June 1995. 2 9. on 0. alba and Ostrya 
viri^iniana (Miller) K. Koch; 14 June 

1995, 1 9, on 2. alba (GSNP). Site B- 
19: 8 June 1995, 2 6 . on Q. sp.; 7 July 
1995, 1 9 (GSNP). Sevier Co.: Site SE- 
4: 25 May 2000, 1 9, [Malaise trap[ 

Enchenopa binotata (Say 1824) complex. 
NC: Swain Co. (NEW COUNTY RE- 
CORD): Site .SW-9: 14 Oct. 1986, 1 9, 
sweeping (GSNP). TN: Blount Co.: Site 
B-10: 2 July 1995, 1 9, 3 cJ. on Liriod- 
enclron tulipifera L. (GSNP). Site B-12: 

1 5 June 2000. 1 9 . 3 d , on Robinia pseu- 
doacacia (NCSU). Cocke Co.: Site C-1: 
14 Nov. 2000, 1 9, [Malaise trap] 

Eiitylia carinata (Forster 1771). Fig. 4. NC; 
Haywood Co.: Site H-9; 9 June 1995, 2 
6, on Betula nigra L. (GSNP). Swain 
Co.; Site SW-1; 13 Oct. 2001, 3 9, 2 (5. 
on Eupatorium (as Joe-pye-weed) sp. and 
Solidago sp. (NCSU). Site SW-5; 29 May 
1999, 1 9,1 (5, on Erigeron sp. (NCSU). 
Site SW-11; 21-24 June 1964, 5 9, 2 cJ 
(NCSU). Site SW-1 6: 9-16 June 2001. 1 


9, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
3: 28 May 1999, 7 ?, on Verbesiiui al- 
ternifolia (L.) Britten ex Kearney 
(NCSU). Site B-5: 14 Oct. 2001, 1 9, on 
V. iiltenujoliu (NCSU). Site B-6: 28 May 

1 999, 3 9,3 cJ , on Que reus alba, sweep- 
ing, sweeping weeds (NCSU). Site B-7: 
28 May 1999, 1 9, on V. sp.; 14 OcL 
2001. 3 9, V. olternifolia (NCSU). Site 
B-10: 19 April 1995. 1 9 on V. altenii- 
folia: 26 April 1995, 2 9, on Aeer nih- 

niin L. (GSNP). Site B-15: 19 April 
1995. 3 9.5 S. on grass (GSNP). Site 
B-20: 21 May 2001. 2 9. [Malaise trap] 
(GSNP). Sevier Co.: Site SE-1: 21 May 
1943. 1 9 (GSNP). Site SE-2: 15 June 
1950. 1 9 (GSNP). 

Glossonotus iinivittatus (Harris 1841). TN 
(NEW STATE RECORD): Blount Co.: 
Site B-2: 28 May 1999, 1 9, on Quercus 
faleata (NCSU). Site B-I3: 13 June 
1981, 1 S (GSNP). 

Hadrophallus borealis (Fairmaire 1846). 
NC: Swain Co.: Site SW-11: 21-24 July 
1964, 1 9, 2 d (NCSU). TN: Sevier Co.: 
Site SE-4: 3 July 1995, 1 9 (GSNP). 

Helonica exeelsa (Fairmaire 1846). TN: 
Blount Co.: Site B-2: 28 May 1999, 1 9. 
on Qiieretis alba (NCSU). [First reported 
from GSMNP by Wallace and Deitz 
(1999) as Heloniea treehopper]. 

Microcentrus earyae (Fitch 1851). TN 
Records: Site B-2: 14 Oct. 2001. 1 9. on 
Caiya sp. (NCSU). 

Mieroeenrnts perditus (Amyot & Serville 
1843). NC: Swain Co. (NEW COUNTY 
RECORD): Site SW-1: 29 May 1999. 1 
9. Qaereus sp. (NCSU). Site SW-3: 14 
June 2000, \ S . Q. sp. (NCSU). Site SW- 
14: 30 June to 30 July 2001. 1 6. [Mal- 
aise trap) (GSNP). TN (NEW STATE 
RECORD): Blount Co.: Site B-2: 15 June 

2000. 1 9. on Q. sp. (NCSU). Site B-3: 
15 June 2000. 1 9 . on Q. alba (NCSU). 
Site B-8: 15 June 2000. 2 9.3 S.on Q. 
alba (NCSU). Site B-10: 10 June 1995. 
2 S.on Q. alba: 14 June 1995. 3 9. on 

Q. alba: 15 June 1995. 1 9. 1 6. on Q. 
alba (GSNP). Site B-16: 15 June 2000. 1 
9, Q. alba (NCSU). 
Mierutalis ealva (Say 1830). NC: Swain 
Co.: Site SW-11: 21-24 June 1964. 4 9 
(NCSU). Site SW-15: 9 July 1999. 1 9, 
[Malaise trap] (GSNP). TN: Blount Co.: 
Site B-2: 28 May 1999. 1 9. on Quercus 
alba (NCSU). Site B-5: 15 June 2000, 1 
9, sweeping (NCSU). Site B-6: 28 May 

1999, 1 9, on Q. alba (NCSU). Site B- 
7: 28 May 1999. 1 d. on Robinia pseu- 
doaeacia (NCSU). Site B-10: 24 May 
1995. 1 i: 10 June 1995. 1 d: 14 June 
1995, 1 9 (GSNP). Site B-16: 15 June 

2000, 1 9, sweeping meadow (NCSU). 
[First recorded for TN in Davidson Co. 
(Meyer 1937).[ 

Mierutalis dorsalis (Fitch 1851). NC: 
Swain Co.: Site SW-1 1: 21-24 July 1964, 

1 9, 2 d (NCSU). 

Ophidenna definita Woodruff 1919. NC: 
Site SW-2: 10 June 1994. 19.1c?. on 
Quercus rubra, insecticidal fogging 
Blount Co.: Site B-2: 28 May 1999, 1 9, 
on Q. faleata (NCSU). Site B-7: 28 May 
1999, 1 9. on Q. imbricaria Michaux 
(NCSU). Site B-12: 15 June 2000, 1 6 
(NCSU). Site B-19: 31 May 1994. 3 9. 

2 d. 1 [sex ?]. on Q. rubra, insecticidal 
fogging (ECUT). Cocke Co.: Site C-3: 16 
May 2001. 1 6 (GSNP). Sevier Co.: Site 
SE-4: 25 May 2000. 2 9. [Malaise trap] 

Ophiderma evelymi Woodruff 1919. TN: 
Blount Co.: Records: Site B-2: 28 May 
1999. 3 9, 1 6. on Quercus faleata: 15 
June 2000. 6 9. on G. sp. (GSNP 

Ophiderma flava Coding 1893. NC: Hay- 
wood Co.: Site H-10: 17 June 1993. 4 i. 
on Quercus rubra, insecticidal fogging; 
22 July 1993. 1 9. on (?. rubra, insecti- 
cidal fogging: 12 Aug. 1993. 1 9. on Q. 
rubra, insecticidal fogging; 17 June 
1994. 1 9. on (?. rubra, insecticidal fog- 
ciniz: 1 July 1994, 1 c?. on Q. rubra, in- 



secticidal fogging (ECUT). Swain Co. 
1 Sept. 1992, 1 9 . on Q. rubra, insecti- 
cidal fogging: 23 July 1993, 1 9, on Q. 
rubra, insecticidal fogging; 9 Aug. 1993, 
1 9 . on Q. rubra, insecticidal fogging 
(ECUT): 10 June 1994. 4 9 , \5 i , on Q. 
rubra, insecticidal fogging (ECUT, 
GSNP). Site SW-17: 3-17 July 2001, 1 
S, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
19: 2 June 1993, 2 9 . on Q. rubra, in- 
secticidal fogging; 18 June 1993, 3 9, on 
Q. rubra, insecticidal fogging; 31 May 
1994, 4 9,3 6 , on Q. rubra, insecticidal 
fogging; 26 Aug. 1994, 1 9.on Q. rubra, 
insecticidal fogging (ECUT). Cocke Co.: 
Site C-3: 16 May 2001, 2 i: 18 May 
2000, 1 i (GSNP). 

Opiiiclernia flavicephala Coding 1893. NC: 
Site SW-1: 29 May 1999, 1 9, sweeping 
(NCSU). Site SW-6: 29 May 1999, I 9, 
on Quercus sp. (NCSU). 

Ophiderma pubescens (Emmons 1854). TN 
(NEW STATE RECORD): Blount Co.: 
Site B-19: 31 May 1994, I 9, on Quer- 
cus rubra, insecticidal fogging (ECUT). 

Ophiderma salainandra Faiimaire 1846. 
NC: Haywood Co. (NEW COUNTY RE- 
CORD): Site H-IO: I July 1994, 1 d, on 
Quercus rubra, insecticidal fogging 
(ECUT). Swain Co. (NEW COUNTY 
RECORD): Site SW-1 3: 9-16 June 2001, 
1 cJ, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
19: 2 June 1993. 1 9 , on Q. rubra, in- 
secticidal fogging (ECUT). 

Platycotis vittata (Fabricius 1803). NC: 
Haywood Co.: Site H-2: 29 May 1999, 1 
9, on Quercus sp. (NCSU). Site H-3: 29 
May 1999, 1 9, on Liriodendron tulipi- 
fera (NCSU). Site H-6: 5 June 1986, I 9 
(GSNP). Site H-7: 30 May 1990, 1 i 
(GSNP). Site H-U; 26 Mar. 2001, I 9, 
[Malaise trap] (GSNP). Swain Co.: Site 
SW-1: 29 May 1999, 1 9, 2 cJ,on Q. sp.; 
13 Oct. 2001. 1 i. on Q. falcata 
(NCSU). Site SW-2: 1 Sept. 1992, 2 9, 

on Q. rubra, insecticidal fogging; 5 Oct. 
1993, 1 9, on Q. rubra, insecticidal fog- 
ging (ECUT); 15 July 1994, 1 9 . on Q. 
rubra (GSNP). Site SW-7: 28 June 1995, 
I (?, on Betula alleghaniensis Britton 
(GSNP). Site SW-14: 21-28 June 2001, 
1 6, [Malaise trap] (GSNP). Site SW-17: 
12 Nov. 2000, 1 9, [Malaise trap]; 3-17 
July 2001. 1 6, [Malaise trap] (GSNP); 
9-24 Oct. 2001, 5 9,3 3, [Malaise trap] 
(GSNP); 8-24 Nov. 2001. 1 9, [Malaise 
trap] (GSNP). Site SW-1 8: 13 Nov. 2000, 

1 9. [Malaise trap]; 24 May-6 June 
2{J0I, I 9. [Malaise trap]; 6-25 June 
2001, I 9, [Malaise trap] (GSNP). TN: 
Blount Co.: Site B-2; 28 May 1999, 1 9, 
on Q. falcata (NCSU). Site B-3: 28 May 
1999, 22 9, 19 6. on Q. stellata 
(NCSU). Site B-5: 14 Oct. 2001, 6 9,1 
6. 5 nymphs, on Q. alba and Q. slellaia 
(NCSU). Site B-6: 14 Oct. 2001, 7 9,3 
6 . 7 nymphs, on Platainis occidentalis 
and Q. imbricaria (NCSU). Site B-7: 28 
May 1999, 2 9 . \ 6 . on Q. alba (NCSU). 
Site B-8: 15 June 2000, 2 9 , on Q. alba 
(NCSU). Site B-10: 8 June 1995. 1 9 . on 
Q. alba (GSNP). Site B-1 1: 8 June 1995, 

2 9. 1 c5. on C>. alba (GSNP). Site B-14: 
14 Oct. 2001. 1 6. on Q. alba (NCSU). 
Site B-17: 15 June 1995, 1 9 (GSNP). 
Site B-19: 8 June 1995, 1 6 , on Q. ve- 
lutina Lam.; 7 July 1995, 1 9 . on Q. alba 
(GSNP). Site B-20: 14-26 Feb. 2001, 1 
6. [Malaise trap] (GSNP). Cocke Co.: 
Site C-1: 14 Nov. 2000, 1 9, [Malaise 
trap]; 19 Jan. 2001, 1 3. [Malaise trap] 
(GSNP); 8-19 June 2001, 1 <5, [Malaise 
trap] (GSNP); 27 Sept.-16 Oct. 2001, 1 
6, [Malaise trap] (GSNP). Site C-2: 13 
Dec. 2000. 1 6. [Malaise trap); 30 Jan.- 
16 Feb. 2001. 1 9. [Malaise trap| 
(GSNP). Site C-4: 2 Oct. 1995. 6 
nymphs, on Castanea dentata (GSNP). 
Site C-5: 27 Sept. 2000. 1 9 (GSNP). 
Sevier Co.: Site SE-4: 3 July 1995. I 9; 
8 Nov. 1999. 1 9, [Malaise trap|; 29 Mar. 
1999, 1 9, [Malaise trap]; 2 Aug. 2000, 
1 9, [Malaise trap): 12 Sept. 2000, I 9, 
[Malaise trap]; 12 Oct. 2000. I 9. [Mai- 


aise trap]; 24 Oct. 2000, 4 9,3 6. [Mal- 
aise trap]; 27 Nov. 2000, 2 9,2 c? [Mal- 
aise trap]: 12 Dec. 2000, 2 3. [Malaise 
trap]: 16 Jan. 2001. 1 9,5 6, [Malaise 
trap]: 29 Jan.- 12 Feb. 2001, 1 9, [Mal- 
aise trap]: 16-30 July 2001, 1 J. [Mal- 
aise trap] (GSNP): 10 SepL 2001, 1 S. 
[Malaise trap] (GSNP); 15 Oct.-5 Nov. 
2001, 1 6. [Malaise trap] (GSNP). Site 
SE-5: 6 Sept. 1940, 13 9,5 ^, on Cos- 
tanea sp. (GSNP). Site SE-11: 23 Oct.- 
6 Nov. 2000, 3 9,3 6, [Malaise trap]; 6 
Nov. 2000, 1 9. 1 (J, [Malaise trap] 
(GSNP): 21 June-5 July 2001, 1 6. 
[Malaise trap] (GSNP); 13-27 Oct. 2001. 
1 9, 2 (J, [Malaise trap] (GSNP). Site 
SE-12: 10 Nov. 2000, 2 9, 1 (5, [Malaise 
trap): 14 Feb. 2001. 1 9, [Malaise trap]: 
28 Mar.-9 Apr 2001, 1 9. [Malaise trap] 
(GSNP). Site SE-13: 10 Nov. 2000, 1 9, 
1 6. [Malaise trap[ (GSNP): 26 Sept.-24 
Oct. 2001, 1 (J, [Malaise trap] (GSNP). 
Site SE-14: 1 Oct. 1986, 9 9, 11 6,1 
nymphs, on C. dentata (GSNP). 

Publilici concava (Say 1824). NC: Hay- 
wood Co.: Site H-4: 29 May 1999, 1 i. 
on Eiipatoriiiiu sp. (NCSU). Swain Co.: 
Site SW-1: 29 May 1999, 6 9. 2 (5, on 
Erigeron sp. and sweeping; 13 Oct. 2001, 
17 9, 5 (5, on Eupatorium sp. (as Joe- 
pye-weed) and Solidago sp. (NCSU). Site 
SW-6: 29 May 1999, 9 9,2 c?, on Eu- 
patorium sp., Solidago sp.. and sweeping 
(NCSU). Site SW-11: 21-24 July 1964, 
1 9 (NCSU). TN: Blount Co.: Site B-3: 
28 May 1999, 1 9, on Verbesina alter- 
nifolia (NCSU). Sevier Co.: Site SE-6: 5 
Sept. 1995, 1 9, 2 (5, sweeping riparian 
vegetation (GSNP). Site SE-7: 28 May 
1999, 1 9 (GSNP). Site SE-8; 28 May 
1999, 2 9, 3 cJ, sweeping herbaceous 
plants (NCSU). Site SE-10: 28 May 
1997, 1 6 (NCSU). [First recorded for 
TN in Davidson Co. (Meyer 1937)]. 

PublUia reticulata Van Duzee 1908. NC: 
Haywood Co.: Site H-3: 29 May 1999, 1 
S (NCSU). Site H-4: 29 May 1999, 2 9, 
on Eupatorium sp. (NCSU). Swain Co. 

16: 9-16 June 2001, 1 9, [Malaise trap] 
(GSNP). TN: Sevier Co.: Site SE-8: 28 
May 1999, 5 9,2 6. sweeping weeds, 
sweeping herbaceous plants (NCSU). 
Site SE-15: 15 June 1946, 1 9, 1 (? 

Smilia camelus (Fabricius 1803). NC: 
Site SW-1: 29 May 1999. 1 9. on Quer- 
cus sp. (NCSU). 

Spissistilus festinus (Say 1830). TN: Blount 
Co.: Site B-1: 11 July 1995, 1 6 (GSNP). 
Site B-10: 6 April 1995, 2 J ; 24 May 
1995. 1 i: 10 June 1995. 1 6 (GSNP). 

Stictocephala brevitylus (Van Duzee 1908). 
NC: Haywood Co.: Site H-3: 29 May 
1999, 2 9, on Quercus sp. (NCSU). Site 
H-4: 29 May 1999, 1 9, 1 cJ, on Sam- 
bucus canadensis L., sweeping weeds 
(NCSU). Site H-5: 29 May 1999, 3 9,2 
i,onQ. sp. (NCSU). Site H-8: 29 May 

1999. 1 9, on Ostiya virginiana (NCSU). 
Swain Co.: Site SW-1: 29 May 1999, 6 
9,4 (J, on Erigeron sp., Q. sp., Robinia 

pseudoacacia. and sweeping; 14 June 

2000. 3 9, on R. pseudoacacia (NCSU). 
Site SW-6: 29 May 1999, 5 9,3 6. Eu- 
patorium capillifolium (Lam.) Small and 
E. sp. (NCSU). Site SW-11: 21-24 July 
1964, 1 9 (NCSU). Site SW-12: 10 July 
1980, 1 6 (GSNP). Site SW-1 5: 9-16 
June 2001, 1 6. [Malaise trap] (GSNP). 
Site SW-1 6: 9-16 June 2001. 1 <5, [Mal- 
aise trap] (GSNP). TN (NEW STATE 
RECORD): Blount Co.: Site B-2: 28 May 
1999, 1 9, 1 cJ, on Rubus sp. (NCSU). 
Site B-4: 15 June 2000. 3 9, on R. pseu- 
doacacia (NCSU). Site B-7: 28 May 
1999, 1 9, 1 d, on Gleditsia triacanthos 
L.. sweeping (NCSU). Site B-10: 10 June 
1995, 1 9 (GSNP). Sevier Co.: Site SE- 
4; 9 May 1995, 1 9 (GSNP). Site SE-8: 
28 May 1999, 1 9, sweeping herbaceous 
plants (NCSU). 

Stictocephala diceros (Say 1824). NC: 
Haywood Co.: Site H-1: 20 July 1995, 1 
6 (GSNP). TN: Sevier Co.; Site SE-2: 15 
Aug. 1947, 1 S (GSNP). 

Stictocephala lutca (Walker 1851). NC: 



Haywood Co.: Site H-1: 16 May 1995. 1 
9.16 (GSNP). Swain Co. (NEW 
COUNTY RECORD): Site SW-3: 14 
June 2000, 2 9. 1 d. on Qiiercus sp. 
(NCSU). TN: Blount Co.: Site B-6: 28 
May 1999. 1 9. sweeping herbaceous 
plants (NCSU). Site B-19: 15 May 1995. 

2 9. 1 d. on Q. velutiiun 26 May 1995. 

3 9 . on Q. vehitina and Querciis sp. 
(GSNP). Sevier Co.: Site SE-4: 25 May 
2000. 1 (5. [Malaise trap] (GSNP). Site 
SE-9: 17 June 1995. 1 9 (GSNP). 

Stictocephala militaris (Gibson and Wells 
Blount Co.: Site B-22: 20 Aug. 1999. 5 
(J, moth trap — blacklighting (GSNP). 

Telainona collina (Walker 1851). Fig. 3. 
NC: Haywood Co. (NEW COUNTY RE- 
CORD): Site H-9: 9 June 1995 [found 
dead]. 1 [sex ?] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
6: 28 May 1999. 1 nymph, on PUihiims 
occidentalis (NCSU). Site B-7: 28 May 
1999. 1 c?. 1 nymph, on P. occideiihilis 
(NCSU). Sevier Co.: Site SE-4: 22 June 
1999. 1 9. 1 cJ. [Malaise trapj: 4 July 

1999. 1 9. [Malaise trap] (GSNP). 
Teluimma clecomtu Ball 1903. TN (NEW 

STATE RECORD): Blount Co.: Site B- 
3: 28 May 1999. 1 .5. on Querciis stellata 
(NCSU). Site B-19: 31 May 1994. 1 cJ. 
on Q. rubra, insecticidal fogging 
(GSNP). Sevier Co.: Site SE-4: 27 Nov. 

2000. 1 9. [Malaise trap] (GSNP). 
Telainona inaculata Van Duzee 1908. NC: 

Site SW-14: 30 June to 7 July 2001. 1 6. 
[Malaise trap] (GSNP). 
Telanuma nionticola (Fabricius 1803). NC: 
Haywood Co. (NEW COUNTY RE- 
CORD): Site H-10: 17 June 1993. 2 6. 
19,1 [sex ?], on Quervus rubra, insec- 
ticidal fogging (ECUT, GSNP); 17 June 
1994, 1 9 . on Q. rubra, insecticidal fog- 
CORD): Blount Co.: Site B-2: 28 May 
1999, 1 9, on (2. alba (NCSU). Site B- 
5: 15 June 2000. 1 9.1 6 . on Q. stellata 
and Robinia pseudoacacia (NCSU). Site 

B-9: 14 June 2000. 1 6. on Q. sp. 
(NCSU). Cocke Co.: Site C-3: 18 May 
2000. I 9. 1 d (GSNP). 

Telainona recUvata Fitch 1851. NC: Swain 
SW-2: 1 Sept. 1992. 1 9, on Quercus 
rubra, insecticidal fogging (ECUT). 

Telamona unicolor Fitch 1851. TN (NEW 
STATE RECORD): Blount Co.: Site B- 
2: 28 May 1999. 1 d. on Carya sp. 
(NCSU). Site B-6: 28 May 1999. 1 9. on 
C. sp. (NCSU). 

Thelia bimaculata (Fabricius 1794). Fig. 6. 
NC: Swain Co.: Site SW-8: [no date], I 
9 (GSNP). TN: Blount Co.: Site B-4: 15 
June 2000. 2 9. 1 d. on Robinia pseu- 
doacacia (NCSU). Site B-12: 15 June 
2000, 1 9. 1 d. on /?. pseudoacacia 
(NCSU). Site B-18: 1 Sept. 1988, 1 d 
(GSNP). Sevier Co.: Site SE-2: 2 Sept. 
1^>37. 1 9 (GSNP). Site SE-4: 3 July 
1995. I 9 (GSNP). Site SE-15: II Aug. 
1947. 1 d (USNM). 

Tortistilus abnonnus (Caldwell 1949). TN 
(NEW STATE RECORD): Blount Co.: 
Site B-2: 15 June 2000. 2 d. on Quercus 
sp.. sweeping (GSNP NCSU). 

Tortistilus inennis (Fabricius 1775). TN: 
Blount Co.: Site B-16: 15 June 2000. 2 
d. sweeping (GSNP NCSU). 

Tylopelta gibbera (Stal 1869). NC: Swain 
Co.: Site SW-11: 21-24 July 1964. I 9. 
2 d (GSNR NCSU). TN: Sevier Co.: Site 
SE-2: 15 Aug. 1947, 1 9 (USNM). 

Vanduzea arquata (Say 1830). NC: Swain 
Co.: Site SW-1: 28 May 1999, 1 [sex ?], 
70 nymphs [ethanol], on Robinia pseu- 
doacacia: 14 June 2000, 15 9, 2 d, on 
R. pseudoacacia: 13 Oct. 2001, 1 d, on 
Q. alba? (NCSU). Site SW-3: 14 June 
2000, 4 9 , I d , on /?. pseudoacacia 
(NCSU). Site SW-4: 29 May 1999, 2 
nymphs, on R. pseudoacacia (NCSU). 
Site SW-8: [no dale], 2 9 (GSNP). TN: 
Blount Co.: Site B-2: 28 May 1999, 2 9, 
7 d, [1 9. 1 d, and 10 nymphs in eth- 
anol], on R. pseudoacacia (NCSU). Site 
B-10: 20 July 1995. 1 d (GSNP). Site B- 
12: 15 June 2000. 3 9. 6 d. on /?. p.seu- 


doacacki (NCSU). Site B-19: 7 July 
1995, 1 9 (GSNP). Sevier Co.: Site SE- 
4: 8 July 1995. 1 9 (GSNP); 15 Oct. 
2001, 7 9. 1 6.2 nymphs, on R. pseii- 
doacacia (NCSU). Site SE-8: 28 May 
1999, 1 cJ, on R. pseudoacacia (NCSU). 

Xantholobiis intermedins (Emmons 1854). 
NC: Haywood Co. (NEW COUNTY RE- 
CORD): Site H-10: 17 June 1994. 5 9, 
2 cJ, on Quercus rubra, insecticidal fog- 
ging; 1 July 1994, 3 9.1 [sex ?], on Q. 
rubra, insecticidal fogging (ECUT, 
CORD): Site SW-2: 1 SepL 1992, 1 9, 
on Q. rubra, insecticidal fogging; 9 July 
1993. 1 6 . on Q. rubra, insecticidal fog- 
ging; 10 June 1994, 1 9,1 6 , on Q. rub- 
ra, insecticidal fogging; 28 June 1994. 1 
9,4 6 ,on Q. rubra, insecticidal fogging; 
15 July 1994, 1 9. on 0. rubra, insecti- 
cidal fogging (ECUT, GSNP). Site SW- 
17: 3-17 July 2001, 1 9, [Malaise trap] 
(GSNP). Site SW-18: 6-25 June 2001, 1 
9, [Malaise trap] (GSNP). TN (NEW 
STATE RECORD): Blount Co.: Site B- 
19: 31 May 1994. 2 9, on Q. rubra, in- 
secticidal fogging; 26 Aug. 1994, 1 9, on 
Q. rubra, insecticidal fogging (ECUT). 

Xantholobus lateralis Van Duzee 1908. TN 
(NEW STATE RECORD): Blount Co.: 
Site B-19: 15 May 1995, 1 9, on Quer- 
cus velutitm (GSNP). 

Xantholobus muticus (Fabricius Mil). TN 
(NEW STATE RECORD): Blount Co.: 
Site B-5: 15 June 2000. 1 9 . on Quercus 
sp. (NCSU). 


We thank Karen H. Spieler for preparing 
Fig. 7: our colleagues R. L. Blinn, B. K. 
Cassel, and B. M. Wiegmann for collecting 
treehoppers listed in this work; R. L. Blinn 
(NCSU), D. Defoe (GSNP). P L. Lambdin 
(ECUT). S. H. McKamey (USNM), C. 
Parker (GSNP), and I. Stocks (GSNP) for 
allowing us to examine specimens in their 
care; A. Krings and J. M. Stucky (Depart- 
ment of Botany, North Carolina State Uni- 
versity) for assistance with plant identifi- 

cations and botanical nomenclature; and R. 
L. Blinn, W. E Gimpel, P L. Lambdin, S. 
H. McKamey, H. H. Neunzig, D. D. Trieff, 
B. M. Wiegmann, and one anonymous re- 
viewer for helpful comments on the man- 
uscript. This material is based upon work 
supported by the National Science Foun- 
dation under Grant Nos. 9815867 and 
9978026 and by the North Carolina Agri- 
cultural Research Service. 

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World, 2nd ed. Sandhill Crane Press, Gainesville, 
Florida, vi + 310 pp. 

Ball. E. D. 1931. A monographic revision of the tree- 
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Brown. M. L. 2000. The Wild East: A Biography of 
the Great Smoky Mountains. University Press of 
Florida. Gainesville, x.xi + 457 pp. 

Deitz. L. L. 1975. Classification of the higher catego- 
ries of the New World treehoppers (Homoptera: 
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periment Station Technical Bulletin 225: [i-iv], 1- 

. 1989. Bibliography of the Membracoidea 

(Homoptera: Aetalionidae, Biturritiidae. Membra- 
cidae. and Nicomiidae) 1981-1987. North Caro- 
lina Agricultural Research Service Technical Bul- 
letin 290: 1-31. 

Deitz. L. L. and C. H. Dietrich. 1993. Superfumily 
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the Membracoidea (Homoptera: Aetalionidae. Bi- 
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1980. North Carolina Agricultural Research Ser- 
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morpha: Membracoidea!. Systematic Entomology 
26: 213-239. 

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and Polyglyptini (Homoptera: Membracidae). 
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105(3), 2003, pp. 592-598 


Kurt M. Pickett 

Department of Entomology, Mu.seum of Biological Diversity, 1315 Kinnear Road, The 
Ohio State University, Columbus, OH 43212, U.S.A. (e-mail: 

Ahsinici. — Apoica ambracariiia, n. sp., is described from queens, workers, and males. 
It is known from Rio Branco, Brazil, Sao Felix do Xingu, Brazil, and Tuparro Cerro 
Tomas, Colombia. Male genitalia and descriptive characters of all castes are illustrated. 
This is the ninth described species of the nocturnal genus Apoica. 

Key Words: Apoica amhracariiui. new species, Vespidae, Polistinae 

Apoica Lepeletier is a genus of large- 
bodied, swarm-founding social wasps (Hy- 
menoptera: Vespidae; Polistinae, Epiponini) 
distributed from Mexico to Argentina. 
Apoica are distinguished from other epi- 
ponines by their unusually large ocelli and 
associated nocturnality. Some Apoica are 
light in overall color, including the yellow 
A. fiavissiina Vecht, the cream-colored A. 
pallens (¥.). and the white A. gelida Vecht. 
Some Apoica (A. arhorea de Saussure and 
A. strigata Richards) have a "striped" ap- 
pearance on the metasoma, generated by 
pale areas of the terga. Others are more or 
less uniformly dark brown to black in color 
with few pale markings, including A. pal- 
lida (Olivier), A. thoracica du Buysson, and 
A. albimaciila (¥.). Richards (1978) last re- 
vised the group, discussing these eight spe- 
cies. Although the species described here is 
generally a dark amber color, it is easily 
distinguished from other dark-bodied Apoi- 
ca by its possession of a unique suite of 

Apoica ambracarina Pickett, new specie.s 

(Figs. 1-9. lOA, IIB) 

Diagnosis. — Predominantly amber col- 
ored. Posterior border of terminal metaso- 

mal lergum pale yellow. Metanotum 
rounded, with longitudinal, median carina. 
Propodeal bristles erect, not curved. Eye 
bristles present and prominent over the en- 
tire surface of the eyes. Scuta! bristles 
reaching posterior margin of scutum. Hab- 
itus of queen, worker, and male shown in 
Fig. 1. 

Queen. — Morphology: Average wing 
length 1.78 cm (n = 5, ±0.027). Prominent 
longitudinal carina between antennal sock- 
ets (Fig. 2A). Median scutal sulcus wide, 
flaring anteriorly. Metanotum rounded, with 
obvious carina (as in Fig. 5). Color: Ter- 
minal two flagellomeres of antenna pale. 
Body almost entirely amber colored, with 
mesosoma slightly darker amber than meta- 
soma. Scutum with two dark brown par- 
apsidal funows, fusing posteriorly (Fig. 
3A). Posterior border of the terminal meta- 
somal tergum pale yellow (Fig. 4A). 

Worker. — Morphology: Average wing 
length 1.85 cm (n = 5, ±0.035). Prominent 
carina between antennal sockets, as in 
queen (Fig. 2B). Median scutal sulcus as in 
queen. Metanotum rounded, with obvious 
carina (Fig. 5). Color: Scutum with two 
dark brown parapsidal furrows, fusing pos- 
teriorly (Fig. 3B). Posterior border of the 


Fig. 1. Habitus of a queen (A), worker (B). and male (C) of Apoica ciiithrucunna. Scale bars = .5.0 


Figs. 2-5. Apoica amhracarina. 2, Frontal view of heads of a queen (A), worker (B). and male (C) .showing 
carina between antennal sockets; scale bars = 3.0 mm (A), 3.0 mm (B). and 2.0 mm (C). 3, Dorsal view of 
mesosoma of a queen (A), worker (B). and male (C) showing dark brown parapsidal furrows: scale bars = 1.0 
mm. 4. Dorsal view of terminal metasomal tergum of a queen (A), worker (B). and male (C) showing pale 
(yellow) posterior edge: scale bars = 2.0 mm. 5. (A) Oblique posterior view of mesosoma of worker holotype 
of A. amhracarina showing metanoial carina, enlarged in (B) for clarity: scale bars = 2.0 mm (A) and 1.0 mm 


terminal metasomal tergum pale yellow 
(Fig. 4B). Metasoma slightly darker than in 

Male. — Morphology: Average wing 
length 1.63 cm (n = 5, ±0.044). Prominent 
carina between antennal sockets (Fig. 2C). 
Median scutal sulcus less developed than in 
female. Many prominent bristles on entire 
scutum, all curving medially. Metanotal ca- 
rina slight, nearly absent. Genitalia: Distal 
end of aedeagus punctured, with sensilla 
emerging from punctures (Fig. 6a). Central 
region of aedeagus with numerous denticles 
(Fig. 6B). Parameral spine extending far be- 
low body of paramere (Fig. 8) Paramereal 
spine with setae straight or curving slightly 
(Fig. 8A). Tip of digitus attaches to cuspis; 
digitus rounded and curved at tip (Fig. 9). 
Cuspis with prominent, curved setae (Fig. 
9). Color: Scutellum and metanotum most- 
ly pale yellow. Scutum with two dark 
brown parapsidal furrows, fusing posteri- 
orly (Fig. 3C). Posterior border of terminal 
metasomal tergum pale yellow (Fig. 4C). 
Generally lighter in overall color than fe- 

Type material. — Holotype worker, la- 
beled "Rio Branco, Acre State, Brazil, coll. 
S. Mateus and F B. Noll, 10 X 1998" All 
paratypes from same colony, same data as 
holotype. Paratypes include 80 workers, 1 1 
males, and 9 queens, labeled as such. Ho- 
lotype and 40 worker, 5 male, and 5 queen 
paratypes are deposited in the Museu de 
Zoologia, Universidade de Sao Paulo, Bras- 
il. The remaining paratypes are deposited in 
the Ohio State University Insect Collection, 
Columbus (20 workers, 3 males, 2 queens) 
and the American Museum of Natural His- 
tory. New York (20 workers. 3 males. 2 

Distribution. — Apoica anibracarina is 
known from the type locality: Sao Felix do 
Xingu, Brazil; and Tupano Cerro Tomas, 

Other material examined. — 1 9: "Co- 
lumbia Vicnada PNN, Tuparro Cerro Tomas 
140 m, 5°2rOO"N 67°5r36"W, 8-28 aug 
2000 W, Villalba, Sample No 513" (pers. 

coll. C. Sarmiento); 1 9: "Columbia Vic- 
nada PNN, Tuparro Cerro Tomas 140 m, 
5°2 1 '00"N 67°5 1 '36"W, 1 8-28 aug 2000 W. 
Villalba, Sample No 516" (pers. coll. C. 
Sarmiento); 1 9 : "Columbia Vicnada PNN, 
Tuparro Cerro Tomas 140 m, 5°2r00"N 
67°51 '36"W, 29 jun - 15 Jul 2000 Malaise, 
Sample No 269 W. Villalba" (pers. coll. C. 
Sarmiento); 5 9: "Moyen XINGU (Bresil), 
Mission M. Boulard, P. Jauffret et P. Pom- 
panon. Museum PARIS, SAO FELIX DO 
XINGU, 1-4-X-1975" (MNHN); 4 9: 
"Moyen XINGU (Bresil), Mission M. Bou- 
lard. P. Jauffret et P. Pompanon, Museum 
IX- 1975" (one determined A. tltonicica B. 
Sigwalt 1984) (MNHN). 

Nest. — The nest was not collected with 
the colony of A. anibracarina, so a precise 
description is not possible. However, I was 
told that the nest was approximately 15 m 
high in a tree and appeared to be a typical 
Apoica subgenus Apoica (c.f., Apoica ar- 
horea. Vecht 1972; 738. photo I) nest in all 
other respects (pers. comm.. F. B. Noll). 

Etymology. — The name refers to the 
general amber color of the wasp and the 
presence of the longitudinal, medial metan- 
otal carina. 

Remarks. — Apoica anibracarina is dis- 
tinguishable from other Apoica by its amber 
color and the presence of a longitudinal, 
median metanotal carina. Although the me- 
tasoma of some Apoica is light brown or 
yellowish, the presence of the carina distin- 
guishes A. anibracarina from all other 
known Apoica species, except A. albima- 
ciila. A. albimaciila also has a metanotal ca- 
rina, but close examination of the carinae 
reveals that they are not homologous. The 
posterior margin of the metanotum is 
rounded in A. anibracarina (Fig. lOA), as 
is typical of most Apoica, whereas A. al- 
bimaciila has a sharply angled metanotum 
posteriorly (Fig. lOB) and a much more 
prominent metanotal carina. Also, the cari- 
na of .4. albimaciila is near the posterior 
margin of the metanotum (Fig. 1 lA) where- 
as the carina of A. anibracarina is more an- 


Figs. 6-9. Male genitalia of Apoica aiubnuaiiiui. 6. Aedeagus (venlral view), witli enlargements of sensilla 
(6a) and medial denticles (6b). 7, Aedeagus (lateral view). 8, Right paramere (interior view) with enlargement 
of spine showing setae (8a). 9, Right volsella. (lateral view). Scale bars for 6-8 = 0.01 mm. Scale bar for 9 = 
0.005 mm. 


Fig. 10. Lateral view of /l/;o/((( <//;/ 
i.s rmiiidcd posteriorly (see arrow), whe 
Scale bars = 2.0 mm. 

,1,111,1 (.A) and .A «//>/ 
llio iiictanotum ol .V 

'ii,„ ula 

he iiietanoIiHii ol ,\. umliiuiuit 
more shaiply angled (see arro' 

terior (Fig. I IB). Although Richards (iy7,S: 
262) mentioned in his key to Apoica that A. 
thoracicci has a "Metanotum usually round- 
ed, rarely with traces of a keel," this is in- 
accurate. A. thoracicci does not have a nie- 
tanotal keel (Fig. 1 IC). Richards may have 
included A. ambracarina in his concept of 
A. thoracicci if he had seen specimens, hi 
particular, he may have seen some speci- 
mens of A. ambracarina in the Museum 
National D'Histoire Naturelle that were 
identified as A. thoracica (see Material Ex- 

In his discussion of Apoica thoracica. 
Richards (1978: 268) stated 'In the col- 
lection at Paris are two females marked 
type, Guyane francaise: Camopi, 1900 (F. 
Geay). These are specimens of /4. alhinui- 
ciila. The whole gaster, including the end- 
band of gastral tergite 1 to tip of tergite 6 
is yellowish brown (perhaps immature) but 
the pale yellow at the base of tergite 2. 

though faded, can still be distinctly seen if 
carefully looked for. Moreover, the metan- 
otum has a distinct keel." When I first saw 
the colony of the species described here. I 
thought it was equivalent to the two un- 
usual specimens described by Richards 
(1978), because they are light brown to 
amber colored and posses a metanotal ca- 
rina. However, since studying these speci- 
mens, I have concluded that, although they 
are much lighter than any other A. alhi- 
luaciila specimens I have observed, they 
are indeed A. albimaciila and not members 
of A. ambracarina. The posterior edge of 
the metanotum of Richards" specimens is 
sharply pointed, as in A. albimaciila (Fig. 
lOB), but this edge is rounded in A. am- 
bracarina (Fig. lOA). Also, A. aml)rac(0- 
iini has a pale mark on the posterior tip of 
the terminal metasomal tergum and dark 
brown parapsidaj furrows on the scutum. 

Fig. I I . Oblique posterior views ol metanola oi Ai>oiLci albiinacula (A). /I. ,iiiilii,i, ,ii tim (holotypc) (B), and 
A. thnnicica (C). Ttie carina of /i. alliimacula is near the po.sterior margin of the nielanottim (sec arrow) whereas 
the carina of A. ainhracarina is more anterior and more blimt (see arrow); A. llionicicci has no tnetanotal carina. 
Scale bars = 1.0 mm. 


but these are absent on A. alhiinacula and 
Richards' specimens. 


I thank Sidnei Mateus. Fernando B. Noll, 
and Ronaldo Zucchi for advice and speci- 
mens. Conversations with Marc A. Bran- 
ham. James M. Carpenter. J. S. Hans Klom- 
pen. and Peter Kovarik greatly improxed 
my illustrations, and Marc A. Branham and 
Norman F. Johnson kindly reviewed the 
manuscript. This work was supported by 
grants to KMP from the OSU Office of In- 
ternational Travel, the American Museum 
of Natural History, Norman F. Johnson, and 
John W. Wen/el. Material examined for this 
work came from the Museum National 

D'Histoire Naturelle (with thanks to Claire 
Villemont). the American Museum of Nat- 
ural History, The Natural History Museum 
(BMNH), the Nationaal Natuurhistorische 
Museum, and private collections in the lab- 
oratories of Ronaldo Zucchi and J. M. F. de 

Literature Cited 

Ricliarcls. O. W. 1978. The social wa.sps of the Amer- 
icas, excluding the Vespinae. British Museum 
(Natural History). London. 580 pp. 

Vccht. J. van der. 1972. The social wasps (Vespidae) 
collected in French Guyana by the Mission du 
Museum National D'Histoire Naturelle with notes 
on the genus Apoica Lepeletier (1) (2). Annales 
de la Societe Entomologique de France (N.S.) 
S(3): 733-743. 


105(3). 200.3. pp. .599-610 




A. G. WntifiLRR. Jr. 

Department of Entomology. Clemson University, Clemson, SC 29634-0365, U.S.A. (e- 

Abstract. — Specialization on mosses, previously unknown in the Fulgoroidea and ap- 
parently also for any auchenorrhynchan, is reported for the delphacid planthopper Jave- 
sella opcicci (Beamer). New state records for this seldom-collected species are Alabama, 
Georgia, Massachusetts, New York, North Carolina, South Carolina, and Tennessee. 
Nymphs developed mainly on the gametophores of common hair-cap moss. Polytrichum 
commune Hedw. (Polytrichaceae), on granite flatrocks and in other communities, and once 
were found on another polytrichaceous moss, Polytrichastniiu alpinum (Hedw.) G.L. Sm. 
At Boggs" Rock, a granite Hatrock in northwestern South Carolina, late instars overwin- 
tered in mats of Polytrichum commune, and adults developed by mid- to late March. The 
only macropterous adults (n = 5) observed from June 1996 to July 1998 were found 
between 10 April and 2 May 1997; all other adults were brachypters. First instars of the 
season's first generation appeared from mid- to late May, with adults appearing from mid- 
June to early July. The first instars observed in August were those of an overwintering 
generation. Fluctuations in the numbers of planthoppers are discussed in relation to the 
condition of moss colonies (green vs. drought stressed) and weather (precipitation, tem- 
perature, and evaporation). In the laboratory, nymphs fed on thicker stems of the moss. 
The use of mosses is considered a derived host association in the evolution of the Del- 

Key Words: Insecta, Delphacidae, moss feeding, Polytricluini coinnuine. Polytrichustrum 
(ilpinimi. granite outcrops 

The large gaps in our knowledge of the son 1969, Lawrey 1987), the insect fauna 

bryosystem, coupled with the ubiquity of as.sociated with bryophytes evidently is not 

mosses, leaves great scope for original re- ^^tably diverse when only those phytopha- 
search by bryotaunal ecolosists. , ^ i . .i • j ■ 

^ , . . ,,. ,. „„„ 2es known to complete their development 

— I.M. Kmchin 1990 " r- r 

on these plants are considered. Among 

Studies on insect-plant interactions have bryophytes, insects are found mainly on 

emphasized vascular plants, the Tracheo- mosses — Bryophyta sensu stricto — rather 

phyta, including seedless groups such as than on hornworts and liverworts. Moss-in- 

fems, but especially the more economically habiting insects, which are recorded for 

important gymnosperms and angiosperms most of the hemimetabolous and holome- 

among the seed plants. Even though insects tabolous orders (Gerson 1969), can be 

of nonvascular plants (Bryophyta) general- found on both aquatic and terrestrial moss- 

ly have been neglected by researchers (Ger- es. In many cases, whether the species use 



mosses for food or only for shelter is not 

In the Homoptera. used in its traditional 
sense (the group is paraphyletic [e.g.. von 
Dohlen and Moran 1995]). moss feeders in- 
clude several sternorrhynchan groups: 
aphids. such as species of Melaphis Walsh. 
Mnscaphis Borner, and Myzodiiim Borner 
(e.g.. Patch 1938. Borner 1952. Muller 
1973. Smith and Knowlton 1975, Richard- 
son 1981. Moran 1989. Dolling 1991. 
Thomas and Lombard 1991), as well as cer- 
tain ensign scales (Ortheziidae) (e.g., Kozar 
and Miller 1999. 2000) and mealybugs 
(Pseudococcidae) (Williams 1985). The use 
of mosses as true hosts by ortheziids is 
largely undocumented (Dolling 1991). 

Bryophagy apparently has remained un- 
known in the Auchenorrhyncha (evidently 
also a paraphyletic taxon [e.g., Campbell et 
al. 1994, Sorensen et al. 1995, Hamilton 
1996]). Dolhng (1991) stated that all au- 
chenorrhynchans feed on vascular plants. 
Herein bryophagy in the auchenorrhynchan 
superfamily Fulgoroidea is reported. I sum- 
marize data on the seasonality and habits of 
the delphacid planthopper Javesella opaca 
(Beamer) on common hair-cap moss. Po- 
lytrichitm commune Hedw.. and record 
nymphs and adults from the moss Polytri- 
cluistrifiii cilpiiuim (Hedw.) G.L. Sm.: dis- 
cuss fluctuations in the numbers of plant- 
hoppers relative to the condition of host 
colonies (green vs. brown and drought 
stressed) and weather conditions: precipi- 
tation, temperature, and evaporation; and 
suggest that mosses represent an evolution- 
arily derived host association in the Del- 
phacidae. Seven new state records also are 
given for this seldom-collected species. 

This contribution is dedicated to my 
friend and colleague. Craig A. Stoops, who 
called my attention to a delphacid he col- 
lected from moss when we were in the field 
in June of 1996. He helped make field col- 
lections and laboratory observations before 
he left Clemson University in January 

Materials and Methods 

Study area.— Boggs" Rock (34°48.4'N. 
82°41.6'W) in northwestern South Carolina 
was the main study site. It is 2.1 km north 
of Liberty. Pickens County, and lies just 
west of the intersection of U.S. highway 
178 with state secondary road S-39-317 
(Quairy Road). This granite-gneiss flatrock 
ranges in elevation from 262 to 293 m 
(Knox 1974). Several plant species endemic 
to granite flatrocks in the southeastern Unit- 
ed States (e.g., McVaugh 1943, Shure 
1999) are found at Boggs' Rock, but be- 
cause permanent depression pools are ab- 
sent (Knox 1974), flatrock endemics re- 
stricted to such pools also are lacking. 

Host plants. — Polytrichiim commune 
(Musci: Polytrichales: Polytrichaceae) is a 
dark green, robust, perennial moss of wide 
distribution in the New and Old Worlds ex- 
cept in tropical areas. Its leafy stems, rigid 
and erect (acrocarpic), are connected by an 
extensive system of subterranean rhizomes. 
Stems of this tall moss can reach a height 
of 45 cm (Crum 1976, Crum and Anderson 
1981, Derda and Wyatt 1990). This dioe- 
cious moss reproduces asexually by vege- 
tati\e branching and propagation from plant 
fragments, with some populations being 
maintained entirely by vegetative reproduc- 
tion; it can also become established sexu- 
ally from spores (Leslie 1975, Derda and 
Wyatt 1990, Wyatt and Derda 1997). Found 
on various acidic substrates in disturbed 
and natural communities, P. commune often 
is abundant on granite flatrocks. It is an im- 
portant soil builder of the annual-perennial 
herb community of granite outcrops and a 
characteristic plant of that community. Its 
colonies or mats are persistent and often ex- 
tensive (Burbanck and Phillips 1983. Derda 
and Wyatt 1990. Quarterman et al. 1993). 

The moss variety on which the planthop- 
per J. opaca was collected at Boggs" Rock 
and other rock outcrops is P. commune van 
perigoniale (Michx.) Hampe. Plants of this 
variety are shorter and grow in denser tufts 
than those of the nominate variety (Crum 


and Anderson 1981 ). Of the many varieties 
of P. commune that have been described. 
peri^oniale has been the one most consis- 
tently recognized by authorities (Zouhair et 
al. 2000). Polytrichum commune van peri- 
goniale, however, is not genetically distinct 
from the nominate variety (Derda and Wy- 
att 1990). Overlapping in range with P. 
commune van commune, P. commune van 
perigoniale might represent only a "dry 
land phase" of a species that grows mainly 
in wet habitats (Crum and Anderson 1981). 
Derda (1998) considered P. commune var. 
perigoniale merely a phenotype of dry. ex- 
posed soils (cf. Zouhair et al. 2000). 

In the southern Appalachians, J. opaca 
was found on Polytrichum commune var. 
commune and Polytrichastrum alpinum. a 
polytrichaceous moss that also has been 
placed in Pogonatum and Polytrichum 
(Smith 1971. Crum and Anderson 1981. 
Smith Merrill 1992). This coarse species, 
usually 4-16 cm high with erect stems, is 
Holarctic and particularly widespread at 
northern latitudes (Crum and Anderson 

Field sampling. — After J. opaca was dis- 
covered at Boggs" Rock in June 1996. I sur- 
veyed other southeastern granite outcrops 
(flatrocks and monadnocks) for the pres- 
ence of the delphacid on Polytrichum spe- 
cies. The seasonality of J. opaca was fol- 
lowed at Boggs' Rock, the study areas cor- 
responding with granite outcrop divisions B 
and C in Knox's (1974) floristic study. 

I (or C.A. Stoops) visited Boggs' Rock 
seven more times at irregular intervals in 
1996 and collected planthoppers by placing 
a white enamel pan (28 X 40 cm) into a 
thick mat of the moss and using an axe han- 
dle to tap the overlying stems over the pan. 
Planthopper nymphs were field sorted by 
size and recorded as either early (I, II) or 
later (III-V) instars, and the sex of adults 
and the condition of the moss colonies were 
noted. The number of individuals of 7. opa- 
ca observed was not standardized but gen- 
erally was 10. Nearly all individuals were 
replaced on the moss, although voucher ma- 

terial of both nymphs and adults was col- 
lected and deposited in the Clemson Uni- 
versity Arthropod Collection (CUAC); 
adults also were deposited in the collection 
of S.W. Wilson. Central Missouri State Uni- 
versity. Warrensburg. 

Sampling at Boggs' Rock was conducted 
similarly in 1997 and 1998 but at more reg- 
ular intervals than in 1996. I recorded the 
stage of the first 10 individuals of J. opa- 
ca — as early or late instars and adult males 
or females — found on the moss in Knox's 
( 1974) outcrop division C, a colony of the 
host plant that became desiccated and 
brown without frequent rain. I also often 
recorded the stage of the first 10 individuals 
encountered in a supplemental sample from 
moss in a wetter area of the outcrop 
(Knox's division B). In 1997. after an initial 
collection in early January. I sampled week- 
ly or biweekly from early March to late Au- 
gust; I also sampled in mid-September and 
early November. In 1998. samples were tak- 
en weekly (occasionally biweekly or every 
three weeks) from late February to late July 
when no more planthoppers were being col- 
lected, presumably because of drought con- 
ditions. After that, I sampled six more times 
from late August to late November in an 
attempt to collect J. opaca. Six additional 
attempts to collect the planthopper at 
Boggs' Rock were made from early April 
1999 to late May 2002. 

Fluctuations in planthopper numbers. — 
Data from the two nearest weather-record- 
ing stations were used to better interpret the 
numbers of planthoppers observed relative 
to the condition of moss colonies. The 
weather station at Pickens (34°53'N, 
82°43'W) is about 10 km north of Boggs' 
Rock: the Clemson station (34°4rN. 
82°49'W) is about 18 km southeast of the 
sainple site. 

Laboratory observations. — The work 
with J. opaca in the laboratory involved 
placing nymphs of several instars in plastic 
petri dishes or snap-cap vials that contained 
the gametophyte (1-5 stems) of common 
hair-cap moss. The feeding behavior of 


nymphs, presence of exuviae, and number 
of clays of observation were noted, but the 
duration ol nymphal stadia was not deter- 

Jcivi'sella opuca (Beamer) 
(Fig. 1) 

Jdvcscllii opiua was described in the ge- 
nus Delpluicodes Fieber from Connecticut 
by Beamer (1948), who selected as holo- 
lype a male collected at Willimantic t)n 8 
August 1946. His type series also included 
adults taken from 3 August to 2 September 
1946, at Storrs, Conn. (14 6. 8 9): Ding- 
mans FeiTy, Pa. (6 cJ, 5 9); and Mountain 
Lake, Va. (1 6). The only additional pub- 
lished record is that of Maw et al. (2()()()) 
Uom Quebec. Canada; this delphacid was 
listed as a provincial record without a spe- 
cihc k)cality and was recognized as belong- 
ing to the genus Javesella (as ''Javasella") 
Fennah (Maw et al. 2000). According to 
K.G.A. Hamilton (personal communication, 
2002), the Quebec record of J. opaca is 
based on a total of 13 males and 3 females 
from Covey Hill (5 June) and Laniel (5 
June, 20-21 July). New state records, based 
on the present study (see Material exam- 
ined), are Alabama, Georgia, Massachu- 
setts, New York, North Carolina, South 
Carolina, and Tennessee. 

The transfer of opaca to Javesella by 
Maw et al. (2000), made in a checklist 
without discussion of this nomenclatural 
change, is the appriipriate generic place- 
ment (S.W. Wilson, personal communica- 
tion, 1996). Mainly on the basis of genitalic 
characters, the polyphyletic genus Delplui- 
codes should be restricted to about 10 Old 
World species (Asche 1985, Bartlett and 
Dietz 2()()()). 

Seasonal history. — Late instars of 7. opa- 
ca overwintered in mats of common hair- 
cap moss, hi contrast to the yellow or yel- 
k>w-brown nymphs found during spring 
and summer, the overwintered nymphs 
were darker and sometimes fuscous. The 
darker coloration, similar to that observed 
in certain other deiphacids (e.g., Wilson and 

McPherson 1981), was observed at Boggs" 
Rock by early November in 1996 and 1997 
(late instars collected in New York in mid- 
August and some early instars from Mas- 
sachusetts in early September also were 
dark). Adults first were seen in 1997 during 
sampling on 16 March and were present on 
29 March in 1998. In both years, males ap- 
peared first, but sex ratios usually became 
female biased within three weeks after 
adults began to appear. Overwintered fifth 
instars were last seen on 10 April in 1997 
(2 in sample of 10) and 5 April in 1998 (I 
of 20). 

First instars of the season's new genera- 
lion first were observed on 23 May in 1997 
and 12 May in 1998. Adults of this gener- 
ation, almost all males initially, first were 
seen in the 19 June sample in 1997 and the 
22 June sample in 1998. Several late instars 
(7 of 20) of the new generation were still 
present in 1997 on 20 July. Early instars of 
a partial (overwintering) generation were 
found on 4 August 1997. In 1998, appar- 
ently because of drought conditions, no 
nymphs or adults could be collected after 
early July. 

Nearly all the adults observed from June 
1996 to July 1998 were brachypters. Only 
five macropters were seen during the study, 
all between 10 April and 2 May 1997; the 
three that were collected were females. 
Adults were not found after mid-August, 
with only nymphs observed from then until 
mid- to late March of the following year. 

Fluctuations in planthopper numbers. — 
The numbers of J. opaca observed fluctu- 
ated widely during the period of study, de- 
pending mainly on the amount of precipi- 
tation at Boggs" Rock. Available moisture, 
in turn, affected the condition of the moss 
colonies inhabited by the planthopper. 

Despite the wide variation in the num- 
bers of planthoppers that could be beaten 
from Polytrichitnu at least 10 individuals 
usually could be found on each sample 
date; few, however, were found in July of 
1998. Precipitation that year was about 160 
cm at Pickens and about 156 cm at Clem- 


Fig. I. 
rrii liiini < 

Jiivcsc'Ua opcuii. (A) Females (brachypters) on lealy gametophytc 
iiiiiiiiinc: (B) dorsal and (C) lateral views of female. 

immon hair-cap moss, Pol\- 

son, figures that actually exceeded annual 
averages of 142 cm tor Pickens (1952- 
2001) and 134 cm tor Clemson (1930- 
2001). Looking only at total precipitation 
for 1998, fiowever, is misleading; after sub- 
stantial rains on 1 May (nearly 6 cm) and 
8 May (>3.6 cm), rainfall was scant in the 
weeks preceding the planthopper's disap- 
pearance. At Pickens, precipitation for the 
last half of May was only 1.4 cm; rainfall 
for June totaled 13.6 cm, with nearly half 
of the total falling on the month's 20th day. 
After 20 June, only 0.3 cm of rain fell dur- 
ing the next 30 days, a period when daily 
maximum temperatures substantially ex- 
ceeded normal highs. Temperatures were at 
least 32.2°C (90°F) from 21 June to 14 July. 

with nine consecutive days of 36. 1-36.7°C 
(97-98°F). Evaporation at Clemson (data 
are unavailable for Pickens) exceeded rain- 
fall by 15.2 cm for the 30 days following 
20 June. 

Only four adults and no nymphs were 
observed during regular sampling on 4 July 
1998; seven adults were seen when a sup- 
plemental sample was taken on 6 July. No 
additional individuals were found during 
sampling on seven dates from 24 July to 29 
November 1998. Two adult males were 
found in early April 1999, but J. <>i?aca was 
not observed at Boggs' Rock in late March, 
mid-May, and mid-August 2000; in late 
March and early November 2001; or in late 
May 2002. 



Habits. — Jcivesello opaca was found at 
Boggs" Rock in thictc mats of common hair- 
cap moss in marginal areas of rock out- 
crops. Moss colonies iiarboring the plant- 
hopper grew beneath shrubs and small 
trees, particularly Chinese privet (Liiiiis- 
rniiii sinciisc Lour.; Oleaceae) and eastern 
red-cedar (Jiiiiiperus virginiana L.; Cupres- 
saceae). The delphacid was not found in 
more exposed colonies of the moss. Even 
when nymphs and adults were most nu- 
merous, they were difficult to detect on 
mats of PolyUiclnim; only occasionally 
were a few individuals observed on the sur- 
face of P. commune mats. The largest num- 
bers of planthoppers — early instars only — 
were found on 8 June 1997. Javesella opa- 
ca had been difficult to find on 23 May 
1997 when moss colonies were brown (a 
total of 9 individuals was observed in both 
the regular and supplemental samples), but 
on 8 June after recent rainfall, more than .30 
first and second instars were found in each 
of three I -pan samples from now green ar- 
eas of moss. When nymphs were numerous, 
exuviae dislodged from moss also were ob- 
served during sampling. 

In the southern Appalachians, J. opaca 
was found at an altitude of about 1 . 1 30 m 
on Polytrichaslrum alpimim (Alleghany 
Co., NC). The moss was mostly obscured 
by various grasses. Populations of the plant- 
hopper also were found at about 1,685 m 
in mats of Polytriclnim commune van com- 
nnnie (Carter Co., TN). 

When nymphs were placed in petri dish- 
es or vials with sprigs of gametophyte from 
Polytriclnim commune, they quickly (<5 
min.) settled on the thicker stems and in- 
serted their stylets. While feeding, nymphs 
(n = 10+) remained motionless, except for 
an occasional twitching of the legs, and typ- 
ically stayed at one site for two hours or 
more. Only once was a cleai" drop of excre- 
tion or honeydew seen at the end of the 
anus. First instars were not reared to adult- 
hood, but preliminary observations suggest 
that J. opaca could be easily reared on moss 
in the laboratory. Nymphs of various instars 

were maintained in a petri dish on the same 
sprigs of P. commune for 16 days and 
nymphal exuviae were observed. A first in- 
star placed in a 4-dram vial with two moist- 
ened stems of moss molted iince and was 
still alive after 27 days. 

Material examined. — Adults from a lo- 
cality are listed first, followed by a listing 
of any nymphs; when nymphs at Boggs' 
Rock were recorded in the field as early (I, 
II) and later (III-V) instars. the numbers of 
early and later instars are separated by a 
hyphen. An asterisk beside the word 
nyniphis) indicates that voucher material is 
housed in the CUAC. Collections were 
made by the author (no collector indicated), 
C.A. Stoops (C.A.S.). both of us (C.A.S. & 
A.G.W.). or W.K. Reeves (W.K.R.) from 
Polytriclnim commune, except in Alleghany 
County, North Carolina, where the host was 
Polytriclutstrum alpiniim and in Pennsyl- 
vania where the host was identified as Po- 
lytriclnim sp. 

ALABAMA: Cherokee Co.. Rt. 35. SE 
of Fort Payne. 31 Aug. 1996. 2 adults (sex 
undetermined). 10 Apr. 1997 (I d, 9 9); 
Jackson Co., sandstone outcrop, Rt. 117, 
0.3 km NW of jet. Rt. 71. Flat Rock. 10 
Apr. 1997 (3 d. 4 9; 1 fifth instar). GEOR- 
GIA: Columbia Co.. Heggie's Rock, E of 
Appling. 5 Apr. 1997 (4 T, 1 1 2 ), 26 Mar. 
2002 ( 1 d; 1 fifth instar); Gwinnett Co.. No 
Business Creek granitic outcrop. 2.6 km 
SSE of Snellville, 7 July 1996 (1 c? ). 26 
Mar. 2002 (1 cJ, 4 9; 2 fifth instars). MAS- 
SACHUSETTS: Montague Sand Plains. 3.5 
km N of Montague. 5 Sept. 2002 (\ i , \ 
9; early instars). NEW YORK: Sullivan 
Co.. Rt. 42, Forestburg, 3 Sept. 2002 (8 first 
through third instars); Ulster Co., Sam's 
Point Dwarf Pine Ridge Preserve, NE of 
Cragsmoor. 13 Aug. 2000 (12 S. 10 9; 
nyniphs). NORTH CAROLINA: Alleghany 
Co., ca. 7 km ENE of Laurel Springs, 16 
June 2002 (4 d, 1 9; late instars); Swain 
Co., Great Smoky Mountains National 
Park, nr. Ravens Fork. 8 Jan. 2001. W.K.R. 
(late instars). PENNSYLVANIA: Potter 
Co.. NE of Keatina Summit. 18 May 1997. 


C.A.S. (3 late instars). SOUTH CAROLI- 
NA: Pickens Co.. Boggs" Rock. 2.1 km N 
of Liberty, 16-17 June 1996, C.A.S. & 
A.G.W. (adult.s, sex undetermined; 25 + 
nymphs); 22 June 1996, C.A.S. (adults; 
nymphs=^: 1-2); 6 July 1996, C.A.S. & A.G. 
W. (7 (5, 10 9; nymphs*:0-ll); 20 July 
1996, C.A.S. (adults; nymphs^^O-4); 18 
Aug. 1996 (5 adults, sex undetermined; 
nymphs:! 1-4); 29 Sept. 1996, C.A.S. & 
A.G.W. (nymphs:3-7); 13 Oct. 1996, 
C.A.S. & A.G.W. (nymphs: 10-0); 9 Nov. 
1996, C.A.S. & A.G.W. (nymphs: 10-0); 4 
Jan. 1997 (nymphs*:0-19); 7 Mar. 1997 
(nymphs*: 0-11); 16 Mar 1997, C.A.S. & 
A.G.W. (2 6: nymphs: 0-10); 21 Mar. 1997 
(3 adults, sex undetermined; nymphs: 0-7); 

4 Apr 1997 (4 cj, 4 9; nymphs*:0-2); 10 
Apr 1997 (3 (5, 5 9; nymphs; 0-2); 18 
Apr 1997 (4 S.b 9); 25 Apr 1997 (3 6. 
1 9); 2 May 1997(3 6,1 9); 10 May 1997 
(3 (5, 6 9; nymph:(3-l); 23 May 1997 (2 
cJ, 4 9; nymphs:2-0); 8 June 1997 
(nymphs*:7-4); 19 June 1997 (3 <5, 2 9 
nymphs*:0-9); 29 June 1997 (2 (5, 4 9 
nymphs*:l-3); 5 July 1997 (3 6: nymphs* 
2-5); 13 July 1997 (11 d, 3 9; nymphs* 
2-4); 20 July 1997 (10 d. 3 9; nymphs* 
2-5); 4 Aug. 1997 (6 d, 2 9; nymphs*: 11- 
1); 17 Aug. 1997 (nymphs*:8-l); 14 Sept. 

1997 (nymphs*:2-0); 2 Nov. 1997 
(nymphs: first through third instars): 28 
Feb. 1998 ( nymphs* :0-6); 15 Mar 1998 
(nymphs:0-10); 22 Mar 1998 (nymphs:0- 
10); 29 Mar 1998 (3 i: nymphs:0-7); 5 
Apr 1998 (7 i. 3 9); 12 Apr 1998 (5 S, 

5 9); 24 Apr 1998 (1 .5,9 9); 5 May 1998 
(1 c?, 2 9); 12 May 1998 (2 9; nyniphs:8- 
0); 18 May 1998 (nymphs: 10-0); 27 May 

1998 (1 9; nymphs:9-0); 6 June 1998 
(nymphs:8-12); 22 June 1998 (12 <5, 1 9; 
nymphs:l-6); 4 July 1998 (3 d, 1 9 ); 6 
July 1998 (6 d, 1 9 ); 4 Apr 1999 (2 d); 
York Co., granite flatrock. Clover, 20 Apr 
1997 (3 9; 1 fifth instar). TENNESSEE: 
Carter Co., Carver's Gap, Roan Mountain, 
27 October 2002 (late instars). 


When J. opaca was first collected from 
mats of Polytrichum in June of 1996, its 
presence on moss was considered inciden- 
tal. I was unaware of any moss-feeding ful- 
goroids, a group in which adults of many 
species, including delphacids, can be found 
on plants that do not support nymphal de- 
velopment (e.g., DuBose 1960, Wilson et 
al. 1994). Because seeds of various angio- 
sperms accumulate, germinate, and survive 
in mats of mosses (McVaugh 1943), even 
the planthopper nymphs collected from 
moss might have been associated with vas- 
cular plants, such as grasses or sedges, that 
were growing within the Polytrichiiiii col- 
onies. Delphacids develop mainly on mono- 
cots, especially grasses and sedges (Wilson 
et al. 1994). Javesella kilnuini (Van Duzee) 
and J. stall (Metcalf) are associated with an 
aberrant host, the vascular cryptogam 
(sphenophyte) horsetails, Equlsetwn spp. 
(Equisetaceae) (Strickland 1940, Ossian- 
nilsson 1978). 

A consistent collection of nymphs and 
adults of the delphacid from P. coinimine at 
Boggs" Rock and elsewhere, coupled with 
observations of its moss-feeding behavior 
in the laboratory, soon indicated that J. 
opaca was a true bryophage. It can be re- 
garded as a bryophilous species or a bry- 
obiont {sensit Gerson 1982) — that is, an an- 
imal associated exclusively with bryo- 
phytes. As such, J. opaca becomes the first 
known bryophagous fulgoroid and appar- 
ently also the first auchenorrhynchan 
known to develop on mosses. Somewhat 
similarly, the first known bryophagous spe- 
cies of the largest family of Heteroptera, the 
Miridae, only recently was discovered (Ya- 
sunaga 2000; see also Wheeler 2001). 

My collection of J. opaca in North Car- 
olina on Polytrichastriim alpinum might 
shed light on how Beamer ( 1948) was able 
to collect 22 adults of this planthopper at a 
locality in Connecticut and 1 1 adults in 
Pennsylvania without associating the spe- 
cies with mosses. When 1 collected the first 


specimens of J. opaca in North Carolina 
while sweeping grasses. I was unaware of 
the polytrichaceous moss growing under the 
grasses. Beamer probably would not have 
looked for delphacids in extensive mats of 
Polytrichum commune because no auchen- 
orrhynchans were then known from mosses. 
He more likely would have swept the plant- 
hopper from the erect stems of Polytrichas- 
triim alpinum (or another moss of similar 
growth habit) that grew beneath grasses, 
plants that are common hosts of Delphaci- 
dae. Delphacids should be more readily 
swept from a moss such as Polylrichostnim 
alpinum that grows in loose tufts than from 
one such as Polytrichum commune that has 
a more densely tufted (cespitose) growth 
habit. Even if Beamer had noticed the 
mosses while he swept grasses, he likeh 
v\ould not have associated the planthopper 
u ith a nonvascular plant. 

The Delphacidae appear not to have di- 
versified on mosses. Except for J. opaca. 
all other delphacids for which nymphal 
hosts are known develop on vascular plants. 
Fulgoroids and other homopterans that feed 
on plants other than angiosperms or on fun- 
gi likely are derived from angiosperm feed- 
ers (e.g., Wilson et al. 1994; see also Ham- 
ilton 1990). Even though J. opaca belongs 
to an "advanced" lineage (Stenocraninae- 
Plesiodelphacinae-Delphacinae) of the Del- 
phacidae and belongs to the most morpho- 
logically advanced tribe of the most ad- 
\anced delphacid subfamily (see Asche 
1985, 1990: Emeljanov 1995), moss feed- 
ing by this planthopper might represent an 
ancient or relictual host relationship for the 
family. More likely, however, is that in the 
Delphacidae the use of mosses reflects a 
one-time host shift and an evolutionarily 
derived host association. 

Bryophagy in the Delphacidae, a phlo- 
em-feeding group (e.g., O'Brien and Wil- 
son 1985. Denno et al. 1987, Wilson et al. 
1994, Watanabe and Kitagawa 2000), inight 
have been predicted for polytrichaceous 
mosses. Although mosses are considered 
nonvascular plants (Buck and Goffinet 

2000), the long-distance conducting sys- 
tems of certain members of the Polytricha- 
ceae are similar in structure and function to 
those of simple vascular plants (Tansley and 
Chick 1901, Thomas et al. 1988). Water 
moves in elements called hydroids; photo- 
synthate, principally sucrose, moves in lep- 
toids that are similar to sieve elements in 
the phloem of higher plants. Among mosses 
that have been studied, the conducting cells 
of Polytrichum commune, Polytrichastrum 
alpinum, and certain other polytrichaceous 
species are the most structurally complex 
(Collins and Oechel 1974, Behnke 1975, 
Scheirer 1980, Reinhart and Thomas 1981, 
Richardson 1981, Thomas et al. 1990). 

The moss-feeding mirid described as a 
new species by Yasunaga (2000) belongs to 
a group of heteropterans that are mainly 
cell-content feeders on mesophyll rather 
than vascular-tissue feeders (Wheeler 
2001). The mirid would not necessarily 
need to feed on a moss with a conducting 
system similar in structure to that of vas- 
cular plants. Javesella opaca. however, as a 
presumed phloem feeder, is likely restricted 
to polytrichaceous mosses with well-devel- 
oped conducting cells. It might be more 
host restricted than are many other bry- 
ophagous insects (Gerson 1982, Lawrey 
1987). In fact, most delphacids (74%) are 
recorded from a single plant genus (Wilson 
et al. 1994). 

Javesella opaca might feed on gameto- 
phores of common hair-cap moss in a man- 
ner similar to that of the aphid Myzodium 
modestum Hottes on the same moss species 
(Thomas and Lombard 1991). The plant- 
hopper, however, does not feed in aggre- 
gations as does the aphid and might not al- 
ter the translocation of sugars in internal 
conducting tissues of the moss, as Thomas 
and Lombard ( 1 99 1) demonstrated for the 
aphid. Occasional production of honeydew 
droplets by the sucrose-feeding M. modes- 
tum (Thomas and Lombard 1991) agrees 
with my observations on J. opaca, which 
presumably also feeds on photosynthate of 
common hair-cap moss. 



Javcsellci Dpiica has been collected main- 
ly on thick mats of Polytrichiim commune 
in both natural and disturbed communities. 
This planthopper might also be found on 
Polytrichastriim ohioense (Ren. & Card.) 
G.L. Sm. In fact. Knox (1974) recorded Po- 
lytrichastriim ohioense as the only polytri- 
chaceous moss at Boggs' Rock, but hi.s 
voucher material actually is Polytrichum 
commune (L.E. Anderson, personal com- 
munication. 2002). Both polytrichaceous 
species can be found on granite outcrops 
(e.g.. Costing and Anderson 1939, Mc- 
Vaugh 1943). but Polytrichum commune is 
a characteristic plant of granite flatrocks 
and apparently is the dominant hair-cap 
moss in granite outcrop communities in 
Georgia and elsewhere in the southeastern 
states (e.g.. Burbanck and Phillips 1983. 
Quarterman et al. 1993, Murdy and Carter 

Given the persistence of Polytrichum col- 
onies and the stability of granite flatrock 
communities, the observed low incidence 
{<3i%) of macroptery might have been ex- 
pected for J. opaca because dispersal might 
not be critical to the continuity of a plant- 
hopper population that lives in such a rel- 
atively stable or permanent habitat (e.g., 
Wagner and Liebherr 1992). From June 
1996 to July 1998, the delphacid typically 
was observed at Boggs" Rock in small 
numbers when rainfall was scant and its 
host was metabolically inactive, but it could 
be collected in greater numbers from still 
green colonies of the moss in wetter areas 
of the flatrock. The more consistent avail- 
ability of moisture in these wetter colonies 
apparently allowed egg hatch to continue, 
which sometimes led to asynchrony in the 
development of hopper populations in des- 
iccated compared to greener host patches. 

In 1998, a year of above-average precip- 
itation, the delphacid was not found after 
early July. Apparently the low and uneven 
distribution of rainfall in late spring and 
early summer, combined with consecutive 
days of unusually high temperatures and 
evaporation, severely reduced the densities 

of J. opaca in the two sample areas at 
Boggs" Rock. Adults and nymphs were no 
longer seen, even in the usual wetter moss 
colonies, which had become desiccated. Al- 
though temperatures within green mats of 
P. commune generally are lower than those 
at the surface, desiccated mats are marked 
by internal temperatures higher than those 
of the ambient air (Leslie 1975). Other 
groups of arthropods, such as collembolans 
and mites, were scarce in 1998 when moss 
colonies were brown and dry. In contrast to 
the planthopper, collembolans and mites re- 
appeared in numbers following abundant 
rainfall. That only two adults of J. opaca 
have been collected at Boggs" Rock since 
July 1998. even when moss was lush, sug- 
gests that its populations were substantially 
reduced during periods of low rainfall in the 
summer of 1998 rather than being found 
deeper in moss and. therefore, perhaps 
more difficult to dislodge from desiccated 
colonies of the host. 

In the southern Appalachians, mats of 
Polytrichum commune in high-elevation 
mesic communities, such as grassy balds, 
would be constantly moist and should be 
considered an even more permanent habitat 
than the periodically dry mats of the moss 
in granite outcrops. The degree of brachyp- 
tery in J. opaca might be studied in popu- 
lations occupying both permanent and less 
stable habitats. That this planthopper pri- 
marily ingests sucrose translocated in moss 
leptoids, as apparently is the case in several 
other moss-feeding hemipterans, requires 
verification. Also needed are surveys for 
additional moss species that might serve as 
hosts and studies of basic life-history pa- 
rameters such as fecundity, longevity, and 
duration of the immature stages (eggs were 
not found in the present study), as well as 
observations on mating and oviposition be- 
havior. In addition, the studies on nutrition- 
al ecology that Lawrey (1987) suggested be 
conducted on moss-feeding arthropods in 
general are appropriate to the research 
needs for J. opaca. 



The m(«s-t"eeding habits of J. opaca can 
be recorded because of the initial collection 
and curiosity of Craig Stoops, now an en- 
tomologist with the U.S. Navy. I also am 
grateful to S. W. Wilson (Department of Bi- 
ology, Central Missouri State University, 
Warrensburg) for identifying the delphacid 
and informing me of its correct generic 
placement, L. E. Anderson (Department of 
Botany, Duke University, Durham, N.C.) 
for identifying mosses, P. D. McMillan (De- 
partment of Biological Sciences, Clemson 
University) for preparing moss specimens 
and sending them to L. E. Anderson, W. K. 
Reeves (Department of Entomology, Clem- 
son University) for ci)llecting the planthop- 
per in Great Smoky Mountains National 
Park, K. G. A. Hamilton (Agriculture and 
Agri-Food Canada, Eastern Cereal and Oil- 
seed Research Centre, Ottawa) for provid- 
ing collection data for Canadian specimens 
of J. opaca, D. E. Linvill (Department ol 
Agricultural and Biological Engineering, 
Clemson University) for providing meteo- 
rological data, J. F. Stimmel (Bureau of 
Plant liuiuslry, Pennsylvania Department of 
Agriculture, Harrisburg) for photographing 
J. opaca. P. C. Huth (Mohonk Preserve, 
New Paltz, N.Y.) for facilitating access to 
Sam's Point Dwarf Pine Ridge Preserve, J. 
R. Allison (Georgia Department of Natural 
Resources, Social Circle) for accompanying 
me in the field at granite outcrops in Geor- 
gia, R. W. Brooks (Department of Ento- 
mology, University of Kansas, Lawrence) 
for attempting to locate R. H. Beamer's 
field notes, and P. H. Adler (Department of 
Entomology, Clemson University) for pro- 
viding comments that improved the manu- 
script. Two reviews secured by the editor — 
one submitted anonymously, the other by 
C. R. Bartlett (Department of Entomology 
& Applied Ecology, University of Dela- 
ware, Newark) — also improved the manu- 

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1()S(3). 2()()3, pp. 61 1-629 


Aubrey G. Scarbrough 

Department of Biological Sciences, Towson University, 8000 York Road. Baltimore. 
MD 21252. U.S.A. (e-mail: 

Abstract. — A new Afrotropical species group. Omiuatius jlcivipeniiis. including six spe- 
cies (O. aiigustus. n. .sp.. O. callidus, n. sp., O. ftavipeiinis. n. sp.. O. nigrantis. n. sp., 
O. pernecessarius, n. sp.. and O. perscieiitus, n. .sp.) are reported. The species group and 
species are diagnosed, described, and compared with similar species. Illustrations of the 
right wing of the male and terminalia of all species and a key to the species are provided. 
Comments on the distribution are included. 

Kev Words: 

Diptera. Asilidae. Omiiuitiiis Wiedemann, ftaripcimis species group, new 
species, Afrotropical 

Oininatiiis Wiedemann ( 1S21 ) is a large, 
widespread Oinmatiine (Diptera: Asilidae) 
genus in the Afrotropics that has received 
little attention until recently (Scarbrough 
and Marascia 1996, 2000; Scarbrough 
2002a). Eighty species (Oldroyd 1980) are 
assigned to this genus of which 46 are from 
Africa. However, preliminary studies indi- 
cate that a considerable number of unde- 
scribed species from Africa will require for- 
mal names and descriptions. Several addi- 
tional taxonomic problems exist within this 
genus that must be resolved before phylo- 
genetic relations can be accessed and com- 
prehensive keys to the African species can 
be constructed. Thus far the genus Einpliy- 
somerci Schiner (1996, 1999) and the sub- 
genus Metoiiimatius (Scarbrough and Mar- 
ascia 2000) have been revised. Further, we 
redescribed two species, designated types 
for O. tcnelliis van der Wulp and designated 
a replacement name (O. ahdelkiiriensis 
•Scarbrough) for O. tibialis Richardo (Scar- 
brough 2002a). The purpose of this paper 
is to report a new Afrotropical species 
group, O. fiavipennis. including six new 


The descriptive terminology follows 
McAlpine (1981). Dissections of the ter- 
minalia and preparation of illustrations of 
external and internal parts follow Scar- 
brough and Marascia (1999, 2000) and 
Scarbrough (2002b). Descriptions are com- 
posites of all specimens examined. In re- 
cording label data, each line is demarcated 
by a slash (/). Square brackets are used for 
additional information not present on la- 
bels. Ratios used in the descriptions are as 
follows: Head-face ratio [HFWR] = the 
greatest anterior distance across divided by 
the width of the face at the base of the an- 
tenna; flagellum width-length ratio [FWLR] 
= the length divided by the greatest width 
of the flagellum; cell m, width ratio 
[m,WRj = width at the basal third [widest] 
and apical two-thirds [narrowest] divided 
by the basal width; hind femoral width- 
length ratio [HFWLR] = the length of the 
hind femur divided by its greatest width. 
All measurements were made of pinned 
specimens. Structures in illustrations are la- 
beled only for the first species and are not 


repeated for comparable structures of the 
following species. Moiphological structures 
illustrated in different positions have a sin- 
gle scale line. 

Specimens used in this study are housed 
in the following Institutions: The Natural His- 
tory Museum. London (BMNH); California 
Academy of Sciences, San Francisco 
(CASC); Canadian National Collection. Ot- 
tawa (CNCI); Natal Museum of South Africa. 
Pietermaritzburg (NMSA); Musee Royal de 
L'Afrique Centrale, Tervuren (MRAC); Mu- 
seum National d" Histoire Naturelle, Paris 
(MNHN); Hope Entomological Collection, 
The University Museum. Oxford (OXUM), 
and The Natural History Museum of Zim- 
babwe. Bulawayo (NMZB). Acronyms of in- 
stitutions herein follow Amett et al. (1993). 


The Afrotropical Omniatiiis flavipeiiiiis 
species group 

Diagnosis. — Omnuitiiis flavipeiuiis spe- 
cies are readily recognized by the following 
combination of characters. 1 ) Abundant, 
long setae below the head and proboscis, 
length about as long as proboscis; long fla- 
gellum, 1.7-3.0 longer than wide; dorsal fa- 
cial setae usually long, Vi-Vi as long bris- 
tles. Anepimeral bristle present. 2) Wing 
(Figs. 1—6) mostly or entirely dense micro- 
trichose. Cell Tj long, narrowly triangular 
with base at or just beyond apex of cell d; 
vein R4 strongly arched basally, slightly 
concave or angled posteriorly before wing 
margin. 3) Middle and femora with 2 ven- 
tral rows of stout bristles, those of hind fe- 
mur extend from base to or near apex. 4) 
Apical sternites of abdomen with sparse 
erect bristles except in nigrantis and per- 
scientus. 5) Epandrium (Figs. 7, 17, 27, 47, 
57) usually narrow except wide apically in 
nigrantis. apex pointed, curved dorsally in 
nigrantis (Fig. 37). 6) Aedeagus (Figs. 10. 
20, 30, 40, 50, 60) tubular, flattened dor- 
soventrally, with wide base and apex, nar- 
rowed medially; ventral keel and flared 
apex present Figs. 12, 22, 32, 42, 52, 62). 

7) Spermatheca (Figs. 15, 25, 35, 55) long 
and tubular, with wide base and narrow 
apex, usually recurved or looped, except 
short, carrot-shaped in nigrantis (Fig. 45). 

Remarks. — Whereas most Ommatius 
species are superficial similar externally, 
the structure of the terminalia is the primary 
means of identification of species and spe- 
cies groups. The long setation of the head, 
presence of two rows of ventral bristles be- 
low the middle and hind femora, length and 
shape of cell r^. and shape of the aedeagus 
readily separate the flavipennis species 
group from all known Afrotropical conge- 
ners. The length and shape of cell r4 of O. 
longipennis (Linder 1955) is similar to all 
members of the O. flavipennis but differs in 
the other characters listed above. It is fur- 
ther similar to O. perscientus in the the nar- 
row, mostly sparse setose face, thin brown 
tomentose mesonotum, and brown veins 
and microtrichia. However, Ommatius per- 
scientus differs significantly from O. lon- 
gipennis in the combined characters of the 
terminalia. Species of the subgenus Metom- 
matius Hull (Scarbrough and Marascia 
2000) and O. tenelhis van der Wulp (Scar- 
brough 2002a) also differ in the presence of 
glabrous stripes or spots on the mesonotum, 
wing venation, and characters of the ter- 
minalia. The former differs further in the 
absence of marginal scutellar bristles and 
rows of ventral bristles below the middle 
and hind femora. 

Distribution. — The species group is 
widespread in middle Africa, ranging from 
southern Ethiopia westward to Nigeria and 
southward to Angola and Zimbabwe. How- 
ever, five species (O. callichts. O. flavipen- 
nis. O. nigrantis, and O. pernecessarius. O. 
perscientus) are concentrated in the eastern 
half of the continent although O. perneces- 
sarius ranges westward to Nigeria. Omma- 
tius angustus is known only from the Lu- 
bango and Huambo regions of Angola. All 
species were captured [where data given] 
between 1100 and 2100 meters in savanne 
and low mountain forests. 


Fig. 1-6. Oinimiliiis spp.. right wing. 1. O. uiiiiiism 
pemecessariiis. 6, O. perscientus. R4 = radial vein, rj 
m, = medial cell, m, = medial cell, CuA| + M, vein. 

radial cell, r-m 

O. ticiyipt'iinis. 4. O, nii^niiili.s 
= radial-medial crossvein. d = 

Key to the Afrotropical Ommat/us 
FLwiPENNis Species Group 

Mesonotum dorsally brown tomentose. dif- 
fused parumedial stripes sometimes present; 
ocellar tubercle with 4-5 long setae: postocular 
bristles long. thin, strongly proclinate. apex of 
bristles well forward of posterior margin of 
eye; sternites of female with only thin setalike 


Mesonotum dorsally with two. narrow, con- 
trastingly brown tomentose paramedial stripes; 
ocellar tubercle with only 2 long setae; posto- 
cular bristles thick and short, tips usually just 
forward of eye margin posteriorly, well behind 
ocellar tubercle; body yellow tomentose: apical 
sternites of female with few to several yellow 


Fore and middle temora black, narrow base 
sometimes yellow; abdomen unusually flat, 
clavate apically: mesonotum posteriorly, scu- 
tellum entirely, and basal segments of abdomen 
gray tomentose: wing and terminalia as in Figs. 

4. .^7 — 16); Democratic Republic of Congo. 
Tanzania niifiantii. n. sp. 

- Fore and middle femora yellow ventrally and 
posteriorly, dark brown dorsally and anteriorly: 
abdomen round in cross-section, neither flat 
nor spatulate: thorax and abdomen dorsally 
brown tomentose; wing and terminalia as in 
Figs. 6. 57-62); Zambia. Zimbabwe 

persclcnrii.s. n. sp. 

-'!. Mesonotum with most lateral bristles yellow; 
gonostylus with a prominent concavity poste- 
riorly (Figs. II. 21. 31); aedeagus narrow api- 
cally. base much wider in dorsal view, mid- 
basal "shoulders" absent (Figs. 10. 20, 30), ven- 
tral keel variable (Figs. 12, 22, 32); sternite 8 
of female with 1-4 prominent bristles apically, 
preapical, medioapical tubercle ab.sent (Figs. 
14. 24. 34); genital fork with narrow, constrict- 
ed base (Figs. 26, 36, 46) 4 

- Mesonotum usually with lateral bristles black; 
gonostylus without prominent concavity pos- 
teriorly (Fig. 51); aedeagus wide in dorsal 


Figs. 7-12. Ommatiiis angustus. male terminalia. 7-9. Lateral, dorsal, and ventral views. 10. Aedeagus. dorsal 
view. 11. Lett gonostylus. 12. Aedeagus. lateral view. AE = aedeagus. EP = epandrium, CE = cercus. GS = 
gonostylus. GC = gonocoxite. HY = hypandrium. 

view, wide mid-basal "shoulders" (Fig. 50) and 
strong ventral keel present (Fig. 52); stemite 8 
of female without prominent bristles apically. 
a prominent, preapical. medioapical tubercle 
present (Fig. 54); spermatheca with a preapical 
constriction (Fig. 55); genital fork with wide 
base (Fig. 56); Burundi. Democratic Republic 
of Congo, Kenya, Malawi, Nigeria, Tanzania, 

Uganda, Zambia peniecessorius. n. sp. 

Gonostylus angular (Figs. 27, 31); gonocoxite 
long, capitate apically in lateral view (Fig. 27); 
aedeagus unusually narrow in dorsal view, base 
abruptly wide (Fig. 30 1, prominent ventral keel 

present (Fig. 32); apical margin of stemite 8 of 
female evenly contoured, hornlike process ab- 
sent. 2 flat bristles present (Fig. 34); sperma- 
theca as in Fig. 35; Democratic Republic of 
Congo. Zambia. Zimbabwe . . . flavipennis. n. sp. 
- Gonostylus curved forward (Figs. 11,21); gon- 
ocoxite shorter, sides even apically (Figs. 7, 9, 
17, 19); aedeagus triangular in dorsal view, 
sides gradually widens posteriorly (Fig. 10, 
20), keel as in Figs. 12, 22); apical margin of 
stemite 8 of female hornlike, each 
bearing prominent bristles (Figs. 14, 24); sper- 
matheca as in Fills. 15, 25 5 


Figs. 13-16. Oininaliii.s 
Genital fork. Abbr. TG8 = 

aiii;iisiii.\. female terminalia. 13-14. Dorsal and ventral viev 
tergite 8. TG9 = tergite 9, CE = cercus, ST8 = sternite 8. 

15. Sperniatheca. 16. 

Gonostylus narrow, only slightly arched forward 
(Figs. 7. 1 1 ) aedeagus wide apically, apex oval, 
about as wide dorsally as laterally (Fig. 10), a 
moderate ventral keel present (Fig. 12); tergite 
8 of female with lateral bristles oval, sides grad- 
ually narrowed from base to apex (Fig. 13); ster- 
nite 8 with deep, medioapical notch between 
prominent hornlike processes (Fig. 14); apex of 
sperniatheca slightly swollen (Fig. 10); genital 

fork as in Fig. 1 1 ; Angola aiigusliis. n. sp. 

Gonostylus wide, strongly arched forward (Fig. 
2 1 ); aedeagus narrow apically. apex much wid- 
er dorsally than laterally, a low ventral keel 
present (Figs. 20-22); tergite 8 of female with 
base of lateral bristles flat and wide, side 
abruptly narrows beyond (Figs. 23. 24); ster- 
nite 8 only slightly recessed, not strongly 
notched between short, hornlike processes 

(Fig. 24); apex of .spermatheca not noticeable 
swollen (Fig. 25); genital fork as in Fig. 26; 
Democratic Republic of Congo . . callidus. n. sp. 

Tax A 

Ommatiiis angiistus Scarbrough, 
new species 

(Figs. 1. 7-16) 

Male.— Body 15.9-17.0 mm; wing 12.3- 
12.5 mm. Head: Face yellow tomentose. 
dorsal half with black vestiture, dorsal setae 
abundant. Vi-Vi as long as bristles; FHWR 
1.0;7.1-1.0:7.5. Antenna, frons. and ocellar 
tubercle black setose. Flagellum about as 
long as scape and pedicel combined; FWLR 


1.0:2.0-1.0:3.2. Frons light-brown tomen- 
tose, setae longer than scape. Ocellar tuber- 
cle with 2 long setae, length siibequal to 
scape and pedicel combined. Dorsal posto- 
cular bristles short, mostly black, longest 
seta with tip just forward of eye posteriorly. 

Thorax: Mesonotum with 2 wide, 
brown, tomentose. paramedial stripes, fused 
anteriorly, and 2 lateral spots; 4 dorsocen- 
tral bristles present, most black. 1-2 yel- 
low; 5 lateral bristles yellow, 1 notopleural 
bristle black. Scutellum with scattered, 
long, yellow setae and 2 yellow marginal 
bristles; weak preapical groove present. 
Anepimeral bristle yellow. 

Winfi (Fig. 1 ): Entirely yellow to yellow- 
ish-brown microtrichose. Anterior radial 
vein mostly yellow, brown apically. Cell 
m.WR 1.0:1.8:1.6-1.0:1.9:1.6. 

Lesis: Femora mostly yellow with yellow 
bristles; fore and middle femora mostly 
black anteriorly, nanow apex black poste- 
riorly; hind femur with narrow apex and 
narrow anterior stripe black, stripe present 
on apical Vi-Vr. apical anteroventral bristle 
sometimes black. Middle femur with 4-5 
ventral bristles in each row. HFWLR 1 .0: 
4.3-1.0:5.1. Nanow apex of middle and 
hind tibiae brown. Tarsi yellow setose; bas- 
al 3 tarsomeres yellow, narrow apices 
sometimes brown; fore and middle tarsi 
with numerous yellow bristles laterally; 
hind tarsus with only 1 yellow bristle. 

Abdomen: Mostly yellow to yellowish- 
gray tomentose, yellow setose; tergites 5-7 
sparse, brown tomentose dorsally, black se- 
tose; apical corner of tergites 2-8 with 2-4 
stout, usually yellow, bristles; sternites 2-7 
usually with 1-4 stout yellow bristles api- 

Tcrminalia: Black (Figs. 7-12). Gonos- 
tylus naiTow in lateral view, concave pos- 
teriorly. Aedeagus triangular dorsally, basal 
corners wide; wide in lateral view; low ven- 
tral keel present. 

Female. — Differs as follows. Body 14.5- 
19.5 mm; wing 10.8-12.5 mm; FHWR 1.0: 
6.6-1.0:6.7; FWLR 1.0:2.8-1.0:2.9; m,WR 
1.0:2.1:1.9-1.0:2.5:2.4: HFWLR 1.0:5.1- 

1 .0:5.6. Head: Occiput with 2-3 dorsal pos- 
tocular bristles yellow. Legs: Fore tarsus 
with yellow bristles present on outer side 
only; hind femur with setigerous bristles 
only on basal Vi: hind tarsus with only black 
bristles. Abdomen: Most tergites laterally 
and sternites apically with 2-4 stout, bris- 
tles, latter mostly yellow. Terminalia (Figs. 
13-16): Sternite 8 mostly black, narrow 
apex red; apical margin strongly produced 
as hornlike processes, apex with 3-4 stout, 
black bristles. 

Specimens examined. — Holotype S, 
ANGOLA: (A6)/Tundavala/9 mis NW Sa/ 
da Bandeira [= Lubango, Hufla Plateau]/ 
23.vii.1972/Southern/African Exp./B. M. 
1972-1 (BMNH). Paratypes: 1 <5. 3 9, 
same data as holotype (BMNH); 1 9. 
(A22), Nova Lisboa [= Huambo, Bie Pla- 
teau]/5-7.viii.l972/Southern/African Exp./ 
B. M. 1972-1 (BMNH). 

Etymology. — Latin aiigii.snis for nairow 
passage, referring to the gorge at Tunda- 

Distribution. — Montane species [eleva- 
tion 1676 m] capture in July and August on 
the Hin'la and Bie Plateaus of west central 
and si>uth central Angola. 

Remarks. — The long narrow flagellum, 
yellow thoracic and femoral bristles, and 
combined characters of the terminalia (Figs. 
7-16) especially the gonostylus and aedea- 
gus of the male and segment 8 and sper- 
matheca of the female distinguish O. an- 
giisnt\ from congeners. 

Oiumatiiis callidiis Scarbrough, 
new species 

(Figs. 2. 17-26) 

Male.— Body 19.3-20.3 mm; wing 12.5- 
12.7 mm. Head: Face with yellow vestiture, 
setae on dorsal % short, Vi as long as bris- 
tles; HFWR 1.0:6.7-1.0:8.0. Antenna, 
frons. and ocellar tubercle black setose; 
pedical rarely scattered yellow setose. Fla- 
gellum less than scape and pedicel com- 
bined; FWLR 1.0:1.9-1.0:2.2. Frons ven- 
trally with setae slightly longer than scape. 
Ocellar tubercle with 2 setae about % as 


Pigs. 17-22. Oinniauiis iitlticlii 
dorsul view. 21. Lett gonoslylus. 2'. 

male terminulia. l7-|y. Lateral. 
Aedeagus, lateral view. 

dcirsal. and venlral view.s. 20. Aedeagus. 

long as scape and pedicel comhined. Pos- 
tocular bristles yellow, rarely with 1-2 
black; bristles short, longest with tip just 
forward of eye posteriorly. 

Thorax: Mesonotuin with 2 wide, 
brown, tomentose, paramedial stripes, fused 
anteriorly, and 2 lateral spots; bristles most- 
ly yellow; 7-8 dorsocentral bristles present; 
2-3 dorsocentral and I notopleural bristles 
usually black. .Scutellum with long, yellow 

setae and yellow marginal bristles; preapi- 
cal groove absent. Ancpimeral bristle yel- 

Wing (Fig. 2); Entirely yellow to brown- 
ish-yellow micrt)trichose. Anterior radial 
veins yellow basally. Cell niiWR 1.0:2.3: 

Legs: Femora mostly yellow, narrt)w 
apices brown; ventral bristles usually yel- 
low; fore and middle femora mostly brown 


Figs. 23-26. Oininatiii. 
Genital fork. 

callidits. teniale termiiialia. 23-24. Dorsal and ventral views. 25. Spermatheca. 26. 

anteriorly, hind femur anteriorly with nar- 
row brown sometimes on apical V3-V2. Mid- 
dle femur with 2-3 black anteroventral bris- 
tles. HFWLR 1.0:4.1-1.0:4.3. Narrow apex 
of 2, sometimes 3, tibiae brown. Tarsi most- 
ly yellow setose; basal 3 tarsomeres yellow, 
narrow apices often brown; each tarsus with 
several yellow bristles. 

Tenninalia (Figs. 17-22): Gonostylus 
wide, strongly arched forward, wide con- 
cavity posteriorly. Aedeagus with wide 
apex and base in dorsal view, triangular, 
narrow in lateral view; ventral keel weak. 

Female. — Body 17.3-18.3 mm; wing 
15.0 mm; HFWR 1.0:6.2-1.0:7.3; FWLR 

1.0:2.3-1.0:2.6; m,WR 1.0:2.3:3.0-1.0:3.0: 
2.6; HFWLR 1.0:4.9-1.0:5.3. Thorax: Two 
to 4 dorsocentral and 2 notopleural bristles 
sometimes black. Legs: Fore and middle 
femora yellow, only naiTOw apex black; 
middle femur with 1 anterior and 2 anter- 
oventral bristles black. Abdomen: Lateroap- 
ical margin of tergites and scattered on 
most sternites with yellow bristles. Tennin- 
alia (Figs. 23-26): Sternite 8 red, apical 
margin produced medially, short processes 
with 1-2 black bristles apically, each pro- 
cess separated medially by shallow concav- 
ity bearing a divided tubercle. 

Specimens examined. — Holotype S. 



Musee du Congo/Lulua Riv. Luele/1928/Dr. 
Walker (MRAC). Paratypes: 3 6.4 $, 
same data as holotype (MRAC). 

Etymology. — Latin callidus, meaning 
skillful, refers to the hunting ability of asi- 

Distribution. — Captured along the Lulua 
Rivers in southeastern Democratic Republic 
of Congo. 

Remarks. — The short facial, frontal, and 
ocellar setae, the yellow postocular and me- 
sonotal bristles, and combined characters of 
the terminalia (Figs. 17-26). especially the 
gonostylus and aedeagus of the male and 
segment 8 and spermatheca of the female 
distinguish O. ciillichis from congeners. 

Ominatius flavipeitnis Scarbrough, 
new species 

(Figs. 3. 27-36) 

Male.— Body 13.9-17.3 mm; wing 9.8- 
11.2 mm. Head: Face with yellow and 
black setae dorsally. setae about Vi as long 
as bristles; FHWR 1.0:5.9-1.0:7.0. Anten- 
na, frons. and ocellar tubercle black setose. 
Flagellum as long as scape and pedicel 
combined; FWLR 1.0:2.7-1.0:3.0. Frons 
brownish yellow to red-brown tomentose. 
Ocellar tubercle with longest setae about as 
long as scape and pedicel combined. Occi- 
put with black postocular bristles usually 
extends to dorsolateral or midlateral margin 
of eye; dorsal bristles with apices just be- 
yond posterior margin of eye. 

Thorax: Mesonotum dorsally largely 
brown tomentose. paramedial stripes and 
lateral spots present in best specimens; 5 
lateral and 3-6 dorsocentral bristles present, 
most black, often 2-3 dorsocentral and 2- 
3 lateral bristles yellow. Scutellum with 
scattered, long setae and yellow marginal 
bristles; shallow preapical groove present. 
Anepimeral bristles yellow. 

Wing (Fig. 3): Entirely yellow to brown- 
ish-yellow microtrichose. Anterior 2-3 ra- 
dial veins yellow basally. Cell niiWR 1.0: 

Legs: Fore and middle femora dark 

brown anteriorly and narrow apices poste- 
riorly, yellow otherwise. Middle femur with 
4 black bristles. 1 anteriorly and 3 antero- 
ventrally; posteroventral bristles yellow. 
Hind femur yellow with narrow apex black, 
black variable anteriorly as a narrow or 
wide streak, often mostly black anteriorly; 
ventral bristles yellow; HFWLR 1.0:4.6- 
1.0:5.4. Tarsi mostly yellow setose; basal 
2-3 tarsomeres light, usually yellow with 
apex brown; each tarsus with 5-12 yellow 

Abdomen: Dark brown with yellow to 
yellowish-gray tomentum and yellow ves- 
liture; each tergite dorsally with brown to- 
mentum. light brown on basal segments, 
grading to dark tomentum on apical ter- 
gites. Apical margins of tergites 2-7 later- 
ally with 2 yellow bristles; apical corner of 
sternites 3 and 4 with 1-2 thick yellow bris- 
tles, remaining sternites with bristly setae. 

Terminalia (Figs. 27-32): Gonostylus 
strongly angular, L-shaped, deep concavity 
posteriorly. Gonocoxite apically capitate. 
Aedeagus narrow in dorsal view, base 
abruptly wide; wide in lateral view; prom- 
inently ventral keel present. 

Female. — Differs from males as follows. 
Face: Dorsal Vz-Vj, black setose; FWLR 1.0: 
3.1. Legs: Middle femur with only yellow 
bristles. Hind femur often with only narrow 
apex black. Basal 3 tarsomeres of fore and 
middle tarsi and basal 2 of hind tarsus 
mostly yellow. Abdomen: Apical margin of 
tergites 2 and 3 laterally with 3-4 and ter- 
gites 4-7 with 2 yellow bristles; tergite 8 
with 3 black bristles. Sternites 3-7 with 
several, scattered, yellow bristles. Termin- 
alia (Figs. 33-36): Sternite 8 with apical 
margin slightly produced medially. 2 flat, 
black bristles present mediolaterally. 

Species examined. — Holotype S. ZAM- 
BIA: Sakeji River [1400 m. elev.]/Ikelenge/ 
N. Mwinilungo/Nat Mus Bulawayo/13- 
14.iv.l972/Pinley-de Moor Exp. (NMZB). 
CONGO: 1 cJ. 2 9. Congo Beige: RN.U./ 
Mbuywe-Bala (1.750 m)/25-3 1 .iii. 1948/ 
Mis. G. E de Witte (MRAC). ZAMBIA: 2 


Figs. 27-32. Oinniatiiis flayipennis. male terniinalia. 27-29. Lateral, dorsal, and ■ 
dorsal view. 31. LcCl aonoslylus. 32. Aedeaaus. lateral view. 

entral views. 30. Aedeagus 

d, 1 9, same data as holotype; 2 6 , Mrushi 
R./80 m E of Kapin/7.iii.l969/Pinhey-de 
Moor Exp (NMZB); 1 d, 35 mi S of Ndola/ 
30.i.l965/Nat Museum S Rhodesia 
(NMZB). ZIMBABWE: 1 cJ. 1 2. Croco- 
dile Pool Farm Blanket/Nat Museum S 
Rhodesia/15. X!i.l977/D.K.B. Wheeler 

Etymology. — Latin, fiavipennis, combi- 

nation of flavus for yellow and pennis for 
wing, referring to the color of the wing. 

Distribution. — Captured from December 
through April at altitudes between 1400- 
1750 meters along the river basin Plateaus 
of southern Democratic Republic of Congo, 
Zambia, and Zimbabwe. 

Remarks. — The nanow fiagellum, black 
setae on the dorsal half of the face, and 


Figs. .^.^-.^(i. Onumuiiis fiavi/n 
36. Genital fork. 

female terminalia. .^.^-34. Dorsal and ventral views. 35. .Spermatheea. 

combined characters of the terminaHa (Figs. 
27-36), especially the wide aedeagus with 
a inidbasal constriction and prominent ven- 
tral keel, the short angular gonostylus. the 
capitate gonocoxite of the male and char- 
acters of segment 8 and spermatheea of the 
female distinguish O. fiavipennis from con- 

Ommatius nigraiitis Scarbrough, 
new species 

(Figs. 4, 37-46) 

Male. — Black body with unusually long, 
yellow and black setae, black femora, and 
a Hat. spatulate abdomen. Body 11.7-12.9 

mm: 7.8-10.0 mm. Head: Face entirely 
with long vestiture of uniform thickness, 
thick bristles absent; vestiture ventrally 
more dense, about as long as proboscis, and 
yellow; that on dorsal % of face black; 
HFWR 1.0:6.2-1.0:6.5. Antenna, frons, and 
ocellar tubercle black setose. Flagellum 
subequal to scape and pedicel combined; 
scape with long ventral setae extending be- 
yond base of flagellum; FWLR 1.0:2.1-1.0: 
2.2. Frons yellow brown to brown tomen- 
tose; setae as long as scape and pedicel 
combined. Ocellar tubercle with several 
long setae, apex of longer seta well beyond 
base of flagellum. Numerous black posto- 



cular bristles present, bristles extending to 
or beyond midlateral margin of eye; bristles 
thin and strongly proclinate with apex of 
longest dorsal bristle at or just before ocel- 
lar tubercle. 

Thorax: Mostly yellow to yellowish- 
gray tomentose. Mesonotum with wide, 
brown tomentose stripe medially and 2 
spots laterally, paramedial lines indistinct: 
prescutellum broadly gray tomentose; ves- 
titure mostly black and thin, anterior setae 
about as long as pedicel and scape com- 
bined; 9-10 long, setalike dorsocentral bris- 
tles present; 5-6 thick lateral bristles pre- 
sent. 1 often yellow. Scutellum with scat- 
tered black or yellow setae and 2 thin black 
bristles; longest setae as long as marginal 
bristles. Anepisternum dorsally with scat- 
tered black setae. Anepimeral bristle yel- 

Wing (Fig. 4): Entirely brownish yellow 
microtrichose. Veins brown. Cell m, WR 

Legs: Femora usually mostly dark brown 
or black with naiTOw bases yellow, rarely 
entirely dark; most bristles and setae yel- 
lowish to white; long ventral setae of fore 
and middle femora sometimes black; 
HFWLR 1.0:5.6-1.0:5.8. Hind tibia with 
narrow apex black. Tarsi yellow setose; 
basal 2 tarsomeres yellow, narrow apices 
brown; bristles mostly black. 1—2 yellow 
fore tarsal bristles usually present. 

Abdomen: Spatulate, junction of seg- 
ments 2-3 % as wide as segments 6—7; ter- 
gites 1-4, tergite 5 narrow base and sides, 
and most sternites with thin gray tomentum. 
apical tergites shiny brown or black with 
purple highlights. Setae mostly yellow, long 
laterally with sparse black or yellow bristles 
on most tergites; tergites 1-3 entirely yel- 
low setose, tergite 4 yellow setose basally 
and black apically, tergites 5-8 entirely 
black setose. 

Tenninalia (Figs. 37—42): Narrow, about 
Vi as wide as segment 6. Epandrium with 
apex flared, about as wide as base, small 
digitate ventroapical process present. Gon- 
ostylus narrowed apically, posterior con- 

cavity absent. Aedeagus narrow apically, 
sigmoid, apex flared. 

Female. — Differs as follows. Body 1 1.2- 
13.1 mm: wing 11.0-11.3 mm; FHWR 1.0: 
6.2-1.0:6.3; FWLR 1.0:2.2-1.0:2.7: m,WR 
1.0:2.4:2.0-1.0:2.3:2.1; HFWLR 1.0:5.3- 
1.0:5.4. Abdomen: Only slightly constricted 
at junction of segment 2-3, latter about % 
as wide as segments 6-7; tergites dorsally 
mostly black setose; dull gray tomentum 
gradually thinning apically, apical segments 
bare; sternites with only scattered yellow 
setae and sparse, thin bristles laterally. Ter- 
minalia (Figs. 43-46): Tergite 8 with only 
thin bristles. Sternite 8 evenly contoured 
apically. Spermatheca short, straight, carrot- 
shaped. Genital fork with apex of arms cap- 

Specimen examined. — Holotype 6, 
TANZANIA: Iwawa/15 mis. N. W. of/ 
Mwakete/7000 ft/21. i. 52 WP (BMNH). 
CONGO: 1 cJ, Congo beige: RN. U., Mu- 
kana (Lusanga) (1,810 m.)/6.iii.l948/Mis. 
G. E de Witte. 1352a (MRAC): 2 9 [1 
without tip of abdomen]/Congo beige: P.N. 
U./Mukana (1.810 m.)/4.iii. 1948/22. 
23.iv.1949/Mis. G. F de Witte. 1389a/ 
2550a (MRAC); 1 6. Congo beige: RN. U./ 
Kabwekanon (1,815 m.)/6.iii.l948/Mis. G. 
E de Witte. 1367a (MRAC); 2 9, Congo 
beige: P.N. U. /Lusanga (pres Mukana) 
(1,810 m.)/ G. E de Witte. 
80-81 (MRAC). 

Etymology. — Latin nigrantis for black or 
dusky, referring to the color of the legs. 

Distribution. — Captured in montane for- 
ests (1810 to 2121 m. elev.) during March 
and April in western Tanzania and eastern 
Democratic Republic of Congo. 

Remarks. — The mostly black femora 
with yellow bristles; the unusually long, 
black setae of the head: gray tomentum of 
the thorax and basal abdominal segments; 
the flat, spatulate abdomen; and the com- 
bined characters of the terminalia (Figs. 
37-46) distinguish O. nigrantis from con- 
geners. The unusually wide, dorsally ori- 
ented apex of the epandrium; narrow gon- 


Figs. 37 — 1.2. Ommatius nigranris. male terminalia. 37-39. Lateral, dorsal, and ventral views. 40. Aedeagus, 
dor.sal view. 41, Left gonostylus. 42, Aedeagus, lateral view. 

ostylus: and sigmoid aedeagus further dis- 
tinguishes the male. Absence of scattered, 
stout bristles on the abdomen; a short, car- 
rot-shaped spermatheca; and capitate arms 
of the genital fork further distinguish the 

Ommatius pernecessarius Scarbrough, 
new species 

(Figs. 5. 47-56) 

Male. — Body 11.0-14.4 mm; wing 9.6- 
10.5 mm. HeucI: Dorsal % of face with 


Figs. 43 — 16. Oiiiimiliu: 
Genital fork. 

nifininlis. t'eniaie lemiinalia. 4.1 — 14, Dorsal and ventral views. 4.'i. Spermatheca. 46. 

sparse black or mixed black and yellow se- 
tae, about Vi as long as 6 black bristles; 
FHWR 1.0:6.0-1.0:7.0. Antenna, frons, and 
ocellar tubercle black setose. Flagellum 
slighter shorter than scape and pedicel com- 
bined; FWLR 1.0:1.7-1.0:2.3. Frons with 
setae as long as scape. Ocellar tubercle with 
setae slightly shorter than scape and pedicel 
combined. Occiput dorsally with 6-8 black 
postocular bristles; apex of longest just be- 
yond posterior margin of eye. 

Thorax: Mesonotum with 2 wide, 
brown, tomentose, paramedial stripes, fused 
anteriorly, and 2 lateral spots; 5 lateral and 
5-6 dorsocentral black bristles, 1-2 rarely 
yellow. Scutellum with scattered yellow se- 
tae and 2 black marginal bristles; preapical 
groove absent. Anepimeral bristle yellow. 

Wing (Fig. 5): Dense yellow microtri- 
chose; radial veins brownish yellow. Cell 
m,WR 1.0:2.3:2.0-1.0:5.0:4.5. 

Legs: Fore and middle femora anteriorly 

and narrow apices posteriorly black, other- 
wise yellow; hind femur variable, usually 
mostly yellow with narrow anterior streak 
and narrow apex black; all femora some- 
times black anteriorly with naiTOw base yel- 
low. Middle femur usually with only black 
bristles; 1-3 short black bristles usually 
present posteroventrally. Hind femur usu- 
ally with most or all anterior and anterov- 
entral bristles black, rarely bristles entirely 
yellow; posteroventral bristles usually yel- 
low; HFWLR 1.0:4.4-1.0:5.0. Tibiae with 
narrow apices brown, sometimes light 
brownish yellow anteriorly. Tarsi yellow se- 
tose; basal tarsomere, soinetimes second 
tarsomere, mostly yellow, apices narrowly 
brown; only black bristles present. 

Abdomen: Mostly dense yellow tomen- 
tose, yellow setose; tergites medially brown 
with sparse black setae; sternites 3—7 and 
anterior margin of tergites laterally with 
yellow bristles. 


Figs. 47-52. Oiiiiiuiliii\ pcnicccssarius. male terminalia. 47-49. Lateral, dorsal, and ventral views. .50, Ae- 
deagus. dorsal view. 31, Left aonostylus. 52, Aedeagiis, lateral view. 

Terminalia (Figs. 47-52): Gonostylus 
narrow, posterior concavity absent. Gono- 
coxite long, narrow apicaliy. tightly 
wrapped abound base of gonostylus. Ae- 
deagus wide, strong prebasal constriction 
and prominent ventral keel present. 

Female. — Differs as following. Body 
10.8-14.2 mm; wing 10.5-11.7 mm: 
FHWR 1.0:5.8-1.0:6.3; FWLR 1.0:1.6-1.0: 
2.7; m, WR 1.0:2.5:2.4-1.0:4.7:4.7; 

HFWLR 1.0:5.0-1.0:5.4. Head: Face with 
10-12 black bristles and dorsally mostly 
black setae. Thorax: Scutellum with mar- 
ginal bristles always black, preapical 
groove often absent. Wing: Base of cell r4 
well beyond apex of cell d; r-m crossvein 
just beyond middle of cell d. Legs: Color 
of femora variable, blackish with narrow 
bases yellow to hind femur yellow with 
only narrow apex dark brown to black. 


Hind femur posteroventrally with 3-3 lon- 
ger, thinner yellow bristles. Abdomen: Most 
tergites apically with 3-4 thick, long, yel- 
low bristles, tergite 8 with only short, black 
bristles; sternites 2-7 with few to several 
yellow bristles. Sternite 8 anteromedially 
reddish yellow to yellow, with a raised, 
oval tubercle, usually divided longitudinal- 
ly. Terminalia: (Figs. 53-56). Sternite 8 
with prominent medioapical tubercle: sper- 
matheca constricted before apex, apex wid- 
er. Genital fork with short, wide base. 

Specimens examined. — Holoype 6\ 
UGANDA: Budongo Forest nr Lk. Albert/ 
1000 m/iv.l972/E. B. Babyetaqara (CNCI). 
CONGO: 1 6,1 9 , Musee Du Congo/Uele 
Aba/14, 20.iv.l914/Dn Rodhain (MRAC): 1 
9. Musee Du Congo/Uele: Van Kerkhevan- 
ville/Degreef (MRAC); I 9 , Musee Du Con- 
go/Uala/xii.l932/A. Corbisier(MRAC): 1 9, 
Musee Du Congo/Iturl/Niarembe v. 1921/Ch. 
Scops (MRAC): 1 9, Musee Du Congo/Dun- 
gu/iii.l920/R Van den Plas (MRAC); 1 9, 
7052 Lubwe/sur feuille Arbre/viv. 7.xii.55 
Lips E" Villa. (MRAC): 1 9. 10 mi. S. of/ 
Kapona/13.i.58/1570 m/E. S. Ross &, R. E. 
Leech. (CASC). BURUNDI: 1 6, Mwaro/ 
Gigoli/21.x.80/J. G. Pointel (MNHN). ETHI- 
OPIA: 1 6, Sidamo/Provience Dilla/ O. DeMasil/C. A. S. Acces- 
sion (CASC). KENYA: 1 J, 2 9, Kakamega/ 
v-vi.l950/Mrs. Adamson (BMNH): 1 9. 
KENYA; Kapsabet/10.v.l991/A. Freidberg 
& Fini Kaplan (NMSA). MALAWI: 1 <5. 1 
9, Nyasaland/Usangha distr. 35-4500 ft/ 
29.xi-15.xii.l910/S.A. Neave (BMNH). NI- 
GERIA: 1 <5, 2 9, Obridu cr/SE state/ 
25.iii.l971/J.T Medler(BMNH). RWANDA: 
1 6, Gabiro, Kegera Park/1325 m 8.xii.57/E. 
S. Ross & R. E. Leech (CASC). UGANDA: 
1 9, N. Uganda: van Someren/Madi Opei n. 
u/v.5I/Com. Inst. Ent. coll No. 12301 
(BMNH); 1 6. 1909/Col. Sir D. Bruce/1909- 
155 [terminalia on slide S.58] (BMNH); 1 9. 
May 1914/Uganda Uny-/oro Hoima/3700 
feet/Coll. Kitter-/bergerd dd ■19/C. A. Wig- 
gins (OXUM). TANZANIA: 1 9. Njombe/ 
6000-6500 ft/Tanganyika/9.xii.l957 W. R 

(BMNH): 1 6. 1 9. 41 mi. N./of Kasulu 
1150 nV19.xi.l967/K. S. Ross &/A. R. Ste- 
phan (CASC). ZAMBIA: 1 9, Lusaka/8.i.41/ 
Frank M. Hull Collection (CNCI). COUN- 
TRY UNKNOWN: 1 9, Yl, presented by/ 
Imp. Bur. Ent./1921-9. [Prey mounted on 
separate pin] same data except Lectislhe- 
quaertilV.a\\erJdei. in B. M.AV. E. China 
[Cercopidael (BMNH). 

Etymology. — Latin pernecessoriiis for 
'very close by relation" referring to its sim- 
ilarity to others in this group. 

Distribution. — Captured in October 
through June at altitudes between 1150 to 
1900 meters from Southern Ethiopia west- 
ward to Nigeria and southward to Zambia. 

Remarks. — The short flagellum; sparse, 
short setae on the dorsal half of the face; 
black thoracic, femoral, and tarsal bristles; 
and combined characters of the terminalia 
(Figs. 47—56) distinguish O. pernecessarius 
from congeners. The shapes of the gonos- 
tylus, gonocoxite, and aedeagus readily 
characterize the male. The raised, oval, 
preapical tubercle on sternite 8, the preap- 
ically constricted spermatheca, and wide, 
short base of the genital fork further char- 
acterize the female. 

Ommatius perscientiis Scarbrough, 
new species 

(Figs. 6. 57-62) 

Male.— Body 8.7-14.2 mm; wing 7.3- 
9.8 mm. Head: Face yellow tomentose, sev- 
eral, thin, black bristles and sparse setae 
present on dorsal %; thick yellow bristles 
and abundant long setae present on ventral 
Vs: FHWR 1. 0:8. T-l. 0:9.8. Antenna, frons, 
and ocellar tubercle black setose. Flagellum 
short, as long as scape; FWLR 1.0:1.5. 
Frons yellowish-brown tomentose; setae 
short, longest seta slightly longer than 
scape. Ocellar tubercle with 2 setae as long 
as 3 antennomeres combined. Occiput 
mostly thin brownish-gray tomentose, eye 
margin narrowly dense yellowish gray; 
scattered black setae present dorsally; long, 
thin, black postocular bristles extend be- 
yond midlateral margin of head, longest 


Figs. 53-56. Oninuiliiis peniecessariu 
1. 56, Genital forl<. 

female tenninalia. 53-54. Dorsal and ventral views. 55. Sperniathe- 

dorsal bristle with tip well before posterior 
margin of eye. 

Thorax: Mesonotum brown tomentose 
dorsally, distinct paramedial stripes and lat- 
eral spots absent: bristles black; 4 dorsocen- 
tral bristles present, wide spaced with first at 
of just behind transverse suture. Scutellum 
brown tomentose dorsally, apical margin 
gray: scattered black setae and 2 black mar- 
ginal bristles present: weak preapical groove 
present. Anepimeral bristle black: anepister- 
num dorsally scattered black setose. 

Wing (Fig. 6): Dense, black microtrichia 
present beyond r-m crossvein: veins dark 
brown. Cell m,WR 1.0:2.1:1.9-1.0:2.9:2.6. 

Legs: Coxae with stout bristles. Femora 
mostly black; fore and middle femora yel- 
low ventrally and posteriorly except at 

apex: hind femur yellow on basal fourth to 
third. Middle femur with black bristles an- 
teriorly and anteroventrally: posteroventral 
bristles variable, all yellow, black, or mixed 
with apical bristles black. Hind femur with 
anterior and anteroventral bristles variable, 
entirely yellow or black, if mixed then 1—3 
basal anteroventral bristles yellow: poster- 
oventral bristles entirely yellow or 3-4 api- 
cal bristles black; HFWLR 1.0:5.0-1.0:5.2. 
Tibiae yellow except brown as follows: nar- 
row apex of fore and middle tibiae and api- 
cal half of hind tibia; bristles black. Tarsi 
mostly black, basal tarsomere of fore and 
middle tarsi mostly yellow, narrow apex 
black; bristles black. 

Abdomen: Black, apical margin of most 
segments dull yellow to yellowish brown. 



Figs. 57-62. OnumiUiis pernecessarius, male terminalia. 57-59, Lateral, dorsal, and ventral views. 60. Ae- 
deagus. dorsal view. 61. Left gonostylus. 62. Aedeagus. lateral view. 

Terga mostly thin, brown tomentose. basal 
segments brownish gray laterally; black se- 
tose; apical corner of terga 4-8 with 1-4 
thin, black bristles. Sterna brownish-gray 

tomentose, mostly pale yellow setose; ster- 
nites 6-7 with scattered, short, black setae; 
sternum 8 with only black vestiture. 

Tcniiliuilla (Figs. 57-62): Gonostylus 


narrow apically, posterior groove absent. 
Aedeagus slightly sigmoid with a promi- 
nent ventral keel. 

Female. — Unknown. 

Specimens e.\amined. — Holotype d , 
ZIMBABWE: Mt. Selinda/Nov-Dec 1930/ 
R. H. R. Stevenson (OXUM). Paratype. 
ZIMBABWE: 1 6. [without abdomen], 
same data label as holotype (OXUM); 1 6. 
Vumba Mts./S. Rhodesia/29.xi.l934/Major 
Dry.sdale (OXUM). ZAMBIA: 1 6. South- 
ern Rhodesia/Chitinda Forest/25. viii- 
2.V.1907/C. F M. Swynnerton/1915-426 

Etymology. — Latin perscientus for 'very 
skillful", alluding to the predaceous habit of 

Distribution. — A montane species cap- 
tured during August and November-De- 
cember in eastern Zambia and Zimbabwe. 

Remarks. — In addition to the characters 
in the key, the nanow face with sparse setae 
dorsally; dorsal postocular bristles procli- 
nate with apices well forward of the pos- 
terior margin of the eyes; mesonotum with 
thin brown tomentum, paramedial stripes 
absent; wing with dense brown microtrichia 
beyond r-m crossvein; hind femur black ex- 
cept for nanow yellow base; apical half of 
hind tibia black, and terminalia (Figs 57- 
62) characterize O. perscientus. 


I thank John Chainey (BMNH), Keve Ri- 
bardo (CASC). J. M. Gumming (CNCI), Ja- 
son G. H. Londt (NMSA), Eliane De Con- 
inck and Jos Debecker (MRAC), Lenonidas 
Tsacas, (MNHN), D. Sithole (NMZB), and 
Darren Mann (OXUM) for the loan of spec- 
imens. Thanks are also extended to D. Sith- 
ole for donating paratypes of O. flavipennis 
to the NMNH, Washington, DC; to Darin 
Mann, A. C. Pont, J. W Ismay and Dorothy 
Newman for permitting access to the ento- 
mological collection and library, to exam- 
ine indispensable type specimens, and dis- 
cussing questions regarding type status and 
nomenclature during our visit to the Oxford 
Museum. Special thanks are extended to Dr 

and Mrs. M. J. Smart of Wolverhampton, 
England, for advise relating to our visit to 
Oxford, their kindness and friendship dur- 
ing our visit to their home; to Rosa Scar- 
brough for encourage and assistance during 
our visit to the Oxford Museum. William 
Grogan, Department of Biological Scienc- 
es, Salisbury University. Maryland, and 
Raymond Gagne, Systematic Entomology 
Laboratory, ARS, USDA kindly reviewed 
the manuscript. 

Literature Cited 

Arnett, R. H., G. A. Samuelson. and G. M. Nishida. 
1993. The Insect and Spider Collections of the 
World, 2nd edition. Sandhill Crane Press, Gaines- 
ville, FT., USA. 310 pp. 

Lindner, E. 1955. Oslatrkanische Asiliden (Dipt.). Er- 
gebnisse der Deutschen Zoologischen Ostafrika- 
Expedition 1951-1952. Jahreshefte Verein vater- 
landische Naturkunde in Wurttemberg, Stuttgart 
1 10: 24-46. 

McAlpine. J. F. 19X1. Morphology and terminology — 
adults, pp. 9-63. In McAlpine. J. E et al.. eds. 
Manual of Neartic Diptera. Biosystematic Re- 
search Institute. Ottawa Agriculture Canada. 
Monograph No. 27. Volume I, 674 pp. 

Oldroyd. H. 1980. Family Asilidae. pp. 334-373. /n 
Crosskey. R. W.. ed. Catalogue of the Diptera of 
the Afrotropical Region. London: British Museum 
(Natural History). 1347 pp. 

Scarbrough. A. G. 2002a. Rede.scriptions of two spe- 
cies of Ominatiiis Wiedemann (Diptera: Asilidae) 
with lectotype and paratype designations for Om- 
inalins tenelhis van der Wulp. range extension, 
and a replacement name for Ommatiiis tibialis Ri- 
cardo. Proceedings of the Entomological Society 
of Washington 104: 680-686. 

. 2002b. Synopsis of the Neotropical lioloseri- 

ceiis complex of the genus Ommalius Widemann 
(Diptera: Asilidae): ampliatus and Itolosericeiis 
species groups. Transactions of the American En- 
tomological Society 128: 133-222. 

Scarbrough A. G. and C. Marascia. 1996. Status of the 
genus Emphysomera Schiner, 1866 (Diptera: As- 
ilidae), with a synopsis of Afrotropical species. 
Annals of the Natal Museum 37: 191-213. 

. 1999. Synopsis of the Oriental and Australian 

species of Emptiysomera Schiner (Diptera: Asili- 
dae). Deutsche Entomologisches Zeitschrift 46: 

. 2000. Revision of (Mmiiariii.s Wiedemann 

(Diptera: Asilidae): Subgenus Metoiiiinatius Hull, 
1962. Annals of the Natal Museum 41: 157-179. 
Wiedemann. C. 1821. Diptera exotica. Kiliae. Part I, 
pp. i-xix. 1-244 |in Latin). 2 plates. 8 figures. 
(Asilidae pp. 179-242.) 

105(3). 2003. pp. 630-640 


John W. Brown 

Systematic Entomology Laboratory, PSI, Agricultural Research Service. U.S. Depart- 
ment of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, 
RO. Box 37012, MRC-168, Washington, DC 20013-7012 (e-mail: jbrown@sel.barc. 

Abstract. — Three new genera in the tortricid tribe Euliini are described and illustrated: 
Eiiryeiilia. with type species E. hiocellata (Walsingham 1914), new combination, from 
Mexico; Pseudapina. with type species P. lanceovalva, new species, from Venezuela; and 
Circapina. with type species C. flexalaiia, new species, from Costa Rica. Three new 
generic synonymies are proposed: Pycnospina Razowski 1997 is a junior synonym of 
Lohogenesis Razowski 1990, with the new combination L. centrota (Razowski 1997); 
Osmaria Razowski 1991 is a junior synonym of Aiiopinci Obraztsov 1962, with the new 
combination A. psaeroptera (Razowski and Becker 1991); and Chirotes Razowski and 
Becker 1999, along with its replacement name, Prochi rotes Razowski 2001, are junior 
synonyms oi Strophotina Brown 1998, with the new combinations S. chorestis (Razowski 
and Becker 1999) and S. niphochondra (Razowski and Becker 1999). 

Ke\ Words: genitalia, biodiversity, synonymy, new combinations, classification, Eii- 
ryeulia. Pseudapina, Circapina, Pycnospina, Lohogenesis, Osmaria. Ano- 
pina, Chirotes. Prochirotes, Strophotina 

Since 1980, more new genera (n = 107) 49), relationships with other genera ;ue be- 
have been proposed in the tribe Euliini than coming more clear, and there is a need to 
in any other tribe of the family Tortricidae. synonymize those that likely represent ex- 
All of these new genera are restricted to the tremes within a single monophyletic lineage 
New Worid tropics, presenting compelling (ie., genus). The purposes of this paper are 
evidence of the hyperdiversity of the group to describe three new genera, propose the 
within this large and complex geographic re- synonymy of three previously described gen- 
gion. Based primarily on material collected era, describe two new species, and propose 
by Vitor Becker of Planaltina, Brazil, the vast ^ve new combinations. The overall goal is to 
majority of these senera have been described continue to build upon the growing generic 
by Jozef Razowski. who has nearly single- framework for the tnbe and to clanly rela- 
handedly created a generic framework mto tionships in an eftort to establish a stable, 
which new species of Neotropical Euliini can 
be placed. However, this task is far from 
complete, with numerous described and un- 
described species still defying confident ge- Specimens were obtained from or ex- 
neric assignment. In contrast, for some of the amined at the following institutions: The 
monotypic genera, which are numerous (n = Natural History Museum, London, England 

meaningful classification. 

Materials and Methods 



(BMNH): Institiito Nacional de Biodiver- 
sidad, Santo Domingo de Heredia, Costa 
Rica (INBio): Essig Museum of Entomol- 
ogy, University of California. Berkeley. 
California. U.S.A. (UCB): National Muse- 
um of Natural History. Smithsonian Insti- 
tution, Washington. D.C., U.S.A. (USNM); 
and Zoological Museum. Copenhagen. 
Denmark (ZMC). Genitalia preparations of 
representative individuals were made fol- 
lowing the methodology summarized in 
Brown and Powell (2000). Illustrations of 
genitalia are photographs of slide mounts 
taken with a SONY DKC5000® digital 
camera and enhanced using Adobe Pho- 
toshop® and Adobe Illustrator® software. Il- 
lustrations are of a single representative 
preparation, but some have been slightly 
modified where the parts are disarticulated. 
Descriptions are composite, based on all 
available specimens. Forewing measure- 
ments were made with a transparent milli- 
meter ruler under low power of a Leica 
MZ12® dissecting microscope, and estimat- 
ed to the nearest one-tenth millimeter Ter- 
minology for wing venation and genitalia 
structures follows Horak (1984). Abbrevi- 
ations are as follows: FW = forewing; HW 
= hindwing; DC = discal cell; n = number 
of individuals examined; x = mean; ca. = 
about (approximately). Elevations present- 
ed in feet on data labels are converted to 
meters and given in brackets. 


Euryeulia Brown, new genus 

Type species: Torth.x hiocelkitci Walsing- 
ham 1914. 

Diagnosis. — Superficially. Euryeulia is 
like no other genus in Euliini. Its forewing 
pattern, with a pair of rounded, maroon spots 
in the distal half of the wing (Fig. 1 ), is more 
similar to some species of Episimus Wal- 
singham (Olethreutinae) than to other euli- 
ines. The genitalia, likewise, are moderately 
divergent from other members of the tribe. 
The elongate, sinuate acces.sory rods origi- 
nating from the base of the gnathos (Fig. 4) 

are reminiscent of a similar stmcture in Gna- 
theiilia Razowski; the capitate uncus is sim- 
ilar to that of Anopina hilasma (Walsingham) 
and Gaitrunciis Razowski; and the narrow 
valvae are reminiscent of Emocomutia Ra- 
zowski. The inflated apophyses posteriores in 
the female genitalia (Fig. 7) are similar to the 
those of Paraptila Meyrick. Putative auta- 
pomorphies include all the unusual characters 
mentioned above because most appear to be 
derived independently within Eiiiyeulia. The 
relationship of Eiiiyeiilici to any of the men- 
tioned genera is obscured by the large num- 
ber of autapomoiphies. Eutyeiilia is assigned 
provisionally to Euliini on the basis of its 
possible relationship with other genera in the 
tribe based on the characters mentioned 

Description. — Head: Antennal cilia 1.2- 
1 .3 times width of fiagellomere in male; 
short, unmodified in female. Labial palpus 
(all segments combined) ca. 1.2-1.4 times 
horizontal diameter of compound eye, seg- 
ment 11 weakly upturned, rough scaled, ex- 
panded distally by scaling; segment III ca. 0.2 
times as long as II, smooth scaled, with tip 
exposed. Maxillary palpus rudimentary. Dor- 
sal portion of frons with short overhanging 
tuft of scales; lower portion smooth scaled. 
Ocellus moderate in size. Chaetosemata pre- 
sent. Proboscis present, presumably function- 
al. Thorax: Dorsum smooth scaled; legs un- 
modified, male without foreleg haiipencil. 
FW length ca. 2.5 times width; DC length ca. 
0.65 times FW length: DC width 0.16-0.18 
times DC length; CuA, originates ca. 0.7 
along length of DC; all veins separate beyond 
DC; chorda absent; M-stem absent; CuP pre- 
sent only at margin. Raised scale tufts absent; 
male without costal fold. Hindwing with 
Sc-HR and Rs approximate at base; Rs to ter- 
men; Rs and M, approximate at base; M, and 
M, separate; M, and CuA, connate. Abdo- 
men: Dorsal pits absent; no modified coreth- 
rogyne scaling in female. Male genitalia (Fig. 
4) with uncus weakly curved, enlaiged and 
somewhat cordate apically. Socius moderate- 
ly long, slender, ca. 0.9 times length of gna- 
thos amis, pendant; not fused to gnathos. 


Gnathos complete, arms joined distally into 
short trianguUir plate; an elongate, slender, 
sinuate, accessory arm arising at base of each 
arm of gnathos: accessory arms ca. 1.5 times 
length of gnathos proper Siibscaphium and 
hami absent. Transtilla a broad band, with a 
few small thorns. Valva narrow, costa cur\ ed 
\ entrad just before apex; sacculus ill-dehned. 
without free process. Pulvinus absent. Vin- 
culum present: tegumen unmodified. Juxta an 
iUTOw head-shaped, sclerotized plate. Aedea- 
gus moderate in size. ca. 0.66 length of \ ah a; 
vesica with small patch of tiny comuti: phal- 
lobase simple, rounded. Female genitalia 
(Fig. 7) with papillae anales moderately slen- 
der: apophyses anteriores long and slender: 
apophNses posteiiores slightly shorter, broad- 
l\ inflated in anterior tv\o-fifths, w ith narrow 
tip at anterior end. Sterigma a broad trans- 
\erse band, niirrowest at middle, dilated lat- 
erally; area immediately anterad of papillae 
anales a large, broad cup: antrum a large 
sclerotized bowl. Ductus bursae moderately 
long and broad, with a slender accessory bur- 
sa arising from right side at antrum. Corpus 
bursae relatively short, oblong, finely punc- 
tate: signum absent. 

Etymology. — The genus name is derived 
from the Greek "eurys." meaning broad, 
and "Eiilla". the type genus of the tribe, 
and is interpreted as feminine in gender 

Eiiiyt'ulia biocellata (Walsingham 1914). 
ne>v combination 

(Figs. 1, 4. 7) 

Tornix biocellata Walsingham 1914: 278 
(description and illustration of adult). 

••Eitlia" biocellata: Powell et al. 1995: 146 

Diagnosis. — As mentioned above, Eu- 
lyeidia biocellata cannot be confused with 
any other euliine. It is recognized easily by 
its EpisiiniisA'ike facies and unique genita- 
lia. The species, originally described in the 
Linnaean genus Tortrix, has nothing in 
common with that genus, which includes 
one species from Europe and one from 
Asia. Its transfer to "Eulia" in the Check- 

FigN. l-.v Adults ol new lortricid geneni. 1. Eii- 
ryeulia biocellalci. 2. P\tiulapina lancemutvu. y. Cir- 
capiria flexalana. 

list of Neotropical Lepidoptera (Powell et 
al. 1995) was merely for convenience — a 
place for many species of Euliini that 
lacked meaningful generic assignment. Eu- 
lia is a monotypic genus that is Holarctic 
in distribution. 

Redescription. — Male. Head: Frons 
white: vertex cream: labial palpus cream on 
inner surface, cream and pale brown on out- 
er surface. Thorax: Mixed cream and pale 
brown, with dark brown. V-shaped band 
bordered at posterior edge by cream: tegula 
red-brown. FW (Fig. 1) length 5.9-7.2 mm 
(x = 6.4: n = 4): ground color gray, with 
narrow, transverse, gray-brown lines, 
broadly bordered by pale yellow : two con- 
spicuous, circular, tawny maroon spots, one 
at costa ca. 0.6 distance from base to apex, 
extending to lower angle of DC. the second 
smaller, situated midway between the costal 
spot and mid-termen. Fringe whitish. Ab- 


domen: Pale brown. Genitalia (Fig. 4; based 
on 2 preparations) as described for genus. 

Female. Head and thorax: As described 
for male. FW length 8.0 mm (n = 2); as 
described for male. Abdomen: Pale brown. 
Genitalia (Fig. 7; based on 2 preparations) 
as described for genus. 

Types. — Lectotype 9, Mexico, Guenero, 
Amula, 6000' |1846 m], Aug 1918. H. H. 
Smith (BMNH). Paralectotype 6. same data 
as lectotype (BMNH). Walsingham (1914) 
stated "Type 9 (66513); 6 (66514)" which 
has been interpreted as the designation of a 
female type and a male type. Hence, a lec- 
totype designation is necessary in order to 
stabilize the nomenclature and identity of this 
species. The female already bears a BM "lec- 
totype" label. Walsingham's (1914) type se- 
ries consists of the two specimens cited 
above; a third identically labeled specimen in 
BMNH was not included. 

Additional specimens examined. — MEX- 
ICO: Durango: 3 mi E Revolcaderos. 1 1 
Aug 1972. J. Powell (\ S . UCB). Guerrero: 
Amula. 6000' [1846 m]. Aug [no year). H. 
H. Smith (\ i. BMNH): 5 km W Tixtla. 
1710 m. 18 Sep 1982. J. Powell & J. Chem- 
sak (1 6.2 9. UCB). Sinaloa: 2 mi SW 
Potrerillos. 4200' [1285 m], 12 Aug 1986, 
J. Brown & J. Powell (Id, UCB). 

Distribution and biology. — Euryeulia 
biocellata has been recorded only from the 
middle elevations ( 1290-1850 m) of the Si- 
erra Madre Occidental of western Mexico. 
The general habitat of the collecting local- 
ities in Durango and Sinaloa is pine-oak 
forest. Collection dates are August and Sep- 
tember All recent specimens were taken at 

Pseudapiiia Brown, new genus 

Type species: Pseudapiiui lamcovcdva 
Brown, new species. 

Diagnosis. — Superficially, Pseudapiiui is 
nearly indistinguishable from Circapiiia. 
new genus (Figs. 2, 3), described below, 
and based on facies alone it is hard to imag- 
ine that they are not congeneric. However, 

characters of the male and female genitalia 
(Figs. 5-6, 8-9) provide convincing evi- 
dence that the two are distinct. Putative au- 
tapomorphies for Pseiidapina include the 
slender, dorsally projecting socii and the 
broadly lanceolate valvae (Fig. 5). Pseii- 
dapina shares with Odonthaliiiis bisetaniis 
Brown and O. inipropriiis Brown a patch of 
strong, curved spines from the basal portion 
of the valva at the lateral base of the tran- 
stilla, and a distally attenuate valva (Brown 
2000). The female genitalia of Pseiidapina 
share with many species of Odonthalitiis 
extremely short apophyses anteriores (Fig. 
8). The erect, sclerotized structures from the 
dorsum of the tegumen interpreted as a 
modification of the socii in Pseiidapina 
may be homologous with what has been in- 
terpreted as a paired or bifurcate uncus in 
Odonthalitiis. but this is not without ques- 
tion. Their position and configuration also 
are reminiscent of the hami of Chlidanoti- 
nae, but it is highly unlikely that these 
structures are homologous with those found 
in Chlidanotinae. The putative homology of 
the spines of the valva and the socii/uncus, 
along with the extremely short apophyses, 
argue for a close relationship between 
Odonthalitus and Pseiidapina. Assignment 
of Pseiidapina to Euliini is based on the 
possession of a male foreleg hairpencil, 
which has been identified as a putative syn- 
apomorphy for the tribe (Brown 1990). 

Description. — Male. Head: Antennal cil- 
ia 1.2-1.5 times width of flagellomere in 
male. Labial palpus (all segments com- 
bined) ca. 1.5 times horizontal diameter of 
compound eye; segment II weakly up- 
turned, rough scaled, expanded distally by 
scaling; segment III ca. 0.5 times as long as 
II. smooth scaled, exposed. Maxillary pal- 
pus rudimentary. Dorsal portion of frons 
with short overhanging tuft of scales; lower 
portion smooth scaled. Ocellus small. Chae- 
tosemata present. Proboscis present, pre- 
sumably functional. Thorax: Dorsum 
smooth scaled; male with foreleg hairpen- 
cil. FW length ca. 2.5 times width; DC 
length 0.65-0.70 times FW length; DC 


width 0.16-0.18 times DC length: CiiA, 
originating ca. 0.7 along length of DC; all 
veins separate beyond DC; chorda absent; 
M-stem absent; CuP present at margin. 
Raised scale tufts absent; male without cos- 
tal fold. Hindwing with Sc + R and Rs ap- 
proximate at base; Rs to termen; Rs and M, 
approximately at base; M, and M, separate; 
M, and CuA, connate or short-stalked. Ah- 
clomcii: Dorsal pits absent; no modified cor- 
ethrogyne scaling in female. Male genitalia 
(Fig. 5) with uncus weakly club-shaped, 
gradually expanded in distal three-fourths, 
sparsely setose in distal one-ft)urth. Struc- 
ture interpreted as base of socius elongate, 
scleroti/.ed. attenuate, projecting dorsally. 
hooked apically. nearly as long as uncus; 
lower portion of socius short, broad, pen- 
dant. Cinathos ci)mplete, somewhat mem- 
branous basally. arms joined distally into 
short, hoodlike process. Subscaphium and 
hami absent. Transtilla with stout, sclero- 
tized lateral bases, membranous mesally. 
Valva broad at base, attenuate apically. with 
narrow apical beak; group of 10-15 stiff, 
curved setae arising near base of transtilla: 
sacculus not developed. Pulvinus absent. 
Vinculum complete, well developed. Juxta 
a scleroti/.ed plate. Aedeagus moderately 
small, narrow, weakly curved in distal one- 
(ifth, ca. i>ne-half length of valva: phallo- 
hase slightU elongate, simple, rounded at 
base; \esica without cornuti. Female geni- 
talia (Fig. 8) with papillae anales broadly 
slipper-shaped; apophyses extremely short, 
especially apophyses anteriores. Sterigma a 
narrow, spiculate, transverse band. Ductus 
bursae extremely long, slender. Coipus bur- 
sae round, densely and evenly spiculate; 
signum absent. 

Etymology. — The genus name is a ci)n- 
traction of the Greek "pseudos,'" meaning 
false, and "Anopitui." a genus with which 
Pseudupiiui is superficially similar. It is in- 
terpreted as feminine in gender. 

Pseudapina lanceovalva Brown, 
new species 

(Figs. 2, 5, 8) 
Diagnosis. — As mentioned abo\e, P. 
lanceovalva is superficially nearly indistin- 

guishable from Circupina flexalaiui. de- 
scribed below. Male genitalia of P. lanceo- 
valva are easily distinguished by the slen- 
der, attenuate, dorsally-projecting socius, 
weakly club-shaped uncus, broadly lance- 
olate valva, and unmodified aedeagus. The 
group of curved spines near the base of the 
valva is similar to that present in two spe- 
cies of Odonlhaliliis Razowski. Female 
genitalia are distinguished by the extreiTiely 
long, slender ductus bursae, which is simi- 
lar to that of Odouthalitns poas Brown. 

Description. — Male. Head: Frons, ver- 
tex, and labial palpus with cream to pale 
tan scales. Thorax: Cream to pale tan. FW 
(Fig. 2) length 5.8-6.1 mm (x = 5.9; n = 
2); ground color white, sprinkled with 
brown and red-brown scales; ill-defined 
basal patch of pale red-brown scales; a sub- 
rectangular patch of pale red-brown scales 
near middle of costa; three tiny dashes from 
costa between costal patch and apex; apex 
with ill-defined pale brown patch. Fringe al- 
ternating pale gray and pale orange. Abdo- 
men: Pale brown. Genitalia (Fig. 5; based 
on 2 preparations) as described for genus. 

Female. Head: As described for male. 
Thorax: FW length 7.0 (n = I), pattern as 
described for male. Abdomen: Genitalia 
(Fig. 8; based on 1 preparation) as de- 
scribed for genus. 

Holotype. — 6. Venezuela. Aragua. Ran- 
cho Grande [1100 m]. 1-7 Aug 1967. R. 
W. Poole (USNM). 

Paratypes. — Venezuela: Aragua: Rancho 
Grande. 1 100 m. 24-31 Oct 1966. S. S. & 
W. D. Duckworth (19. USNM). 8-14 .lul 
1967, R. W. Poole (1 cJ, USNM). 

Distribution and biology. — Pseudapina 
lanceovalva is known only from the type 
locality. Niithing is known of the early stag- 
es. Adults have been collected in July, Au- 
gust, and October. 

Etymology. — The species name refers to 
the broadly lanceolate shape of the \al\a in 
the male genitalia. 

Circapiiia Brown, new genus 

Type species: Circapina fiexalana Brown, 
new species. 


Figs. 4—6. Male genitalia. 4. Eiiryeulia hiocellata. 3, Pseiidapina lanccovalva. 6. Ciicapiini flexalana. 


Diagnosis. — Adults of Circapina are 
most similar to those of Pseudapinci in fore- 
wing length and pattern — the two are nearly 
indistinguishable. However, as discussed 
above, the two are easily separated by nu- 
merous features of the male and female 
genitalia. Putative autapomorphies for Cir- 
capinci include the slender, weakly sclero- 
tized, sparsely hairy uncus; the erect, sem- 
icordate socii; and the broad, apically 
hooked aedeagus. It is likely that Circapina 
is related to Anopina on the basis of the 
elongate antennal cilia in the male, the pres- 
ence of a male foreleg haiipencil, and the 
forewing pattern with a semicircular costal 
blotch, but the same could be said of P.seii- 
dapina. Circapina is assigned to Euliini on 
the basis of the male foreleg hairpencil, 
which may represent a synapomorphy for 
this tribe (Brown 1990). 

Description. — Male. Head: Antennal cilia 
1.2-1.5 times width of flagellomere in male; 
simple, unmodified in female. Labial palpus 
(all segments combined) ca. 1.5 times hori- 
zontal diameter of compound eye; segment 
II weakly upturned, rough scaled, expanded 
distally by scaling; segment III ca. 0.5 times 
as long as II, smooth scaled, mostly exposed. 
Maxillary palpus rudimentary. Dorsal por- 
tion of frons with short overhanging tuft of 
scales; lower portion smooth scaled. Ocellus 
small. Chaetosemata present. Proboscis pre- 
sent, presumably functional. Thorax: Dor- 
sum smooth scaled; male with foreleg hair- 
pencil. FW length ca. 2.5 times width; DC 
length 0.65-0.70 times FW length; DC 
width 0.16-0.18 times DC length; CuA, 
originating ca. 0.7 along length of DC; all 
veins separate beyond DC; chorda absent; 
M-stem absent; CuP present at margin. 
Raised scale tufts absent; male without cos- 
tal fold. Hindwing: Sc4-R and Rs approxi- 
mate at base; Rs to termen; Rs and M, ap- 
proximate at base; M, and M, separate; M, 
and CuA I connate or short-stalked. Abdo- 
men: Dorsal pits absent; no modified coreth- 
rogyne scaling in female. Male genitalia 
(Fig. 6) with entire capsule somewhat ro- 
bust, with legumen broad, especially dorsal- 

ly. Uncus simple, rodlike, very slender, 
weakly sclerotized, sparsely hairy. Socius 
sclerotized, short, erect, with fine hairs, sem- 
icordate, broad at base, attenuating apically, 
not fused to gnathos. Gnathos complete, 
arms moderately broad, short, joined distally 
into short, stout, hooklike mesal process. 
Subscaphium and hami absent. Transtilla 
digitate at lateral bases, membranous mes- 
ally. Valva broadly expanded at base, with 
rounded ventral portion, nanow. parallel-sid- 
ed in distal two-thirds, rounded apically; 
densely hairy, semicircular flap in middle of 
basal one-third; sacculus not developed. Pul- 
vinus absent. Vinculum complete, well de- 
veloped. Juxta a sclerotized plate. Aedeagus 
moderately long, ca. as long as valva, broad, 
with distal one-fifth hooked and strongly 
toothed; phallobase elongate, simple, round- 
ed. Female genitalia (Fig. 9) with papillae 
anales moderately broad, mostly parallel-sid- 
ed; apophyses short, especially apophyses 
anteriores. Sterigma membranous, with 
quadrate lobe mesally. Ductus bursae mod- 
erate in width, weakly sinuate, lightly scler- 
otized, coiled one revolution. Corpus bursae 
oblong, finely punctate but without conspic- 
uous spiculae; signum absent. 

Etymology. — The genus name is a con- 
traction of the Latin "circa," meaning near, 
and "Anopina," a genus with which Cir- 
capina is superficially similar. It is inter- 
preted as feminine in gender. 

Circapina flexalana Brown, new species 

(Figs. 3. 6, 9) 

Diagnosis. — Superficially, Circapina 
flexalana is most similar to Pseiidapina lan- 
ceovalva. It can be distinguished from all 
other Euliini by the slender, hairy uncus, 
broad tegumen with erect semicordate socii, 
and broad aedeagus with curved and 
toothed apex. The latter feature is the most 
striking autapomoiphy for this species. 

Description. — Male. Head: Frons, ver- 
tex, and labial palpus cream to pale tan. 
Thorax: Pale tan with upraised posterior 
crest of shiny gray scales; tegulae whitish. 
FW (Fig. 3) length 4.9-6.6 mm (x = 5.8; 






" v>*^ 


Figs. 7-9. Female genitalia. 7, EtiryruUa hiact'llata. .S. I'sciuUiptiui Uuuctivalva. 9, Circapiiui lltxalaiui. 


n = 12); ground color white, sprinkled with 
brown and red-brown scales; ill-defined 
basal patch of pale brown scales; a semi- 
circular patch of pale red-brown scales near 
middle of costa, with an irregular, pale 
brown, rectangular patch immediately be- 
low; apical patch extending to mid-tornus, 
pale brown with red-brown overscaling. 
Fringe alternating gray and bronze. Abdo- 
men: Pale brown. Genitalia (Fig. 6; based 
on 3 preparations) as described for genus. 

Female. Head: As described for male. 
Thorax: FW length 7.0 mm (n = 2); as de- 
scribed for male, except basal region of 
dorsum overscaled with gray. Abdomen: 
Genitalia (Fig. 9; based on 1 preparation) 
as described for genus. 

Holotype. — i. Costa Rica, Puntarenas 
Province, Monte verde, 1400 m, 22-24 Jul 
1990, S. Meredith & J. Powell (UCB). 

Paratypes. — Costa Rica: Guanacaste 
Province: Est. Cacao, lado suroeste del Vol- 
can Cacao, 1000-1400 m, Sep 1989, C. 
Chavez (\i. INBio), Jun 1990, II Curso 
Parataxon. (Id. INBio), 25 Sep-11 Oct 
1990, C. Chavez (1(5, INBio), Nov-Dec 
1990, C. Chavez & R. Espinoza (IcJ, IN- 
Bio). Puntarenas Province: Monteverde, 
1400 m, 22-24 Jul 1990, S. Meredith & J. 
Powell (4(J, 12, UCB), 29-31 Mar 1992, 
S. McCarty & J. Powell {26, 12, UCB, 
USNM), 8-10 Dec 1978, D. Janzen (IcJ. 
INBio). San Luis, Monteverde, 1000-1350 
m, Feb 1995, Z. Fuentes (IcJ, INBio). 

Distribution and biology. — Cercapina 
flexalana is known from 1000-1400 m el- 
evation on the western side of the central 
Cordillera of Costa Rica. All specimens ex- 
amined are from two localities: Monteverde 
(Puntarenas Province) and SW side of Vol- 
can Cacao (Guanacaste Province). Capture 
dates are scattered throughout the year from 
February to December 

Etymology. — The specific epithet refers 
to the reflexed or curved distal portion of 
the aedeagus. 

Lobogenesis Razowski 1992 
Lobogenesis Razowski 1992: 213; Powell 
et al. 1995: 144; Brown 2000: 25. Type 

species: Lobogenesis lobata Razowski 
1992, by monotypy. 
Pycnospina Razowski 1997: 84. Type spe- 
cies: Pycnospina centrota Razowski 
1997, by monotypy. New synonym. 

Razowski (1997) proposed Pycnospina 
to accommodate the species P. centrota Ra- 
zowski, described in the same paper, known 
only from the holotype male. Based on the 
description and illustration (Razowski 
1997), the species centrota belongs in Lo- 
bogenesis Razowski 1992; I overlooked 
this species in my revision of the latter ge- 
nus (Brown 2000). Lobogenesis centrota 
(Razowski), new combination, shares the 
following synapomorphies with other Lo- 
bogenesis: ( 1 ) uncus bifurcate and finely 
spined in distal one half; (2) socius with a 
rounded lobe extending dorsad of the point 
of attachment to the tegumen; (3) valva 
with a linear row of tiny spinelike teeth ex- 
tending from subbasal spinose lobe to near 
apex; and (4) valva with a strong, digitate 
flange from basal one-third of costa. The 
expanded arms of the uncus beyond the bi- 
furcation, and the club-shaped enlargement 
at the distal ends of the gnathos arms in- 
dicate that L. centrota belongs to a species 
group that includes L. peruviana Brown, L. 
antiqua Brown, and L. contrasta Brown 
(Brown 2000). Inclusion of centrota in Lo- 
bogenesis relegates Pycnospina to a junior 
synonym of Lobogenesis. 

Anopina Obraztsov 1962 

Anopina Obraztsov 1962: 2, 1967: 2; Pow- 
ell 1964: 118, 1983: 39, 1986: 374; Pow- 
ell et al. 1995: 142; Brown and Powell 
1991: 5, 2000: 12. Type species: Tortrix 
triangulana Kearfott 1908, by original 

Osmaria Razowski 1991: 177. Type spe- 
cies: Phtheochroa psaeroptera Razowski 
and Becker 1986. by monotypy. New 

In 1986 Razowski and Becker described 
Phtheochroa psaeroptera from Huatusco, 
Veracruz, Mexico. In 1991, Razowski rec- 



ognized that the species had been placed 
erroneously not only in Phtheochroa. but in 
the tribe Cochylini. To remedy this situation 
he described a new monotypic genus, Os- 
inaria. and indirectly transferred it to Euli- 
ini (i.e.. "it [Osmaria] belongs in the group 
of genera close to Popayanita Razowski."). 
Based on the description and illustration 
(Razowski and Becker 1986), psaeroptera 
is almost certainly a member of the tricm- 
giilana species group of Anopitui Obraz- 
tsov, new combination, as defined by 
Brown and Powell (2000). The overall 
shape of the uncus, gnathos, and valva, and 
the elongate antennal cilia of the male are 
characteristic of Anopiiia. The shape of the 
sacculus and the white color of the fore- 
wing are synapomorphies supporting the re- 
lationship of psaeroptera to a group of 
white Aiwpina species that includes A. des- 
mataiui (Walsingham) (from Guerrero), A. 
albominima Brown and Powell (from Guer- 
rero), and A. alhoinaculana Brown and 
Powell (from Sinaloa and Nayarit). See 
Brown and Powell (2000) for comparisons 
of genitalia. Inclusion of psaeroptera in An- 
opiiia relegates Osmaria to a junior syno- 
nym of Anopina. 

Strophotiiia Brown 1998 

Strophotina Brown 1998: 44. Type species: 
Eiilia strophota Meyrick 1926, by origi- 
nal designation. 

Chirotes Razowski and Becker 1999: 417. 
Preoccupied by Chirotes Cuvier 1817. 
Type species: Chirotes chorestis Razows- 
ki and Becker 1999, by original desig- 
nation. New synonym. 

Prochirotes Razowski 2001: 277. Replace- 
ment name for Chirotes Razowski and 
Becker 1999. New synonym. 

I described Strophotina (Brown 1998) to 
accommodate 5. cnrvidagiis Brown (from 
Costa Rica) and Eitlia strophota Meyrick 
(from Colombia, Ecuador, Peru, and Vene- 
zuela), with the latter designated as the type 
species. The following year, Razowski and 
Becker (1999) described Chirotes to ac- 

commodate C. chorestis Razowski and 
Becker (type species) (from Ecuador), C. 
niphochondra Razowski and Becker (from 
Ecuador), and Eiilia strophota Meyrick. 
There is no doubt that the two genera rep- 
resent the same concept, and the synonymy 
is the result of a lack of communication 
among the authors rather than differing tax- 
onomic opinions. The synonymy of Chi- 
rotes and its replacement name, Prochiro- 
tes. with Strophotina results in two new 
combinations: Strophotina chorestis and S. 

My interpretation of species limits within 
the genus differs from that presented by Ra- 
zowski and Becker (1999). I treated all spec- 
imens {43. 69) from South America (i.e., 
Colombia, Ecuador, Peru, and Venezuela) as 
conspecific (i.e., strophota). although I indi- 
cated that it is possible that more than one 
species was represented in the material 
(Brown 1998). Razowski and Becker (1999) 
considered specimens (2 9) from Carchi, 
Maldonado, Ecuador, to represent strophota. 
and the specimen ( 6 ) from Baeza, Napo. Ec- 
uador, to represent chorestis. Although I sus- 
pect that the two names are synonyms, the 
paucity of material inhibits reliable conclu- 
sions. In addition, I suspect that niphochon- 
dra represents a different, undescribed genus 
closely related to Strophotina. I have exam- 
ined two males from Chile (ZMC) that are 
congeneric with niphochondra. These males 
(including the type of niphochondra) have 
short, more attenuate valvae, a broad, stout 
aedeagus that is quite different from that of 
Strophotina. a unique tonguelike mesal por- 
tion of the transtilla, and lack the dense row 
of spines from the costa of the valva. While 
it is possible that these features are autapom- 
ophies at the species level, I suspect that they 
represent characters that define a new genus. 


I thank the following for allowing me to 
examine specimens in their care: Kevin 
Tuck (BMNH), Eugenie Phillips (INBio), 
Jerry Powell (UCB), and Ole Karsholt 
(ZMC). Linda Lawrence, USDA System- 


atic Entomology Laboratory. National Mu- 
seum of Natural History. Washington. D.C., 
skillfully prepared the figures. Karolyn Dar- 
row. Department of Systematic Biology. 
National Museum of Natural History. 
Washington. D.C.. provided the photo- 
graphs of the adults. I acknowledge the fol- 
lowing for helpful comments on the man- 
uscript: Kevin Tuck, The Natural History 
Museum, London, England; David Nickle. 
USDA Systematic Entomology Laboratory, 
Beltsville, Maryland; and Daivd Smith, 
USDA Systematic Entomology Laboratory. 
National Museum of Natural History, 
Washington, D.C. 

Literature Cited 

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logenetic significance of the male foreleg hairpen- 
cil in the Tortricinae (Lepidoptera: Tortricidac). 
Entomological News 101: 109-116. 

. 1998. Description of Stroplwrina. new genus. 

from Central and South America (Lepidoptera: 
Tortricidae). Proceedings of the Entomological 
Society of Washington 100: 43-49. 

. 2000. Revision of Lobogenesis and Oilon- 

thaliliis (Lepidoptera: Tortricidae: Tortricinae). 
with comments on their monophyly. Proceedings 
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Brown. J. W. and J. A. Powell. 1991. Systematics of 
the Chrysoxenii group of genera (Lepidoptera: 
Tortricidae: Euliini). University of California Pub- 
lications in Entomology III. 87 pp. + 143 figs. 

. 2000. Systematics of Anopina Obraztsov 

(Lepidoptera: Tortricidae: Euliini). University of 
California Publications in Entomology 120. 128 
pp. + 174 figs. 

Horak, M. 1984. Assessment of ta.\ononiically signif- 
icant structures in Tortricinae (Lep., Tortricidae). 
Bulletin de la Societe Entomologique Suisse 57: 

Obraztsov. N. 1962. Ano/viui. a new genus of the Cne- 
phasiini from the New World (Lepidoptera. Tortri- 
cidae). American Museum Novitates 2082. 39 pp. 

. 1967. Genera Tortricoidarum. Checklist of gen- 
era and subgenera belonaina to the families Tortri- 

cidae (Ceracidae. Chlidanudae. Schoentenidae and 
Olethreutidae included) and Phaloniidae. Journal of 
the New York Entomological Society 75: 2-1 1. 

Powell. J. A. 1964. Biological and taxonomic studies 
in tortricine moths, with reference to the species 
in California. University of California Publica- 
tions in Entomology 32, 307 pp. 

. 1983. Tortricoidea. pp. 31-42. In Hodges, R. 

W.. ed. Check list of the Lepidoptera of America 
north of Mexico. E. W. Classey. Ltd.. and Wedge 
Entomological Research Foundation. London. 

. 1986. Synopsis of the classification of Neo- 
tropical Tortricinae. with descriptions of new gen- 
era and species (Lepidoptera: Tortricidae). Pan- 
Pacific Entomologist 62: 372-398. 

Powell. J. A.. J. Razowski, and J. W. Brown. 1995. 
Tortricidae: Tortricinae. Chlidanotinae, pp. 138- 
151. In Heppner, J. B.. ed. Atlas of Neotropical 
Lepidoptera. Checklist Part II: Hyblaeoidea — Pyr- 
aloidea — Tortricoidea. Association for Tropical 
Lepidoptera. Scientific Publishers. Gainesville, 

Ra/owksi. J. 1991. Notes on cochyline genus Phrheo- 
cliroci Stephens (Lepidoptera: Tortricidae) with 
descriptions of new American species. Acta Zool- 
ogica Cracoviensia 34: 163-187. 

. 1992. On some peculiar Neotropical tortricine 

genera (Lepidoptera: Tortricidae). SHILAP Revis- 
ta de Lepidopterologia 18(1990): 209-215. 

. 1997. Euliini (Lepidoptera: Tortricidae) of 

Peru with description of new taxa and list of the 
New World genera. Acta Zoologica Cracoviensia 
40: 79-105. 

. 2001. Description of Chamelania Razowski, 

gen. n.. one new species of Proathorybia Ra- 
zowski. 1999, and a proposal for three replace- 
ment names for Neotropical Euliini (Lepidoptera: 
Tortricidae). SHILAP Revista Lepidopterologia 
29: 275-279. 

Razowski, J. and V. Becker. 1986. Cochylidii (Lepi- 
doptera: Tortricidae) collected in Central America 
and Mexico. Acta Zoologica Cracoviensia 29: 

. 1999. Revision of the Neotropical genera Se- 

ticosta Razowski. 1986 and Piinctapinella Brown, 
1991 (Lepidoptera: Tortricidae) with description 
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Pismo Entomologiczne 68: 415-430. 

Walsingham, T de Grey. 1914. Lepidoptera-Heterocera. 
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103(3). 2003. pp. 641-646 


Mai.kiat S. Saini, David R. Smith, and T.ajinder P. Saini 

(MSS. TPS) Department of Zoology. Punjabi University. Patiala — 147002. India (e- 
mail: in); (DRS) Systematic Entomology Laboratory. PSI. Agricultural 
Research Service, U.S. Department of Agriculture, c/o National Museum of Natural His- 
tory. Smithsonian Institution. Washington. DC 20560-0168. U.S.A. (e-mail: 

Abstract. — Anapeptaineiui Konow (Tenthredinidac: Sclandriinae) has been known only 
by its type species. A. iilbipes Konow. from India and Myanniar Two new species have 
been discovered from India and are described: Anapeptamena darjeelingensis, n. sp., and 
A. dhanoiiltiensis, n. sp. A male of the genus, that of A. dhcmoiiltiensis. is described for 
the hrst time. Descriptions, illustrations, and a key are provided for identification of the 

Ke\ Words: India. Mvanmar. Selandriinae 

Intensive survey work in India by the 
first author has documented a large number 
of new records and resulted in the discov- 
ery of many undescribed sawflies. Many 
genera considered monotypic or to include 
few species have been found to be much 
larger and more diverse than previously 
thought. Such is the case \or Anapeptamena 
Konow, a small genus in the Selandriinae 
known only from several specimens of a 
single species from northeastern India and 
Myanmar. Here we review the genus, de- 
scribe two new species from India, and give 
a key to the species. A male of the genus 
also is described for the first time. 

The history of Anapeptamena revolves 
around decisions of synonymy by two au- 
thors. A year after Konow (1898) de- 
scribed Anapeptamena, Cameron (1899) 
described the genus Biisarbia. A year lat- 
er, Konow ( 1900) proposed the synonymy 
of the two genera, and subsequent authors 
followed Konow's treatment, resulting in 
a number of additional species in Anapep- 

tamena. The distinction between Anapep- 
tamena and Biisarbia was not studied un- 
til Malaise ( 1944) was able to examine the 
type species. He discovered and enumer- 
ated a number of differences between the 
two genera and thus treated them as sep- 
arate. Based on Malaise's interpretation, 
all species described in Anapeptamena 
since Konow's (1900) proposed synony- 
my belong either to Biisarbia or other 
genera. From Malaise (1944) to the pre- 
sent, Anapeptamena has remained mono- 
typic. We concur with Malaise's (1944) 
separation of Anapeptamena. 

Holotypes are deposited in the National 
Pusa Collections, Division of Entomology, 
India Agricultural Research Institute, New 
Delhi, India. Paratypes are at Punjabi Uni- 
versity, Patiala, India (PUNJ) and the Na- 
tional Museum of Natural History. Smith- 
sonian Institution. Washington. D.C., USA 

Abbreviations used in the descriptions 
are as follows: EL = eye length: lATS = 



inner apical tibial spur: ICD = intercenchral 
distance: IDMO = interocuhv distance at 
level of median ocellus: ITD = intertegular 
distance: LID = lower interocuhir distance: 
MB = metabasitarsus: OATS = outer api- 
cal tibial spur: OCL = ocello-occipital line: 
OOL = oculo-ocellar line: POL = posto- 
cellar line. 

Aiiapcptaiueim Konow 

Anapeptamena Konov\ 1898: 271. Type 
species: Aiuipepranwiui alhipes Konow. 
By monotypy. 

Description. — Antenna filiform: apical 
segments without ventral membranous ar- 
eas; segments 3 and 4 subequal in length 
or 3 slightly longer than 4: following seg- 
ments gradually decreasing in length to 
apex: pedicel 1.5— 2. Ox as long as its max- 
imum width and almost as long or as long 
as scape. Mandible strongly bent, almost 
at right angle, with a subapical tooth: 
clypeus subconvexly bent from side to 
side with its anterior margin subtruncate 
to slightly circularly emarginate (Figs. 1- 
3); frontal area poorly defined, not sur- 
rounded by an acute, high carina: hind or- 
bits carinate below only, with postgenal 
carina extending only part way up eye. 
Lateral furrows fine, diverging back- 
wards, abruptly ending just before hind 
margin of head. Hind orbits very short, 
head from abo\e narrowing behind eyes. 
Epicnemium strongly convex, separated 
from mesepisternum by a furrow. Mese- 
pisternum in front view obtusely raised 
without carina, rounded at center. Fore- 
wing without anal crossvein; veins M and 
Im-cu strongly converging, not parallel: 
vein Rs more or less faint. Hind w ing with 
two closed cells. Rs and M: anal cell ses- 
sile. Metabasitarsus shorter than follow- 
ing tarsal joints combined, as about 5:6. 
Tarsal claws with long, slender inner 
tooth, nearly as long as outer tooth, with- 
out basal lobe (Figs. 4-6). Head, thorax, 
and abdomen smooth, shining and im- 

Remarks. — Several generic characters 
are slightly modified from pre\ ious char- 
acterizations of Aiuipeptamena (Malaise 
1944. 1963). These include antennal seg- 
ment 3 sometimes slightly longer than 4, 
the two not always equal in length: the an- 
terior margin of the clypeus may be shal- 
lowly, roundly emarginate, not always sub- 
truncate: and the frontal area is sometimes 
distinct due to well defined frontal ridges 
though not surrounded by a sharp carina 
and not always completely indistinct. 

Anapeptamena may be keyed in Mal- 
aise's (1963) key to world genera of Selan- 
driinae. It is distinguished from Bitsarhia 
(type species. Bitsarhia viridipes Cameron 
from Khasia Hills. India), the genus with 
which it has been confused, by the poorly 
defined frontal area, short postgenal carina 
on the hind orbits extending only part way 
up the eye. and the last four antennal seg- 
ments without ventral membranous areas. 
Bitsarhia has a distinct, pentagonal frontal 
area, surrounded by an extremely acute and 
high carina with a transverse lateral carina 
to inner margin of each eye. the entire hind 
orbits with a postgenal carina, and the last 
four antennal segments with ventral mem- 
branous areas which give the segments the 
appearance of bulging downwards. 

Anapeptamena is known only from 
Myanmar and northeastern India. Food 
plants are not known, but most Selandriinae 
feed on ferns. 

Key TO Species 

1 . Malar space half diameter of front ocellus; la- 
brum black, tegula white; intertegular distance 
3.3 X intercenchral distance; lancet not strongly 
narrowing toward apex, with about 7 semilae 
(Fig. 9); A. alhipes Konow 

- Malar space nearly linear; labruni and tegulae 
either both white or both black; intertegular 
distance 4.0x intercenchral distance; lancet ei- 
ther sharply narrowing toward apex (Fig. 10) 
or with fewer than 7 serrulae (Fig. ID 2 

2. Labrum and tegula while to yellow; clypeus 
subtruncate. anterior margin of labrum acute 
(Fig. 2): frontal ridges laterally connected to 
inner margins of eyes by a low transverse 
ridge: lancet sharply narrowing toward apex. 



with about 8 serrulae (Fig. 10) 

A. Jarjec'lingensis. ii. ,sp. 

- Labrum and tegula black; clypeus shallowly 
roundly emarginale. anterior margin of labrum 
rounded (Fig. 3); frontal ridges without tran.s- 
verse ridge to inner margins of eyes; lancet 
short, not strongly narrowing toward apex, 
with about 6 serrulae (Fig. 11) 

A. dhanuulliensis. n. sp. 

Aiuipeptaiucuu alhipes Konow 
(Figs. 1,4, 9) 

Anapeptaniena albipes Konow 1898: 
271.— Malaise 1944; 10-11 (Myanmar 

Female. — Average length, 5.0 inm. 
Black with tegula and legs, except infus- 
cated apical tarsal segments, yellow to 
whitish. Wings hyaline; costa, stigma, and 
venation dark brown to black. 

Antennal length 2x head width; pedicel 
2X longer than its apical width, almost as 
long as scape; segment 3 longer than 4, as 
8;7. Anterior margin of clypeus subtrunca- 
te, labrum broader than long, as 2:1. with 
rounded anterior margin (Fig. 1 ); malar 
space equal to half diameter of front ocel- 
lus; inner margins of eyes parallel in mid- 
dle, faintly converging below; LID:lDMO: 
EL = 2.0:2.4:1.9; OOL:POL:OCL = 1.7: 
1.0:2.0. Frontal area slightly higher than 
level of eyes; supraantennal tubercles and 
frontal ridges indistinct, without transverse 
ridge to inner margin of eye; supraantennal 
pits distinct and removed from antennal 
sockets by a distance equal to diameter of 
pit: median fovea indistinct; circum- and in- 
terocellar funows distinct, postocellar fur- 
row indistinct; postocellar area subconvex, 
broader than long (5:4 at maximum width). 
Mesoscutellum subconvex, appendage not 
carinate. ITD:ICD = 3.5:1.0. Tarsal claws 
as in Fig. 4. I ATS: MB :0 ATS = 2.0:5.5:1.8. 
Abdomen with light golden pubescence. 
Lancet not distinctly narrowing toward 
apex, lateral teeth above serrulae distinct, 
with about 7 serrulae (Fig. 9). 

Male. — Unknown. 

Material examined. — INDIA: West Ben- 
gal: Darjeeling, 2200 m, 26.5.1989, coll. 

M.S. Saini (19. PUNJ); Sikkim: Gangtok, 
1550 m. 14.5.93, coll. M.S. Saini (1 9. 
USNM): Khasia Hills (holotype 9 ). 

Distribution. — India (West Bengal, Sik- 
kim); Myanmar (Kambaiti at 2000 m, close 
to the Yunnan frontier [Malaise 1944)). 

Type. — The holotype female is at the 
Deutsches Entomologisches Institut. Eber- 
swalde. Germany, labeled "Khasia Hills, 
Assam, ■■ "coll. Konow," "Holotypus 
[red]," "Anapeptaniena albipes Knw., As- 
sam," "TYPUS [red]," "Coll. DEI Eber- 
swalde." The head is glued onto a piece of 
cardboard on the same pin; the antennae are 

Anapeptaniena darjeelingensis Saini, 
Smith, and Saini, new species 

(Figs. 2. 5. 10) 

Female. — Length, 5.0 mm. Black with 
labrum, tegula, coxae (except extreme ba- 
ses), midleg (except last tarsal segment), 
and hind leg (except lightly infuscated api- 
cal 3 tarsal segments) yellow to whitish. 
Wings subhyaline; costa, stigma, and ve- 
nation dark brown to black. 

Antennal length 2x head width, pedicel 
2x its apical width, almost as long as 
scape; segment 3 longer than 4, as 5:4. 
Anterior margin of clypeus slightly emar- 
ginate; labrum about as long as broad, 
with acuminate anterior margin (Fig. 2); 
malar space linear; inner margins of eyes 
parallel in middle, faintly converging be- 
low; LID:IDMO:EL = 2.0:2.4:2.1; OOL: 
POL:OCL = 1.5:1.0:1.5. Frontal area 
slightly higher than level of eyes; su- 
praantennal tubercles indistinct but con- 
tinue posteriorly similar to somewhat 
prominent frontal ridges; in middle, fron- 
tal ridge laterally connected to inner mar- 
gins of eye by a low transverse ridge; su- 
praantennal pits distinct and removed 
from antennal sockets by a distance equal 
to diameter of pit; median fovea distinct 
and gently curved in its posterior half 
with bottom almost fiat; interocellar fur- 
row distinct, circum- and postocellar fur- 
rows indistinct; postocellar area subcon- 





vex. broader than long as 5:4 at its max- 
imum width. Mesoscutellum subconvex, 
appendage not carinate. ITD:ICD = 4.0: 
1.0. Tarsal claws as in Fig. 5. IATS:MB: 
OATS = 2.0:6.0:1.8. Abdomen with 
brownish golden pubescence. Lancet 
strongly narrowing toward apex, lateral 
teeth above serrulae indistinct, with about 
8 serrulae (Fig. 10). 

Male. — Unknown. 

Type. — Holotype 9. West Bengal: Dar- 
jeeling, 2200 m. 8.5.1983. coll. M.S. Saini. 

Distribution. — India (West Bengal). 

Etymology. — The species name is de- 
rived from the type locality. 

Anapeptcimena dhanoiiltiensis Saini, 
Smith, and Saini, new species 

(Figs. 3, 6-8. 11) 

Female. — Length. 4.2 mm. Black with 
tibiae yellow and rest of legs fuscous. 
Wings lightly, uniformly infumated: costa, 
stigma, and venation brown. 

Antenna (apical 2 segments missing) 
with pedicel 1.5X longer than broad, as 
long as scape, segment 3 longer than 4, as 
4:3. Anterior margin of clypeus shallowly 
roundly emarginate; labrum broader than 
long, as 2:1 with rounded anterior margin 
(Fig. 3). Malar space linear. Inner margins 
of eyes converging below; LID:IDMO:EL 
= 2.0:2.5:2.0: bdL:POL:OCL = 1.3:1.0: 
1.0. Frontal area almost same height as 
eyes; supraantennal tubercles and frontal 
ridges indistinct; supraantennal pits distinct 
and removed from antennal sockets by 
1.75X diameter of pit; median fovea indis- 
tinct in its posterior half and in anterior half 
represented by a deep median supraantennal 
pit; without transverse ridge extending to 
inner margin of eye; circum- and interocel- 
lar furrows distinct, postocellar funow in- 

distinct; postocellar area subconvex. broad- 
er than long, as 2: 1 at its maximum width. 
Mesoscutellum flat, appendage not carinate. 
ITD:1CD = 4.0:1.0. Tarsal claws as in Fig. 

6. lATS:MB:OATS = 2.0:7.5:1.6. Abdo- 
men with brownish yellow pubescence. 
Lancet short, not strongly nan^owing toward 
apex, without lateral teeth above serrulae, 
with about 6 serrulae (Fig. 1 1 ). 

Male. — Average length. 4.5 mm. Similar 
to female except apical halves of coxae, 
basal half of fore- and midfemora, inner 
surface of hind femur on basal half, and 
apical tarsal segments black. Supraantennal 
pits removed from antennal sockets by 
0.5 X diameter of a pit. Genitalia as in Figs. 

7. 8; harpe triangular, pointed at apex; par- 
apenis produced into a nanow, rounded 
lobe at apex; penis valve oblong, finely ser- 
rate on dorsoapical margin. 

Types. — Holotype 2, Uttar Pradesh: 
Dhanoulti, 2000 m, 25.7.1993, coll. M.S. 
Saini. Paratypes: Same data as holotype ex- 
cept 3.6.1983 (2 (5. PUNJ, USNM). 

Distribution. — India (Uttar Pradesh). 

Etymology. — The name is derived from 
its type locality. 


We thank US PL-480, Project No. FG- 
In-753, and ICAR, New Delhi, for finan- 
cial assistance. Our thanks are extended to 
A. Taeger and S. M. Blank, Deutsches En- 
tomologisches Institut, Eberswalde, Ger- 
many for the loan of the type of Anapep- 
tcimena albipes. We appreciate reviews of 
the manuscript by the following: H. Gou- 
let. Agriculture and Agri-Food Canada. 
Ottawa, and R. A. Ochoa and J. W. 
Brown, Systematic Entomology Labora- 
tory. USDA, Beltsville, MD, and Wash- 
ington, DC, respectively. 

Figs. 1-11. Anapepttiiiwna. 1-3. Clypeus and labrum. 1. A. albipes. 2, A. ikirjeelingensis. 3, A. Jhanoul- 
licnsis. 4-6. Tarsal claw. 4. A. albipes. 5, A. darjeelingensis. 6. A. dhanoiiltiensis. 7. Male gonoforceps of A. 

(lluuuHilriensis. H. Male penis valve of. 4. dhantiuliicnsis. 9-1 I. Female lancets, 9. .4. 
gensis. 11.. 4. dhani>ullicnsis. 

ilhipes. 10. ,4. darjeelin- 



Literature Cited 

Cameron, P. 1899. III. Hymenoptera Orientalis, or con- 
tributions to a knowledge of the Hymenoptera of 
the Oriental Zoological Region. Part VIII. The 
Hymenoptera of the Khasia Hills. First paper. 
Memoirs and Proceedings of the Manchester Lit- 
erary and Philo.sophical Society 43: 1-51. 

Konow, F W. 1898. Neue Chalastogastra-Gattungen und 
Arten. Entomologische Nachrichten 24: 268-282. 

. 1900. Neuer Beitrag zur Synonymie der Ten- 

thredinidae. Wiener Entomologische Zeitung 19: 

Malaise. R. 1944. Entomological results from the 
Swedish Expedition 1934 to Burma and British 
India. Arkiv for Zoologi 35A: 1-58. 

. 1963. Hymenoptera Tenthredinoidea. subfam- 
ily Selandriinae. key to the genera of the world. 
Entomologisk Tijdskrift 84: 159-215. 

105(3). 2003. pp. 647-663 


Samuel A. Wells 

Depaitment of Bidagiicultural Sciences and Pest Management, Colorado State Univer- 
sity, Fort Collins, CO 8().'S2.^-1 177, U.S.A. (e-mail: samwellsCf^ 

Ahstnict. — Fifteen species, forineriy placed in Scmioius Eschscholt/ are tiansferred into 
the genus Semiotiims. which formerly included only S. haniiluuisi Pjatakowa. The 17 
known species of the Neotropical genus Seiniotiniis Pjatakowa are described and keyed. 
Sentiotinus maculatus, n. sp.. is described. Scmioius straminious Cande/e is a new syn- 
onym of S. hrevicollis Candeze, n. comb; S. clyi)tichiis Candeze and .V. siihvirescens 
Schwarz are new synonym.s of S. (/luulricollis Kirsch, n. comb.; S. juvenilis Candeze and 
5. horrei Candeze are new synonyms of .V. fusifonnis Kirsch, n. comb.; S. elciianlulus 
Candeze is a new synonym of S. triliiwiUus Candeze, n. comb.; S. pulchellus Candeze 
is a new synonym of 5. quadriviltis Steinheil, n. comb.; and .V. stiuidiniicri Pjatakowa is 
a new synonym of 5. aeneovituitus Kirsch. n. comb. The three genera of the subfamily 
Semiotinae are also keyed. 

Key Wonis: Coleoptera, click beetle, Eiateridae, Scmioius. Scmioiiuus 

The Net)trt)pica! click beetle genus Sc- 
miotiiius was proposed by Pjatakowa 
(1941) to include a single species, ,S'. Inui- 
gluiasi Pjatakowa from Colombia. It was 
distinguished from the related genus Scmi- 
oius Eschscholtz by having the pronotum 
wider than long, by the presence of carinae 
on the hind angles of the pronotum, and by 
the bulkier body. 

Unfortunately, the type and only known 
specimen of this species was destroyed in a 
fire in Kiev in the 1940's along with several 
hundred other Pjatakowa types (V.G. Dolin, 
personal communication). In the process ol 
revising the Neotropical genus Scmioius. it 
became clear that many of the smaller spe- 
cies assigned to that genus were similar to 
Pjatakowa's description of Scmioiiuus. A 
phylogenetic analysis of all species consid- 
ered to be included in Scmioius indicated a 
basal monophyletic clade for a group of 
species sharing characters with Scmioiiuus 

(Wells 2002). These species are herein as- 
signed to the genus Scmioiiuus. Addition- 
ally, one previously undescribed species is 
included in the genus. 

Of the 17 species here recognized, only 
a few are represented by more than a few 
specimens. A total of only ?>?i specimens 
were available for study from 26 major in- 
stitutions or museums. Accordingly, intra- 
specihc variability is not well understood 
and must be infeixed from the genus Sc- 
miolus. The best approach under these cir- 
cumstances is to be conservative in delim- 
iting species. 

Materials and Mi-thoos 

The following collections provided ma- 
terial for this study (involving both the gen- 
era Scmioius and Scmiotinits). The recog- 
nized four-letter codens are from Arnett et 
al. (199.^). Specimens o\' Scmioiiuus were 
present in only a few of these insliliilions. 


The Natural History Museum, London 
(BMNH); Calilomia Academy of Sciences. 
San Francisco (CASC): Canadian Museum 
of Nature. Ottawa (CMNC); Canadian Na- 
tional Collection of Insects. Ottawa 
(CNCI); Colorado State University, C.P. 
Gillette Museum of Arthropod Biodiversity. 
Fort Collins (CSUC); Cornell University 
Insect Collections. Ithaca. New York 
(CUIC); Deutsches Entomologisches Insti- 
tut. Eberswalde (DEIC); Escuela Agricola 
Panamericana. Zamorano, Tegucigalpa, 
Honduras (EAPZ); E.G. Riley Collection 
(EGRC); Essig Museum, University of Cal- 
ifornia, Berkeley (EMEC); Field Museum 
of Natural History, Chicago (FMNH); Hun- 
garian Natural History Museum, Budapest 
(HNHM); Instituto Nacional de Biodiver- 
sidad, Santo Domingo. Costa Rica (INBC); 
Inslitut Royal des Sciences Naturelles de 
Belgique. Brussels (ISNB): Museum fiir 
Naturkundc. Berlin (MNFD); Museum Na- 
tional d'Histoire Nalurelle. Paris (MNHN); 
Montana Entomology Collection, Montana 
State University, Bozeman (MTEC): Natur- 
historisches Museum, Vienna (NHMW); 
The Ohio State University Collection, Co- 
lumbus (OSUC); University of Oxford, 
Hope Entomological Collections, United 
Kingdom (OXUM); Peter Gate Collection, 
Vienna (PCCV); Snow Entomological Col- 
lections, University of Kansas, Lawrence 
(SEMC); Staatliches Museum fiir Tier- 
Kunde, Dresden (SMTD); Texas A&M 
University, College Station (TAMU); Na- 
tional Museum of Natural History, Smillv 
sonian Institution, Washington, D.C. 
(USNM); Universitetets Zoologiske Muse- 
um. Copenhagen (ZMUC). 

Male and female internal genitalia were 
examined after dissection and being cleared 
in a 1.0 molar solution of KOH at room 
temperature. Dissections were made on 
specimens from non-type material, except 
where available material was limited. The 
type material was too feeble to dissect. Col- 
or termini)logy was standardized by using 
the color charts in ,4 Dictioiuiry of Color 
(Maerz aiul Paul l*^).'S()) and then correlated 

vviih color descriptions found in W. T. 
Slerns Botaniccil Latin (1983). These 
sources should be consulted for precise 
identification of colors in this study. A gen- 
eral description of the main colors refeired 
to herein include aerugineus: verdigris; au- 
rantiacus: apricot orange: badius: dull choc- 
olate brown: fulvus: dull yellow with a mix- 
ture of gray and brown; luteus: buttercup 
yellow; piceus: black as pitch: rufopiceus: 
reddish black (in this study more black than 
reddish); sanguineus: blood red; testaceus: 
brick red to brownish yellow (of unglazed 
earthenware): viridis: untinted green. 

Measurements of body length were made 
from the anterior m;ugin of the frons (includ- 
ing frontal spines when present) to the tip of 
the elytra (including spines, when present). 
Measurements of body width were made at 
the widest part of the body — generally on the 
basal third of the elytra. Pronotal length was 
measured along the center-line of the prono- 
lum. PrcMiotal width was measured at the tip 
of the hind angles. Elytral length was mea- 
sured along the suture. Elytral width was 
measured at the widest point — generally at 
the basal third. The relative width of the eyes 
was determined using the ocular index 
(Campbell and Marshall 1964). This is deter- 
mined by measuring the nanowest distance 
between the eyes divided by the width of the 
head measured across the eyes. This i.|uotient 
is then multiplied by 100. 

Distributional data in the material ex- 
amined sections are from the specimen la- 
bels. Exceptions to this include the dates, 
which have been standardized so as to be 
read: day — month (in Roman numerals) — 
year In cases where these periods are un- 
certain, the data have remained exactly as 
indicated on the labels. Larger political di- 
visiiins have also been added in cases where 
they have not appeared on labels (and 
where they could be deteriuined). Duplicate 
specimens of the same species bearing 
identical labels are only listed once. All 
a\ailable type material was examined, in- 
cluding Chassain's recent lectotype desig- 
luiliims. The Pjatakowa types have been de- 

VOLUMi; 105. NUMBER 3 

slroyed (V.G. DdHii. personal LOiiinumiLa- 


The genus Semioliniis is distinguished 
from all other genera in the Semiotinae hy 
the straight or recurved lateral pronotal mar- 
gin (as in Figs. 1, 14-18) that joins the tloi- 
sal surface of the pronotum and the hyjio 
MieRin at an acute angle (Fig. 12). Other 
characters are also typical of the genus hut 
are not, in themselves, diagnostic for all spe- 
cies. These characters include: the hind an- 
gles of the pronotum that are typically cai- 
inate or subcarinate, the prosternum that is 
typically straight in profile, the Irons that is 
typically lobed and without projecting 
s|iines, the prt)notum that is typically wider 
liian long, the pronotal disk that is typically 
evenly convex across without depressions or 
(ubcrcles, and the elytra that each typically 
lertiiinate in an acute spine and either a sharp 
subapical angle or dentition. These charac- 
ters are not typical of Seiniotus. 

Within the genus itself, the primary char- 
acters used to distinguish species are the 
presence t)r absence of frontal spines, shape 
of the lateral margin of the pronotum, color 
of the head, pronotum, and elytra, and 
shape of the apical elytral spines. 

Male and female genitalic characters also 
ililTer interspecihcally. The male parameres 
range from narrow to broad and bear an 
apical blade (Figs. 2-9). This blade varies 
significantly in size and shape ranging from 
triangular with sharp angles (Figs. 2-4) to 
sinuous with rounded edges (Fig. 6). Pos- 
teriorly the blade has lateral angles that dif- 
fer in si/e and angle of projection. The an- 
terior sclerite of the female bursa copulatrix 
also differs in the only two species exam- 
ined. The pattern includes a median piece, 
or spine, flanked by two lateral arms, oi' 
spines (Figs. 10, I I ). 

Several species of Scmioln.s in the .V. ciir- 
(iiasaniis group (Wells 2002) are superfi- 
cially similar to the species of Semioliniis 
with frontal spines {S. macer {dmde/.c) and 
.S'. (U'lH'dvillalus (Kirsch)). Both groups lack 


marginal sulci. In these cases, species of Se- 
miotus can be recognized by the broader 
pronotal border (joining the hypomera in an 
even curvature (as in Fig. 13) and not 
acutely as in Semiotinus) and by the very 
finely punctate pronotum and elytra with 
faint striae. The pronotal punctures in Se- 
miotinus are deeper, and the striae and strial 
punctures are clearly defined. 

Key to Genera of Semiotinae 

1. Pronotum. viewed from above, with sides 
straight or recurved (Figs. 14-18); dorsal 
surface of pronotum meeting hypomeron at 
an acute marginal angle, without defining 
sulcus (Fig, 12) Semiotinus Pjatakowa 

- Pronotum, viewed from above, with sides 
sinuate or rounded (Fig. 19); lateral pronotal 
margin rounded, incrassate, undulate, or sul- 
cate, never acute (as in Fig. 13) 2 

2( 1 ). Tarsi with thick pubescent pads on segments 

1 to 3 Oistus Candeze 

- Tarsi with thin glabrous pads on segments 1 

to 3 Semiotus Eschscholt/ 

Genus Semiotinus Pjatakowa 

Semiotinus Pjatakowa 1941: 107 (type spe- 
cies, Semiotinus Bang-Haasi Pjatakowa, 
orig. desig.); Calder 1976: 107; Lawrence 
and Newton 1995: 853. 

Description. — Length 7-23 mm, colored 
variously with combinations of orange, yel- 
low, green, brown, and black. Frons with 
two spines or lacking frontal spines (as in 
Figs. 14—18). Antenna seirate extending 
one to four segments beyond hind angle. 
First two segments testaceus to piceus. re- 
maining segments fulvus to piceus. Prono- 
tum as wide as or wider than long, disk 
evenly convex; with or without setae, often 
glabrous, nitidus, punctation evenly scat- 
tered throughout, or becoining denser and 
umbilicate anterolaterally; lateral margins 
straight or recurved (in dorsal view), either 
parallel-sided or convergent anteriorly, 
without defining sulcus, pronotum and hy- 
pomeron joining at a sharp edge (Fig. 12), 
hind angles divergent or not. Hypomeron 
elongate, posterior border sinuous and often 
dentate, partially enclosing procoxae, Pros- 

ternal process lobed anteriorly, prosternal 
process straight or curved in profile, ex- 
tending between procoxae to mesosternal 
fossa. Scutellum subquadrate to elongate, 
longer than wide, flat or convex on disk. 
Legs fulvus to piceus, tarsal segments 1-3 
lobed beneath; tarsal claws broad at base, 
without basal setae. Elytron widest anteri- 
orly then narrowing to apex. Intervals flat 
or convex; elytral base on same plane as 
pronotum, occasionally gibbous and rising 
above level of pronotum; color piceus to 
fulvus, often with alternating light and dark 
bands; glabrous or pubescent: abdomen ful- 
vus to piceus, sternite 5 of females often 
with two foveae. Bursa copulatrix generally 
with two posterior platelike sclerites, and 
one spinose anterior sclerite. Parameres 
generally bearing an apical blade apically. 
Larvae and pupae unknown. 

Key to Species of Semiotinus Pjatakowa 

1. Head with two frontal spines (as in Fig. 

15) 2 

- Head without two frontal spines (as in 
Figs. 14, 16-18) 3 

2( I ). Pronotum and elytra pale yellow to light 

brown with greenish blue (often metallic) 

maculae or bands; apex of each elytron 

with two subequal spines (Pig, 22) , , , , 

aeneovittatus (Kirsch) 

- Pronotum and elytra light brown, if mark- 
ings present, then without greenish blue 
markings; apex of each elytron with a sin- 
gle spine and a subapical tooth or angle 
(Fig, 23) macer (Candeze) 

3( 1 ). Pronotum with three dark longitudinal 
vittae (sometimes confluent posteriorly) 

trilineatus (Candeze) 

Pronotum with 0, I . or 2 dark vittae ... 4 
4(3). Elytra unicolorous, at least on basal half 


Elytra not unicolorous, with light and 
dark contrasting vittae or patterns (or with 
pale elytra contrasting with infuscate stri- 
ae or strial punctures) 10 

5(4), Elytra black, or dark reddish black . , , , 

iiiniio (Candeze) 

Elytra dull yellow, yellowish green, or 

light brown to orange 6 

6(5). Pronotal margin (viewed from above) 

stfongly recurved (Fig. 14) 

conicollis (Candeze) 


Fig. 2-25. Semiouinis and Semioius specie.s. 2-^. Mule genitalia. 2. Seiuiotiiuis quuclricolUs. 3. Semiotinus 
fussiformis. 4. Semiotinus boliviensis. 5, Semiotinus virescens. 6, Semiotinus limhaticollis. 7. Semiotinus aeneov- 
ittatus. 8, Semiotinus maculatus. 9, Semiotinus scitulus. 10-1 1. Female anterior sclerite. 10, Semiotinus aeneov- 
ittatus. 1 1, Semiotinus macer. 12-13, Lateral aspect of pronotum. 12, Semiotinus scitulus. 13, Semiotus zonatus. 
14-19. Head and pronotum. 14, Semiotinus conicollis. 15, Semiotinus aeneovittatus. 16, Semiotinus nigricollis. 
17, Semiotinus virescens. 18, Semiotinus boliviensis. 19, Semiotus nigriceps. 20-25, Elytral apices. 20, Semi- 
otinus limhaticollis. 21, Semiotinus quadrivittis. 22. Semiotinus aeneovittatus. 23. Semiotinus macer. 24, Semi- 
otinus nigricollis. 25, Semiotinus boliviensis. 





Pronotal margin not strongly recurved (as 

in Figs. 1.17) 7 

Pronotiim with two black vittae. elytra 

gibbous at base brevicoUis (Candeze) 

Pronotum without black bands or with a 
single black band: elytra not gibbous at 

base 8 

Elytra (at least on basal half) orange to 

blood red inaciilatiis. n. sp. 

Elytra dull yellow to yellowish green 4 
Lateral pronotal margins straight, hind 
angles divergent (Fig. 17); pronotum im- 
maculate 17 

- Lateral pronotal margins straight and par- 
allel, not diverging posteriorly: pronotum 
maculate .... quadricollis (Kirsch). in part 

10(4). Elytral intervals all dull yellow, striae 
and/or strial punctures infuscate (dark 
brown to black) 11 

- Elytral intervals not all of same color 12 
1 1(10). Pronotum with one dark discal macula or 

vitta fiisifoniii.s (Kirsch) 

- Pronotum with two diirk discal vittae . . 

supplicaiis (Kirsch) 

12( 10). Each elytron with four dark and four pale 
vittae — intervals I, 3. 5. 7. and some- 
times lateral margin, brown, intervals 2, 
4, 6. 8 and 9 dull yellow 

scitnlu.s (Cande/.e) 

- Each elytron with fewer than four vittae 

of either color \?> 

13(12). Pronotum black, without yellow to light 

brown margins 16 

- Pronotum black with yellow to light 
brown margins, or with discal band only 
black 14 

14(13). Head black 15 

- Head orange to pale brown 

Lliiadricollis (Kirsch). in part 

15(14). Elytra gibbous at base: lateral margin of 
elytra concave in outline near apex (Fig. 
21) quadrivirris (Candeze) 

- Elytra not gibbous, as wide (dorsoventral- 
ly) at as on basal third; lateral margin 
of elytra evenly convex in outline near 
apex (Fig. 20) limbalicollis (Candeze) 

16(13). Pronotum narrow, length of anterior pro- 
notal margin approximately % width of 
elytral humeri (Fig. 18); apex of each el- 
ytron bispinose, area between spines 
emarginate (Fig. 25) holiviensis (Candeze) 

- Pronotum less narrow, length of anterior 
pronotal margin approximately % or more 
width of elytral humeri (Fig. 16): apex of 
each elytron blunt and angulate. if small 
dentitions present, then without emargi- 
nation (Fig. 24) nigricollis (Candeze) 

17(9). Body 4 mm wide at widest point 

binii^haasi Pjatakowa 

- Body less than 3.6 mm wide at widest 

point virescens (Candeze) 

Senuotinus aeneovittutiis (Kirsch). 

n. comb. 

(Figs. 7. 10. 15. 22) 

Semiotus aeneovitlatiis Kirsch 1884: 44 
(lectotype, female (Chassain, in press): 
SMTD; type locality: Ecuador, Huam- 

Semiotus sraiidingeri Pjntakowd 1941: 107 
(holotype destroyed; type locality: Ecua- 
dor. Norniandia). New synonym. 

Description, — Length 13-16 mm (length/ 
width ratio 4,4-4,8), Head badius to rufop- 
iceus on disk, fulvus to testaceus peripher- 
ally, with faint dark metallic sheen aerugi- 
neus, with two spines anterolaterally (Fig, 
15): fine decumbent setae scattered 
throughout, punctation subumbilicate and 
scattered throughout. Antenna serrate, 
reaching 2-3 segments beyond hind angle 
of pronotum in male, 1-2 segments in fe- 
male: segments 1 and 2 testaceus, segments 
3-1 1 rufopiceus. Thorax with pronotum 2- 
3 mm (length/width ratio 0,7-1.0) subtra- 
pezoidal, hind angles slightly diverging; 
margin thin, clearly defined; badius to ru- 
fopiceus with dark metallic sheen aerugi- 
nosus; fine scattered setae and deep punc- 
tures throughout, Scutellum fulvus to bad- 
ius, strongly convex, anterior margin de- 
clivitous and wider than rounded posterior 
margin, Prosternum linear in profile, apex 
of prosternal process not divided: fulvus to 
aurantiacus, with or without badius macula 
laterally; few erect setae anteriorly, denser 
and decumbent setae laterally, punctation 
fine medially, thicker laterally. Hypomeron 
fulvus to aurantiacus; glabrous, nitidus with 
fine decumbent setae and deep punctures 
near suture. Mesosternum depressed below 
plane of metasternum; fulvus to aurantia- 
cus. fossa glabrous and impunctate. lateral 
areas with dense setae and fine punctures. 
Mesepisternum and epimeron fulvus to au- 
rantiacus. Metasternum testaceus to auran- 


tiaciis with or without badius to piceus an- 
terolateral areas; mostly glabrous with few 
erect setae and with fine decumbent setae 
anterolaterally. punctation fine medially, 
wider and denser anterolaterally. Femora 
fulvus to testaceus. 

Elytron 9-12 mm (length/width ratio 
3.1-3.3). gibbous anteriorly, subparallel on 
anterior half then nanowing to tip; ranging 
from a metallic aerugineus with circum- 
scutellar area fulvus to luteus. to nearly en- 
tirely fulvus with fine aeruginous macula on 
intervals 6 and 8 basally; glabrous, nitidus. 
striae faint, strial punctures clearly defined; 
apex ending in a terminal spine and a large 
dentition along inner angle subapically 
(Fig. 22). 

Abdomen testaceus to aurantiacus; fine se- 
tae and punctation becoming thicker laterally: 
female without foveae on stemite 5; anterior 
sclerite of bursa copulatrix with lateral arms 
enlarged apically, each bearing 2 to 3 large 
dentitions, lateral arms 1 .2 times longer than 
median piece (Fig. 10). Male with parameres 
diverging apically, apical blades small, 20% 
length of parameres (Fig. 7). 

Material examined. — ECUADOR: Huam- 
boya. D. Stubel (lectotype of S. aneovittatiis. 
SMTD); Napo, Baeza, 2,000 m., 25-11-1979. 
H. & A. Howden (1, CNCl); Napo, 7 km S. 
Baeza, 2,000 m, (20-28 )-II- 1979. H. & A. 
Howden (1, CNCI). 

Diagnosis. — The two frontal spines (Fig. 
15) and the verdigris metallic sheen distin- 
guish S. aeneovittatus from all other Semi- 
oliniis. Seniiotinus nuicer also bears frontal 
spines but is readily distinguished by the 
brown body (and lack of verdigris metallic 
sheen), larger size, and more elongate and 
parallel-sided elytra. 

Note. — The type material of Seiuioiiis 
staiidingeri has been destroyed (Dolin. per- 
sonal communication). Pjatakowa's descrip- 
tion, though, does not differ from the type of 
5. aeneovittatus. There are no other known 
species of Semiotus (or Seniiotinus) that have 
the bluish >:reen metallic coloration. 

Seniiotinus banghaasi Pjatakowa 
Seniiotinus Bang-Haasi Pjatakowa 1941: 
107 (holotype: lost: type locality: Colom- 
bia: San Antonio) 

No specimens of this species were avail- 
able for study, but it apparently represents 
a valid species. The holotype was destroyed 
(V.G. Dolin. personal communication). The 
following description is interpreted from 
Pjatakowa (1941). 

Description. — Length 14 mm, 4 mm 
wide (length/width ratio 3.3). Head flat, ful- 
vus to testaceus. shallowly tri-impressed, 
sparsely punctate. Antenna long, extending 
to hind angle, with segments 1 and 2 Ha- 
votestaceus. segments 3-1 1 fuscus. Thorax 
with pronotum wider than long, nearly qua- 
drangular; testaceus with obscure fulvus to 
viridis border; moderate punctures, anterior 
angles rounded, hind angles short, little di- 
vergent, with a distinct carina. Scutellum 
oblong, anterior margin truncate, rounded 
behind, 3-angled, slightly wider posteriorly. 
Venter, except epipleura and lateral areas of 
prosternum luteus to aurantiacus. Femora 
fulvus to testaceus. 

Elytron convex, striate, striae piceus, 
punctate; fulvus to luteus with viridis tint; 
glabrous; apex emarginate. 

Material examined. — None. 

Note. — Pjatakowa's description of 5. 
banghaasi seems to place the species next 
to S. virescens. Both species are yellowish 
green and similar with respect to the char- 
acters listed by Pjatakowa in the description 
of S. banghaasi. The important difference 
between the two species is the width of the 
body, which in S. banghaasi is 4 mm. and 
in S. virescens, less than 3.6 mm. A speci- 
men (in the BMNH) labelled as 5. ban- 
ghaasi is too narrow and should be consid- 
ered S. virescens. 

Seniiotinus holiviensis (Candeze). 
n. comb. 
(Figs. 4. 18. 25) 
Semiotus boliviensis Candeze 1895: 16 (ho- 
lotype, male: INSB; type locality: Boliv- 


Description. — Length 10-11 mm (length/ 
width ratio 4.4-4.6). Head piceus, frontal 
margin with two angulate lobes anterolat- 
erally. without spines; long setae and iim- 
hilicate punctures throughout. Antenna ser- 
rate, reaching 3-4 segments beyond hind 
angle; badius lo piceus. Thorax with pron- 
otum 2 mm (length/width ratio 0.8-0.9) 
parallel sided on anterior half then strongly 
diverging to hind angles (Fig. 18), margin 
thin, hind angles carinate; piceus; long setae 
and deep punctation throughout, becoming 
denser peripherally. Scutellum subrectan- 
gular, longer than wide, covered with long 
setae. Prosternum nearly straight in profile; 
piceus; long setae and scattered punctures 
throughout. Hypomeron rufopiceus. Meso- 
sternum rufopiceus to piceus; long golden 
setae and scattered punctures throughout. 
Mesepisternum and metasternum rufopi- 
ceus to piceus; long aureus setae and scat- 
tered punctures throughout. Femora rufop- 
iceus to piceus. 

Elytron 8 mm (length/width ratio 3.2- 
3.4), subgibbous, subparallel on anterior % 
then narrowing to tip; badius to piceus with 
intervals 2-4 (or ."i) fulvus; nitidus, with 
short setae, intervals convex, striae and 
strial punctures clearly defined; apex ending 
in a terminal spine and a smaller dentition 
along inner angle subapically (Fig. 25). 

Abdomen badius to piceus; long aureus 
setae and scattered punctures throughout. 
Male with parameres sinuate along apical 
third (Fig. 4), apical blades subtriangular, 
approximately 20% length of parameres. 
Female unknown. 

Material examined.— BOLIVIA: (I, 

Diagnosis. — The completely black head, 
lack of frontal spines and very nanow an- 
terior aspect of the pronotum (Fig. 18), and 
recurved pronotal margins distinguish 5. 
holiviensis from all other Semiotinus. 

Semiotiniis hrevicollis (Candeze), 
n. comb. 
Semiotits hrevicollis Candeze 1857: 332 
(holotype: BMNH; type locality: N. Gra- 
nada); Cande/e 1874: 186. 

Semiotiis stramineiis Candeze 1857: 333 
(holotype: BMNH; type locality: Colom- 
bia); Candeze 1874: 188. New synonym. 

Description. — Length 13-17 mm (length/ 
width ratio 4.0-4.2). Head testaceus to au- 
rantiacus, front without spines; fine decum- 
bent setae scattered throughout and long 
erect setae over eyes and anterior margin, 
punctation scattered throughout; ocular in- 
dex 75.0-78.0. Antenna reaching 1-3 seg- 
ments beyond hind angle; segments 1 and 
2 testaceus, segments 3-11 badius. Thorax 
with pronotum 2-4 mm (length/width ratio 
0.7-0.9), sides straight, diverging to angles, 
hind angles subcarinate; margin thin, clear- 
ly defined; lateral and anterior margins tes- 
taceus, wide, sublateral bands piceus, me- 
dian area aurantiacus to badius; dense de- 
cumbent setae throughout, punctation deep, 
especially deep and umbilicate on piceus 
bands. Scutellum fulvus to testaceus, longer 
than wide, hind margin rounded, anterior 
margin straight. Prosternum linear in pro- 
file, tip of prosternal process not divided; 
testaceus to aurantiacus medially with ru- 
fopiceus macula along suture extending 
onto hypomeron; nearly glabrous and im- 
punctate medially, with erect setae and deep 
punctures anterolaterally. Hypomeron ful- 
vus to aurantiacus laterally, with rufopiceus 
band along suture extending nearly to lat- 
eral margin; with dense decumbent setae 
and punctures along suture, nearly glabrous 
with very fine punctures laterally. Meso- 
sternum testaceus to aurantiacus, fossa gla- 
brous and impunctate, lateral areas with 
dense setae and fine punctures. Mesepister- 
num testaceus to aurantiacus. Metasternum 
aurantiacus to sanguineous, badius to pi- 
ceus anterolaterally; with few setae medi- 
ally becoming dense anterolaterally, punc- 
tation fine medially becoming denser an- 
terolaterally. Femora testaceus. 

Elytron 10-12 mm (length/width ratio 
2.9-3.1), gibbous basally, subparallel on 
anterior % then narrowing to tip; fulvus, 
strial punctures darker; glabrous, nitidus, 
intervals convex, striae an strial punctures 



clearly defined; apex ending in a terminal 
spine and a smaller dentition along inner 
angle subapically. 

Ahdonien testaceus to aurantiacus with 
nebulous infuscation sublaterally; tine setae 
and punctation becoming denser laterally; 
I'emale without foveae on sternite 5. Male 

Material examined. — COLOMBIA: (1. 
BMNH); N. GRANADA (this could be 
anywhere in Ecuador. Colombia. Venezue- 
la, or Panama) (1, BMNH). 

Diagnosis. — Semiotinus hrevicollis is 
distinct from other Semiotinus in having un- 
icolorous elytra, a yellow to orange head, 
gibbous elytral base, and two pronotal vit- 
tae. The type of S. straiuineus is larger (be- 
ing just over 16 mm long) than other known 
examples of S. hrevicollis. No other differ- 
ences could be determined. The two are 
here considered conspecific. 

Seiiiioiiniis loiiicollis (Candeze), n. comb. 
(Fig. 14) 

Semioiiis conicollis Cuniieze 1857: 333 (ho- 
lotype: BMNH; type locality: Colombia); 
Candeze 1874: 188. 

Description. — Length 19—20 mm (length/ 
width ratio 3.7-3.9). Head fulvus to testa- 
ceus, frontal margin often angled without 
spines; nitidus with few decumbent setae 
scattered throughout; ocular index 66.0- 
68.0. Antenna serrate, reaching 3-4 seg- 
ments beyond hind angle; segments 1 and 
2 testaceus, segments 3-1 1 testaceus to 
badius. Thorax with pronotum 3-4 mm 
(length/width ratio 0.7-0.8) wider than 
long, sides strongly recurved (viewed from 
above), hind angles divergent (Fig. 14); tes- 
taceus to aurantiacus. darker on disk; gla- 
brous, nitidus. Scutellum testaceus to auran- 
tiacus, subtriangular Prosternum linear in 
profile; testaceus to aurantiacus, lobe paler; 
glabrous with fine punctation medially, 
denser setae and punctation laterally. Hy- 
pomeron fulvus; covered with fine setae 
and punctures throughout. Mesosternum 
fulvus to testaceus. fossa glabrous and im- 

punctate, lateral areas with dense setae and 
fine punctures. Mesepisternum testaceus tci 
badius. Metasternum testaceus to badius; 
sparse erect setae medially, denser decum- 
bent setae laterally on dark area. Femora 
aurantiacus to testaceus. 

Elytron 15 mm (length/width ratio 2.8- 
3.0) gibbous; unicolorous fulvus with faint 
viridis tint; glabrous, nitidus, striae faint to 
obsolete, strial punctures clearly defined; 
apex bearing two small spines or dentitions. 

Ahdonien testaceus; scattered erect setae 
medially, dense decumbent setae laterally, 
punctation fine becoming dense laterally. 
Genitalia not examined. 

Material examined.— COLOMBIA: (1. 

Diagnosis. — The unicolorous elytra, re- 
curved pronotal margin (Fig. 14), and yel- 
low to light brown head readily distinguish 
S. conicollis from ail other Semiotinus. 

Semiofiiuis fnsljbrmis (Kirsch). n. comb. 
(Fig. 3) 

SemiotHs fiisiformis Kirsch 1866: 181 (lec- 
tt)type, female (Chassain. in press): 
SMTD; type locality: Bogota); Steinheil 
1875; 113; Candeze 1874: 185. 

Semiotus Juvenilis Cixndeze 1874: 1X7 (ho- 
lotype: BMNH; type locality: Colombia). 
New synonym. 

Semiotus borrei Candeze 1878: 9 (type: not 
found; type locality: Ecuador). New syn- 

Description. — Length 12-13 mm (length/ 
width ratio 4.8-4.9). Head piceus, front 
without spines; fine decumbent setae and 
shallow subumbilicate punctures through- 
out. Antenna reaching 3-4 segments be- 
yond hind angle in male; segments 1 and 2 
testaceus, segments 3-11 rufopiceus to pi- 
ceus, Thora.x with pronotum 2-3 mm 
(length/width ratio 0.8.-0.9) subtrapezoidal, 
hind angles diverging; margin thin, clearly 
defined; fulvus to aurantiacus, with ellipti- 
cal piceus macula medially; glabrous. Scu- 
tellum badius to piceus, longer than wide, 
posterit)r border roimded, anterior margin 


straight. Prosteinum linear in profile, tip of 
prosternal process not divided; piceiis, pros- 
ternai lobe and prosternal process fulvus to 
aurantiacus; few erect setae anteriorly, de- 
cumbent setae laterally, punctation fine me- 
dially, thicker laterally. Hypomeron fulvus 
to luteus, piceus laterally along suture; ni- 
tidus with or without few scattered setae 
throughout, punctation of few shallow sub- 
umbilicate punctures throughout. Mesoster- 
num fulvus to testaceus medially, piceus 
laterally, fossa glabrous and impunctate. 
lateral areas with dense setae and fine punc- 
tures. Mesepisternum piceus. Metasternum 
badius to piceus throughout or with fulvus 
to testaceus median coloration; glabrous 
with fine punctation medially, with fine de- 
cumbent setae and denser punctation an- 
terolaterally. Femora fulvus to testaceus. 

Elytron 8-10 mm (length/width ratio 
3.5-3.7). subparallel on anterior half then 
nan\)wing to tip; fulvus to luteus with strial 
punctures badius to rufopiceus; glabrous, 
nitidus. striae and strial punctures clearly 
defined, intervals convex; apex ending in a 
terminal spine and a smaller dentition along 
inner angle subapically. 

Abdomen badius to rufopiceus. sternite 5. 
lateral areas of sternites 3 and 4 and medial 
area of sternite 1 lighter; fine setae and 
punctation becoming thicker laterally. Male 
with parameres subtriangular apically. lat- 
eral margins convex, aedeagus extending 
beyond apical tips of blades (Fig. 3). 

Material examined.— COLOMBIA: Bo- 
gota. Kirsch (1. SMTD); ECUADOR: 
Pich., 35 km E Tandapi. 8,000-9.0{)() ft.. 
24-IV-1975. S. & J. Peck (1. CNCI); E. de 
Ville. 71(1. INSB). 

Diagnosis. — The single pronotal band 
and fulvus elytra with infuscate punctures 
separate S. fiissifonnis from all other Se- 

Note. — The type of 5. borrei has not 
been located. A specimen with the author's 
(Candeze) determination label was located 
in Brussels (INSB). This specimen is clear- 
1\ a synonym of 5. fnsiformis. 

Seniioiinii.s UuibaticolUs (Candeze). 

n. comb. 

(Figs. 6. 20) 

Scniiotiis linibaticollis Candeze 1857: 336 
(holotype: BMNH; type locality: Colom- 
bia): Candeze 1874: 188. 

Description. — Length 9-10 mm (length/ 
width ratio 4.1-4.3). Head piceus throughout 
without spines; few long setae and scattered 
shorter setae, punctation subumbilicate. An- 
tenna serrate, reaching 3-4 segments beyond 
hind angle; piceus. Thorax with pronotum 2 
mm (length/width ratio 0.8-1.0), subquadrate, 
mtugins straight, gradually widening posteri- 
orly, hind angles subcarinate; piceus, with 
thin fulvus to aurantiacus margins becoming 
wider near angles. Scutellum badius to rufop- 
iceus, parallel-sided, angles rounded, longer 
than wide. Prostemum linear in profile; pi- 
ceus, lobe and prosternal process fulvus to 
testaceus; long pale setae, punctation becom- 
ing umbilicate laterally. Hypomeron piceus, 
with lateral mtugin fulvus to aurantiacus; 
long pale setae, punctation close and umbil- 
icate throughout. Mesostemum aurantiacus 
medially, piceus laterally, covered with long 
pale setae. Mesepisternum piceus. Metaster- 
num piceus; glabrous medially, with dense 
decumbent setae laterally, punctation fine be- 
coming dense laterally. Femora fulvus to tes- 

Elytron 6-8 mm (length/width rafio 3.0- 
3. 1 ). subparallel on anterior % then narrowing 
to tip; badius to rufopiceus with intervals 1- 
4. 9, and margin fulvus to luteus on basal %; 
glabrous, nitidus, with fine setae laterally, 
strial punctures well defined, striae fine to ob- 
solete; apex ending in one spine apically and 
a smaller denfition subapically (Fig. 20). 

Abdomen piceus. with lateral margin 
badius. sternite 5 testaceus; pale setae and 
fine punctation throughout. Male with par- 
ameres sinuate, apical blades with lateral 
margins strongly convex (Fig. 6). 

Material examined.— COLOMBIA: (1. 
BMNH): ECUADOR: Chiquinda. Buckley 
(1. BMNH). 

Diaiinosis. — The combination of the 


black head lacking frontal spines, contrast- 
ing elytral bands or patterns (fewer than 4 
of each color per elytron), and evenly con- 
vex elytral margins distinguish S. limbati- 
collis from all other Semiotinits. 

Seinlotlitus nuicer (Candeze). n. comb. 
(Figs. 1 1. 23) 

Sciuiotiis nuicer Candeze 1889: 80 (holo- 
type. female: INSB; type locality: Co- 

Description. — Leni^th 21 mm (length/ 
width ratio 4.5-4.7). Head testaceus to au- 
rantiacus. with two spines anterolaterally; Hne 
decumbent setae and deep punctation 
throughout with longer setae over spines; oc- 
ulai- index 64.0-66.0. Antenna serrate, reach- 
ing 2-3 segments beyond hind angle; testa- 
ceus to piceus. Thorax with pronotum 4 mm 
(length/width ratio 1.0), sides straight, nar- 
rowly diverging posteriorly; margin thin, 
clearly defined; testaceus to aurantiacus, with 
piceus band extending from base to near 
apex; glabrous, nitidus with punctures widely 
separated. Scutellum strongly declivitous an- 
teriorly, without clear anterior margin, lateral 
margins convex; testaceus to badius. Prester- 
num concave in profile, not divided at apex; 
testaceus to aurantiacus with or without bad- 
ius to piceus maculae along suture; glabrous, 
nitidus medially with fine setae and deep 
punctures laterally. Hypomeron fulvus to au- 
rantiacus; glabrous laterally with fine setae 
and deep punctures along suture. Mesoster- 
num aurantiacus to badius, fossa glabrous and 
impunctate, lateral areas with dense setae and 
fine punctures. Mesepistemum badius to pi- 
ceus. Metastemum testaceus to rufopiceus. 
Femora aurantiacus to badius. 

Elytron 15 mm (length/width ratio 3.2- 
3.4), long and naiTOw. subparallel on ante- 
rior % then rounded to tip; testaceus, strial 
punctures infuscate; mostly glabrous, niti- 
dus with occasional setiferous punctures, 
intervals nearly flat, striae obscure to ob- 
solete, strial punctures pronounced; apex 
ending in a terminal small spine and a 

smaller dentition or sharp angle along su- 
ture subapically (Fig. 23). 

Abdomen testaceus to aurantiacus; fine 
setae and dense punctation throughout; fe- 
male with two elliptical and very shallow 
piliferous foveae in center of sternite 5; an- 
terior sclerite of the bursa copulatrix with 
lateral arms enlarged apically, each bearing 
two to three large dentitions, lateral arms 
1.2 times longer than median piece (Fig. 
1 1 ). Male unknown. 

Material examined.— COLOMBIA: (1. 

Diagnosis. — The very elongate and par- 
allel-sided body (at least 4.5 times longer 
than wide) in combination with the two 
frontal spines distinguish S. macer from all 
other Semiotinus. Semiotiniis aeneovittatus 
also bears frontal spines but is less elongate, 
and has a metallic bluish maculae on the 
pronotum and elytra which contrasts with 
the fulvus to testaceus base color. Semiotin- 
us macer is darker testaceus and lacks me- 
tallic blue maculae. 

Semiotinus maciilatus Wells, new species 

(Figs. 1, 8) 

Description. — Length 15 mm (length/ 
width ratio 4.1-4.2). Head aurantiacus with 
basal piceus macula, without spines; gla- 
brous, nitidus with few erect setae above 
eyes and anterior margin, punctation deep 
and scattered throughout. Antenna extend- 
ing three segments beyond hind angle in 
male; segments I and 2 testaceus. segments 
3-1 1 piceus. Thorax with pronotum 3-4 
mm (length/width ratio 1.0-1.1); subtrape- 
zoidal. sides straight, posterior margin wid- 
er than anterior margin, hind angles hardly 
divergent; margin narrowly incrassate with- 
out sulcus; aurantiacus to testaceus with 
narrow piceus band medially; glabrous, ni- 
tidus with deep punctation scattered 
throughout; scutellum piceus, narrowly 
ovate, nearly twice as long as wide; pros- 
ternum nearly straight in profile; aurantia- 
cus to testaceus medially, piceus laterally: 
glabrous with hne punctation medially and 
with decumbent and erect setae and deep 


punctation laterally. Hypomeron aurantia- 
ciis to testaceiis with piceus band along su- 
ture; glabrous, nitidus laterally with tine se- 
tae and deep punctation medially. Meso- 
sternum finely punctate with fine setae, fos- 
sa glabrous and impunctate; aurantiacus to 
testaceus medially, piceus laterally. Mese- 
pisternum piceus. Metasternum aurantiacus 
to testaceus medially, piceus laterally; gla- 
brous with fine punctation medially, with 
decumbent setae and denser punctatit>n lat- 
erally. Femora fulvus to testaceus. 

Elytron 10-11 mm (lengtli/width ratio 
2.9-3.0) subparallel on anterior half then nar- 
rowing to tip; testaceus to aurantiacus with 
interval 1 infuscate on posterior half; gla- 
brous, nitidus, striae and strial punctures deep 
to moderately deep; apex bearing one spine. 

Abdomen aurantiacus to testaceus medi- 
ally, badius laterally; nearly glabrous with 
fine punctation medially, with decumbent 
setae and denser punctation laterally; fe- 
male unknown. Male with parameres nar- 
row medially, apical blades 0.3 times length 
of parameres (Fig. 8). 

Material examined. — Holotype ( i ): EC- 
UADOR: Napo, Baeza. 2000 m. 25-11- 
1979. H. & A. Howden (CNCI). Paratype: 
ECUADOR: Chiquinda, Buckley (1 cJ, 

Diagnosis. — Semiotiis maculaius can be 
separated from all other Seniiotiiuis by the 
uniform orange to light brown dorsal color 
(except the piceus scutellum and pronotal 
and frontal band and the darker infuscation 
on the apical half of the elytra in the para- 
type) and the concave outline of the elytral 
margin near the apex. 

Note. — The specific name 'imuuUitiis' 
refers to the maculate pronotum. 

Scniioriniis morio (Candeze). n. comb. 

Seniiotiis morio Candeze 1857: 336 (holo- 
type: BMNH; type locality: Colombia); 
Candeze 1874: 189. 

Description. — Length 10-11 mm (length/ 
width ratio 3.5-3.6). Head piceus throughout 
without spines; few long setae and subum- 

bilicate punctures. Antenna serrate, reaching 
3-4 segments beyond hind angle; piceus. 
Thorax with pronotum 1-3 mm (length/width 
ratio 0.8-0.9) subcompanulate. lateral margin 
curving inwards to almost straight; margin 
thin, without sulcus, hind angles carinate; pi- 
ceus; nitidus, with long pale setae laterally. 
Scutellum piceus, subtriangular, anterior mar- 
gin not defined. Prosternum linear in profile; 
piceus; vestiture double, punctation becoming 
umbilicate laterally. Hypomeron piceus; long 
pale setae, punctation close and umbilicate 
throughout. Mesostemum piceus, vestiture 
double. Mesepistenium piceus. Metasternum 
piceus; vestiture double, punctation fine 
throughout. Femora testaceus to rufopiceus. 

Elytron 8-9 mm (length/width ratio 
2.3-2.5), subparallel on anterior Vi then nar- 
rowing to tip; deep sanguineous to rufopi- 
ceus throughout; glabrous, nitidus, with fine 
setae laterally, striae and strial punctures 
well defined; apex ending in two dentitions 

Abdomen piceus, without lateral pale ar- 
eas, sternite 5 with or without testaceus to 
sanguineous patches; vestiture double, 
punctation fine throughout. Genitalia not 

Material examined.— COLOMBIA: (1, 

Diagnosis. — The small size and black to 
reddish black coloration of the body sepa- 
rate S. morio from all other Semiotiniis. 

Semiotiniis nigricollis (Candeze), n. comb. 
(Figs. 16, 24) 

Semiotus nigricollis Candeze 1857: 335 
(holotype: BMNH; type locality: N. Gra- 
nada); Candeze 1874: 188. 

Description. — Length 13 mm (length/ 
width ratio 3.7-3.9). Head piceus through- 
out without spines; few long setae and sub- 
umbilicate punctures; ocular index 67.0- 
69.0. Antenna serrate, reaching 3-4 seg- 
ments beyond hind angle; piceus. Thorax 
with pronotum 2-3 mm (length/width ratio 
0.7-0.9) subcompanulate. lateral margin 
curving inwards to almost straight (Fig. 


16); margin thin, without sulcus, hind an- 
gles carinate; piceus; nitidus, with long pale 
setae laterally. Scutellum piceus. longer 
than wide, subrectangular. Prosternum lin- 
ear in profile; piceus; long pale setae, punc- 
tation becoming umbilicate laterally. Hy- 
pomeron piceus; long pale setae, punctation 
close and umbilicate throughout. Mesoster- 
num piceus. covered with long pale setae. 
Mesepisternum and mesepimeron piceus. 
Metasternum piceus; glabrous medially, 
with long setae laterally, punctation fine 
throughout. Femora piceus. 

Elytron 10-11 mm (length/width ratio 
2.9-3.1), subgibbous anteriorly; interval 1 
rufopiceus; intervals 2-5 (and part of 6 ba- 
sally) fulvus to luteus. intervals 7-9 rufop- 
iceus; glabrous, nitidus. with fine setae lat- 
erally, strial punctures well defined, striae 
fine to obsolete; apex ending in two denti- 
tions apically (Fig. 24). 

Abdomen piceus. with lateral margin and 
all of sternite 5 testaceus to sanguineous; 
pale setae and fine punctation throughout. 
Genitalia not examined. 

Material examined. — N. GRANADA 
(this could be anywhere in Ecuador. Colom- 
bia, Venezuela, or Panama) (1, BMNH). 

Diagnosis. — The blunt elytral apices 
(Fig. 24) and completely black pronotum 
distinguish S. nigricoUis from all other Sc- 

Seniiotiniis qmulricollis (Kirsch). n. comb. 
(Fig. 2) 

Sewintiis qiiadricollis Kirsch 1866; 181 

(lectotype. female (Chassain, in press); 

SMTD; type locality: Bogota); Candeze 

1874: 187, 
Semiotus diptychus Candeze 1874: 188 (ho- 

lotype: BMNH; type locality; Ecuador). 

New synonym. 
Semiotits subvirescens Schwarz 1904: 49 

(syntype (1 found), female: DEIC; type 

locality: Ecuador. Balzapamba). New 


Description, — Length 12-15 mm (length/ 
width ratio 4.2-4,3), Head testaceus to au- 

rantiacus, front without spines; fine decum- 
bent setae scattered throughout and long 
erect setae over eyes and anterior margin, 
punctation scattered throughout; ocular in- 
dex 69.0-72.0. Antenna reaching (J-2 seg- 
ments beyond hind angle in female; seg- 
ment one testaceus, segments 2-1 1 piceus. 
Thorax with pronotum 2-3 mm (length/ 
width ratio 0,9-1.0), subtrapezoidal, nearly 
straight, hind angles hardly diverging, car- 
inate; margin thin, clearly defined; aurantia- 
cus with lateral margins paler, with median 
Tiiacula piceus; dense decumbent setae 
throughout, punctation deep becoming 
nearly contiguous along lateral margin. 
Scutellum testaceus, longer than wide, pos- 
terior border rounded, anterior margin 
straight. Prosternum linear in profile to 
slightly concave, tip of prosternal process 
not divided; testaceus to aurantiacus medi- 
ally with or without a rufopiceus macula 
along suture extending onto hypomeron; 
nearly glabrous and impunctate medially, 
with erect setae and deep punctures antero- 
laterally. Hypomeron fulvus to aurantiacus 
laterally, occasionally rufopiceus along su- 
ture; with dense decumbent setae and punc- 
tures along suture, nearly glabrous with 
very fine punctures laterally. Mesosternum 
testaceus to aurantiacus. fossa glabrous and 
impunctate, lateral areas with dense setae 
and fine punctures. Mesepisternum testa- 
ceus to aurantiacus, Metasternum testaceus 
to aurantiacus medially, occasionally badius 
\o piceus anterolaterally; with few setae me- 
dially becoming dense anterolaterally, 
punctation fine medially becoming denser 
anterolaterally. Femora testaceus. tibiae and 
tarsi darker infuscate. 

Elytron 8-1 1 mm (length/width ratio 
2.9-3, 1 ). subparallel on anterior % then nar- 
rowing to tip: fulvus. strial punctures dark- 
er, with or without darker humeral bands; 
glabrous, nitidus, intervals convex, striae 
and strial punctures clearly defined; apex 
ending in a terminal spine and a smaller 
dentition along inner angle subapically. 

Abdomen testaceus to aurantiacus with 
nebulous infuscation sublaterally; fine setae 


and punctation throughout; female without 
foveae on stemite 5. Male with apical blade 
of parameres suhtriangular. lateral margins 
convex (Fig. 2). 

Material examined. — COLOMBIA: (3. 
BMNH; 1. ISNB); N. de S. 2600 m. 
30 km S Chinacota, 14-V-1974, H. & A. 
Howden (1, CNCl): ECUADOR: Balza- 
pamba, R. Haensch S. (1, DEIC). 

Diagnosis. — Semiothms qiiadricoUis is dis- 
tinct from other Semiotinus in having a su- 
bquadrate pronotum with a yellow to orange 
head and a single pronotal band. Semiotinus 
conicollis and S. hrevicoUis also have yellow 
to light brown heads but have two dark pro- 
notal bands (S. brevicoUis) or have a strongly 
recurved pronotum (S. conicollis). 

Note. — As with most other Semiotinus, 
S. qiuulricollis is known from very few 
specimens. Only six specimens were seen 
in this study, two of which are the types of 
the synonyms 5. subvirescens and S. dip- 
Tychus. The venter of the type of S. dipty- 
chiis is paler and the frontal macula is dark- 
er than in S. quadricollis. This difference, 
however, has not proven to be of value 
when comparing variability of other Semi- 
otinus nor in the sister genus Semiotus. 

Semiotinus quadririttis (Steinheil), 
n. tomb. 

(Fig. 21) 

Semiotus quadrivittis Steinheil 1875: 113 

(lectotype (Chassain, in press): MNHN; 

type locality: N. Granada). 
Semiotus pulchellus Candeze 1889: 81 (ho- 

lotype: INSB; type locality: Colombia). 

New synonym. 

Description. — Length 12-16 mm (length/ 
width ratio 3.9-4.0). Head piceus through- 
out, front without spines: fine decumbent 
setae scattered throughout; ocular index 
64.0-66.0. Antenna serrate, reaching 2-3 
segments beyond hind angle; segments 1 
and 2 testaceus. segments 3-11 rufopiceus 
to piceus. Thorax with pronotum 2-4 mm 
(length/width ratio 0.9-1.1), margins 
straight, subparallel, hind angles hardly di- 

verging; margin thin, clearly defined; fulvus 
to aurantiacus laterally with large maculae 
on disk extending from base to (or near to) 
anterior margin; glabrous or with very fine 
setae scattered throughout. Scutellum tes- 
taceus to piceus. longer than wide. Proster- 
num nearly linear in profile; piceus, pros- 
ternal process and lobe testaceus to auran- 
tiacus. Hypomeron badius to piceus along 
suture, fulvus to luteus laterally. Mesoster- 
num testaceus to aurantiacus medially, pi- 
ceus laterally, fossa glabrous and impunc- 
tate, lateral areas with dense setae and fine 
punctures. Mesepisternum piceus. Metaster- 
num testaceus to aurantiacus medially, pi- 
ceus laterally (or completely piceus). Fem- 
ora testaceus to piceus. 

Elytron 8-12 mm (length/width ratio 
2.8-2.9), slightly gibbous; fulvus to auran- 
tiacus with three piceus bands (one along 
interval 1, one around interval 8, and one 
band between intervals 1 and 8); glabrous, 
nitidus; apex ending in a terminal spine and 
a smaller dentition along inner angle suba- 
pically (Fig. 21). 

Abdomen testaceus to aurantiacus with 
scattered piceus maculae. Genitalia not ex- 

Material examined. — COLOMBIA: Me- 
dellin. 1915-38 (1, BMNH); N. Granada (1, 
INSB; 1, MNHN). 

Diagnosis. — The dark brown to black 
pronotal band with yellow to light brown 
margins, fewer than four alternating light 
and dark elytral bands, gibbous elytra, and 
narrow (concave in outline) elytral apices 
distinguish S. quadrivittis from all other Se- 

Seniiotinus scitulus (Cande/e). n. comb. 
(Figs. 9, 12) 

Semiotus scitulus Candeze 1874: 186 (ho- 
lotype; BMNH; type locality; Ecuador). 

Description. — Length 12-13 mm (length/ 
width ratio 4.0-4.4). Head piceus through- 
out or with anterolateral angles aurantiacus. 
front without spines; fine decumbent setae 
scattered throughout, punctation subumhil- 



icate and scattered throughout; ocular in- 
dex 64.0-67.0. Antenna seirate, reaching 
2-3 segments beyond hind angle in male; 
segments 1 and 2 testaceus, segments 3-1 1 
rufopiceus to piceus. Thorax with pronotum 
2-4 mm (length/width ratio 0.8-1.1). sub- 
trapezoidal, hind angles hardly diverging; 
margin thin, clearly defined (Fig. 12); ful- 
vus to aurantiacus with large suborbicular 
to subquadrate piceus macula on disk (often 
pointed apically): glabrous or with Hne se- 
tae scattered throughout. Scutellum badius 
to piceus, longer than wide, posterior bor- 
der rounded, anterior margin straight. Pros- 
ternum linear in profile, tip of prosternal 
process not divided; piceus, prosternal pro- 
cess and lobe fulvus to testaceus; few erect 
setae throughout becoming denser and de- 
cumbent laterally, punctation fine medially, 
thicker laterally. Hypomeron badius to pi- 
ceus along suture, fulvus to luteus laterally; 
glabrous, nitidus, or with few scattered se- 
tae, punctation of few shallow subumbili- 
cate punctures throughout. Mesosternum 
fulvus to testaceus medially, piceus later- 
ally, fossa glabrous and impunctate. lateral 
areas with dense setae and fine punctures, 
Mesepisternum piceus. Metasternum badius 
to piceus throughout or with fulvus to tes- 
taceus median coloration; glabrous with 
fine punctation medially, with fine decum- 
bent setae and denser punctation anterolat- 
erally. Femora aurantiacus to testaceus. 

Elytron 9-10 mm (length/width ratio 
2.9-3.1), subparallel on anterior half then 
nanowing to tip; fulvus to luteus with in- 
tervals 1, 3, 5, 7, and lateral margin badius 
to piceus; glabrous, nitidus, striae faint, 
strial punctures clearly defined; apex ending 
in a terminal spine and a smaller dentition 
along inner angle subapically. 

Abdomen aurantiacus to rufopiceus with 
fulvus areas on sternites 1 and 5; fine setae 
and punctation becoming thicker laterally; 
female not seen. Male with parameres nar- 
rowing apically before blades, apices diver- 
gent, apical blades small, less than 20% 
length of parameres (Fig. 9). 

Material examined.— ECUADOR: (I, 

BMNH); VENEZUELA: Merida. Tabay, 
Mucuy, Send. Lag. Suero cloud forest, 225 
m, (VI-VII)-1989, S & J Peck (1. CMNC); 
Merida, S. Briceno (1, USNM). 

Diagnosis. — The presence of alternating 
light and dark elytral bands (4 each per el- 
ytron) distinguishes 5. scituhts from all oth- 
er Semiotinus. 

Semiotinus sitpplivans (Kirsch), n. comb. 

Semiotiis supplicans Kirsch 1884: 45 (lec- 
totype, male (Chassain, in press): SMTD; 
type locality: Colombia, Pasto, 2800 m). 

Description. — Length 18 mm (length/ 
width ratio 2.9-3.1). Head fulvus to testa- 
ceus, without piceus maculae; frons with 
anterolateral areas rounded to angled, with- 
out spines; scattered setae throughout. An- 
tenna serrate, reaching 2-3 segments be- 
yond hind angle; segments I and 2 testa- 
ceus, segments 3-1 I piceus. Thorax with 
pronotal margins straight, hind angles not, 
or hardly, diverging; fulvus to testaceus 
with two sinuate piceus bands sublaterally; 
with dense setae, especially over piceus 
bands. Scutellum subtriangular, fulvus to 
testaceus. Femora testaceus. 

Elytron subparallel on anterior -A then 
narrowing to tip; fulvus to testaceus with 
strial punctures infuscate; glabrous, nitidus, 
striae and strial punctures distinct; apex 
bearing two subequal spines. Male genitalia 
not examined. 

Material examined. — COLOMBIA: Pas- 
to, 2800 m., D. Stubel (1, SMTD). 

Diagnosis. — The uniform pale elytral 
color interrupted by infuscate strial punc- 
tures, the two pronotal bands, and the two 
subequal spines on each elytron separate S. 
supplicans from all other Semiotinus. 

Semiotinus trilineatus (Candeze), n. comb. 

Semiotus trilineatus Candeze 1857: 324 
(syntypes (3), female: BMNH; type lo- 
cality; Colombia); Steinheil 1875; 112; 
Candeze 1874: 182. 

Semiotus elegantulus Candeze 1857: 325 
(syntypes (2): BMNH; type locality: Co- 


lombia): Caiulc/c 1874; 182. New syn- 

DcsLiipfion. — Leiifith 1 1-15 turn (length/ 
width ratio 4.2-4.6). Head fulvu.s to testa- 
ceus. frontal margin often angled without 
spines: nitidiis with few decumbent setae 
scattered throughout; ocular index 72.0- 
15.0. AiitciiiKi serrate, extending 0-1 seg- 
ments beyond hind angle in female; seg- 
ments 1 and 2 testaceus, segments 3-1 1 tes- 
taceus to badius. Thorax with pronotum 2- 
4 mm (length/width ratio 0.9-1.2). su- 
bquadrate, lateral margins straight, hind an- 
gles diverging; fulvus to aurantiacus with 
long thin median band (extending length of 
pronotum) and two sublateral shorter and 
sinuate badius to rufopiceus bands, hind an- 
gles often obscure; glabrous, nitidus. Scu- 
tellum fulvus to testaceus, longer than wide, 
posterior border rounded. Prosternum linear 
HI prolile; testaceus to aurantiacus. lobe pal- 
er, with or without piceus marking along 
suture; glabrous with fine punctation me- 
dially, denser setae and punctation laterally. 
Hypomeron fulvus, hind angles with or 
without piceus macula; glabrous, nitidus, 
punctation very fine throughout. Mesoster- 
num testaceus, fossa glabrous and impunc- 
tate, lateral areas with dense setae and fine 
punctures. Mesepisternum testaceus. Metas- 
ternimi testaceus, often piceus anterolater- 
ally; sparse erect setae medially, denser de- 
cumbent setae laterally on dark area. Fem- 
ora aurantiacus to testaceus. 

Elytron 8-12 mm (length/width ratio 
2.9-3.6), subparallel on anterior % then nar- 
rowing to tip; intervals 2, 4, 6, and 8 fulvus, 
intervals I, 3, 5, and 7 either fulvus vitreus 
or badius vitreus; glabrous, nitidus, striae 
faint to obsolete, strial punctures clearly de- 
hned; apex ending in a terminal spine and 
a smaller ilentition along inner angle suba- 

Abdomen testaceus medially, badius lat- 
erally; scattered erect setae medially, dense 
decumbent setae laterally, punctation fine 
becoming dense laterally; female without 
foveae. Male uenitalia not examined. 

Material examined.— COLOMBIA: (5, 
BMNH); Bogota (2, BMNH). 

Diagnosis. — The presence of three dis- 
tinct pronotal vittae (sometimes confluent 
basally) distinguishes S. trilineaiiis from all 
other Seniiotiniis. 

Seniioiliiii.\ \'ire.\cen.s (Candeze), n. comb. 

(Figs. 5, 17) 

Semiotiis virescens Cimdc/.c 18.S7: 331 (ho- 
lotype, female: BMNH: type locality: Co- 
lombia): Candeze 1874: 185. 

FJescription. — Length 13 mm (length/ 
width ratio 3.9-4.0). Head fulvus to auran- 
tiacus, front without spines: glabrous, niti- 
dus with few erect setae over eyes and an- 
terior margin, punctation deep and scattered 
throughout: ocular index 72.0-75.0. Anten- 
na seiTate, extending 1-2 segments beyond 
hind angle in male: segments I and 2 tes- 
taceus, segments 3-1 1 piceus. Thorax with 
pronotum 2-3 mm (length/width ratio 0.7- 
0.8), with sides nearly straight, hind angles 
divergent (Fig. 17), margin thin, faintly sul- 
cate (especially posteriorly); fulvus with 
faint viridis sheen: glabrous, nitidus with 
hne punctation throughout. Scutellum ful- 
vus, parallel-sided, anterior and posterior 
margins convex. Prosternum nearly straight 
in profile: fulvus to testaceus: glabrous with 
fine punctures medially, with fine erect se- 
tae and deep punctation laterally. Hypom- 
eron fulvus to viridis: glabrous, nitidus with 
punctures becoming obsolete posteriorly. 
Mesosternum fulvus to testaceus, fossa gla- 
brous and impunctate, lateral areas with 
dense setae and fine punctures. Mesepister- 
num fulvus to testaceus. Metasternum ful- 
vus to testaceus: glabrous with very fine 
punctation medially, decumbent setae and 
denser setae anterolaterally. Femora fulvus 
to testaceus. 

Elytron 10 mm (length/width ratio 3.0- 
3.1) subparallel on anterior half then nar- 
rowing to tip: fulvus to viridis: glabrous, 
nitidus, striae well marked medially, fainter 
laterally, strial punctures moderately deep: 
apex with apical spine and subapical tooth. 


Abdomen aurantiacus to testaceus; gla- 
brous with very fine piinctation medially, 
ileciimbent setae and denser setae antero- 
laterally; female not seen. Male with para- 
meres nanow medially, arcuate in profile, 
apical blades narrower than parameres at 
base, 21-23% length of parameres (Fig. 5). 

Material examined. — COLOMBIA: (1. 
BMNH); nr. Saludito Vaile. 6500 ft.. 20- 
VII- 1970, H. & A. Howden (1. CNCl). 

Diagnosis. — The small size and immac- 
ulate pale fulvus to viridis dorsal coloration 
separate S. virescens from all other Seiiii- 


Specific appreciation is extended to Boris 
C. Kondratieff and Paul A. Opler for en- 
couragement, suggestions, and support (both 
financial and otherwise) during the three 
years of this project. Acknowledgment is also 
made of the support provided by the Depart- 
ment of Bioagricultural Sciences and Pest 
Management at Colorado State University 
and the many lending individuals and insti- 
tutions that provided material. 

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Campbell, J. M. and J. D. Marshall. 1964. The ocular 
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103(3). 2()()3, pp. 664-673 


Terry L. Whitworth 

(TLW) Whitworth Pest Solutions Inc.. 2333 Inter Ave.. Puyallup. WA 98372. U.S.A. 

Ahstnui. — Adult and puparial characters of the bird nest blow fly. Protocalliphora 
sialia Shannon and Dobroscky (Diptera: Calliphoridae) are evaluated as to whether the 
eastern and western North American populations should be considered separate species. 
Based on an analysis of adult and puparial ratios and a variety of morphological features, 
the western North American populations are described as P. occidentalis, new species. 

Kc\ Words: 

bird. nest, blow fly. Proioculliphoni. sialiti. Diptera. Calliphoridae. North 
America, new species 

Larvae of species of Protocallipliom 
Hough (bird blow flies) are obligate blood- 
sucking parasites of altricial nestling birds. 
Twenty-eight species of Protocalliphora 
have been described from North America, 
including two which are Holarctic. while 13 
species have been described from the Pa- 
learctic Region. Sabrosky et al. (1989) re- 
described 1 1 North American species and 
described 15 new species. Whitworth 
(2002) recently described two new species 
of Protocalliphora in North America. 

Protocalliphora sialia Shannon and Dob- 
roscky infests a broad range of hosts and is 
one of the most commonly collected bird 
blow flies in North America. It is found fre- 
quently in the cavity nests of bluebirds, tree 
swallows, and European starlings, as well 
as occasionally in the open nests of Amer- 
ican robins, common ravens, and kingbirds. 
Bennett and Whitworth (1992) found P. 
sialia in 31% of 667 infested nests of 15 
bird species in eastern North America and 
in 7.3% of 964 infested nests of 18 bird 
species nests in the West. 

Protocalliphora sialia is one of several 
similar species where both sexes have the 

same color abdomens and white calypters, 
and males have digitate surstyli. This group 
includes P. shannoni Sabrosky et al., P. hi- 
riindo Shannon and Dobroscky, P. beniietti 
Whitworth, and P. riigosa Whitworth. A 
key to assist distinguishing these species 
was provided in Whitworth (2002). 

Male P. sialia can be distinguished from 
similar species by their broad frons, with a 
frons to head ratio averaging 0.08-0.10, 
and a polished triangular preocellar area. 
Protocalliphora sialia puparia have a long 
prothoracic fringe with an average diameter 
of 500-600(x (compared to 350-400(jl in 
most species), dorsal cuticle with moderate 
to pronounced ridges, and long hyperstig- 
matal spines, averaging 40-50|x. 

Sabrosky et al. (1989) discussed the pos- 
sibility that P. sialia was actually composed 
of two separate species. A comparison of 
adult male and female proportions revealed 
consistent, but overlapping differences be- 
tween speciinens from eastern and western 
North America divided roughly by the 
Great Plains. The most distinctive differ- 
ence between adults is a polished preocellar 
area in eastern females, while western fe- 



I. Distribution o<i Pintocalliplu. 

uilici (•) and l\ 

idcnicilis (*l in Niirtli AiiK-rica. 

males have a dull, microtomentose preocel- 
iar area. Only the eastern form of P. sialia 
puparia had been studied at that time and 
ultimately P. sialia was left as one species, 
ponding future research. The purpose of this 
study is to determine the status of this di- 
verse, widespread species. 

Methods and Materials 

Protocallipliora are rarely found in na- 
ture, except in bird nests. Most of the nests 
for the cuiTent study have been collected 

since 1992, via Internet contacts with bird- 
ers who collected them from bird nest box- 
es after nestlings fledged. Common bird 
names are from the "Checklist of North 
American Birds"', 7th edition, American 
Ornithologists Union (2000). Most material 
from Utah was collected from 1969-1972. 
Methods used to sort adults and puparia 
from nests and to identify Protocallipliora 
species were detailed in Whitworth (2002). 
Taxonomic terms were defined and an ex- 
planation of how to determine ratios in 


Table I . Mean ratios and ranges comparing western, midwestern. and eastern adults of Prouiciillipluini s'uilui 
to data troni Sabrosky el al. (1989). 

Frons Head 
Frons Ocellar Span 
Parafcicial Frons 
Piirafiicia/ Ocellar Span 
Piirafacial VIbrissal Interval 
Cheek Eye 

Frons 3rd Antennal Segment 
Parafacial 3rd Antennal 

0.08 (0.07-0.09) 
1.18 (0.8-1.5) 
1.5 (1,25-2) 
1.75 (1,5-2,33) 
0,92 (0,75-1,18) 
0,42 (0,33-0,49) 
1,23 (,89-1,5) 

1,8 (1,5-2.13) 

0.08 (0,07-0,09) 
1.10 (1-1,25) 
1,71 (1,5-1,87) 
1.88 (1,62-20,07) 
1,00 (0,84-1,18) 
0,40 (0,38-0,44) 

0.098 (0.09-0.1 I) 
1.27 (1.18-1.5) 
1.44 (1.16-1.75) 
1.83 (1., 5-2.11) 
1.02 (.79-1.13) 
0.43 (0.37-0.49) 
1.23 (1.09-1.5) 

1.78 (1.36-2) 

Data from Sabrosky et al., 1989. 

adults and pupaiia were discussed in Sa- 
brosky et al, (1989). See Whitworth (2002) 
for further discussion of measuring protho- 
lacic fringe and cuticular spines, 

A preliminary comparison of puparia re- 
vealed that specimens from the northern 
midwest (Minnesota and Wisconsin) have 
some characters intermediate between the 
eastern and western forms. Therefore, an 
initial analysis of adult and puparial char- 
acters was conducted on three groups: East- 
ern, western, and midwestern. Twenty-five 
adults of each sex were measured from all 
three areas and those measurements are 
compared in Table 1, along with data from 
Sabrosky et al, ( 1989). Twenty-five puparia 
from each area were also measured and are 
compared in Table 2, 


A total of 3,849 nests of 76 bird species 
were collected from 45 states in the contig- 
uous U.S., Alaska and Canada. Of these 
nests. 39% (1,501) from 49 bird species 
were infested with 17 species of Protocal- 
liphoni. Over half of the infested nests 
(992) from 22 bird species were infested 
with 'P. sialia." Nests with this species 
were collected from 33 states in the contig- 
uous U.S., Alaska, and Canada. 

Measurements of adult males, females, 
and puparia suggest a close relationship be- 
tween eastern and midwestern P. sicilia, 
with western P. sialla often significantly 

different (Tables 1, 2), Because the mid- 
western and eastern forms are so similar, 
the two are combined in further discussion 
under the eastern form. The eastern form 
extends northwest from Georgia (recorded 
by Sabrosky et al. 1989) through Minnesota 
and Canada to Alaska, while the western 
form is found from Arizona and New Mex- 
ico to Prince George, British Columbia 
(Fig. 1). Sabrosky et al. (1989) recorded 3 
male P. sialici from south central Mexico 
and, although we have not seen these spec- 
imens, it seems likely they are the western 
form, which extends south from New Mex- 
ico and Arizona. 

Although the stigmatal regions of puparia 
often appeared quite different (Fig. 2), dif- 
ferences in measurements of the eastern and 
western forms were relatively small and 
ranges overlapped (Table 1,2), Some mid- 
western puparia exhibited intermediate 
characters between the two forms, suggest- 
ing possible interbreeding, though the mid- 
western form was closer to the eastern 
form, indicating that two species were in- 
volved. This conclusion was further sup- 
ported by the results of molecular analysis 
which revealed two well-separated groups, 
one composed of the eastern and midwest- 
ern forms and the other of western forms 
(E. Baudry, personal communication). 
Baudry and I are conducting a study of the 
taxonomic relationship between species of 


Table I. Extended 

Sabroskv* Easi 

0.10 (0.09-0.12) 
1.4 (1.2-1.75) 
1.35 (1.2-1.67) 
1.88 (1.46-2.22) 
1.06 (.94-1.25) 
0.44 (0.41-0.47) 
1.26 (1-1.46) 

1.71 (1.33-2) 

0.09 (0.08-0.11) 
1.37 (1.13-1.67) 
1.43 (1.20-1.62) 
1.96 (1.69-2.18) 
1.04 (0.93-1.13) 
0.42 (0.38-0.45) 

0.28 (0.27-0.31) 
3.48 (3.08^.10) 
0.49 (0.43-0.56) 
1.68 (1.45-2.11) 
1.06 (0.68-1.27) 
0.48 (0.40-0.54) 

3.93 (3.4-4.5) 

1.94 (1.67-2.25) 

0.27 (0.25-0.30) 
3.58 (3.06-4.00) 
0,53 (0.48-0.60) 
1.91 (1.73-2.29) 
1.03 (0.90-1.20) 
0.43 (0.41-0.47) 

Protocalliphora, based on molecular anal- 
y.sis, and plan a future publication. 

Protocalliphora sialia was originally de- 
scribed as a new variety by Shannon and 
Dobroscky (1924), and no holotype was 
designated. Sabrosky et al. (1989) selected 
a lectotype and paralectotypes from Penn- 
sylvania, so the eastern and midwestern 
populations are P. sialia. while the western 
populations are hereby named, P. occiden- 
talis. I collected specimens of both species 
unless indicated otherwise. Most were tak- 
en from bird nests donated by birders who 
collected nests from nest boxes. 

Protocalliphora (P.) sialia 

Shannon and Dobroscky 

(Figs. 2a, b, 3a. c) 

Protocalliphora spleudida van sialia Shan- 
non and Dobroscky 1924: 251. 

Apauliiui sialia: Hall 1948. 

Protocalliphora sialia (eastern and western 
form): Sabrosky et al. 1989. 

Diagnosis. — See Tables 1 and 2 for com- 
parisons of ratios and measurements for 
adults and puparia. Male frons to head ratio 
averages 0.10 (0.09-0.12) vs. 0.08 (0.07- 
0.09) in P. occidentalis. Male polished 
preocellar area usually larger in P. sialia 
(Fig. 3a) vs. smaller in P. occidentalis (Fig. 
3b). Female frons to head ratio relationship 
is opposite of the males. For P. sialia it is 
smaller averaging 0.25 (0.22-0.28) vs. 0.28 
(0.27-0.31); preocellar area usually pol- 
ished (Fig. 3c) vs. dull colored and micro- 

tomentose (Fig. 3d). Stigmatal region of pu- 
paria with pronounced folds (Fig. 2a) vs. 
folds faint to absent (Fig. 2c); ventral band 
ratio averages 0.83 (0.71-0.89) vs. 0.72 
(0.57-0.85) (Table 2). 

Material examined. — CANADA. ON- 
TARIO: 2 nests Algonquin Park. Summer 
1955. G.F Bennett collector; 6 puparia. 
barn swallow nest #504; 6 puparia. nest 
#445. no host given. SASKATCHEWAN; 1 
puparium. Saskatoon, kestrel nest #2548. 
Summer 1995. 

banks. Creamer's Field. 2 tree swallow 
nests. July 1994, D.D. Roby collector; 12 
puparia mixed with 27 puparia of P. 
hraiieri (Hendel) and 34 puparia of P. heii- 
netti Whitworth. nest #41. 3 puparia mixed 
with 4 puparia of P. hennetli. and 10 pu- 
paria of P. hraiteri. nest #42. INDIANA: 55 
adults. 60 puparia. Union Co.. Brookville. 
eastern bluebird nest #4754. 5/24/01; 19 
puparia, Terre Haute, eastern bluebird nest 
#3770, 6/29/00. KENTUCKY: 15 puparia 
mixed with 10 puparia of P. deceptor Sa- 
brosky. Bennett, and Whitworth, Ashland, 
warbler species nest UJilill . 6/10/00. 
MARYLAND: 10 puparia mixed with 8 pu- 
paria of P. nietallica (Townsend), Cumber- 
land Co., Allegany, mockingbird nest 
#3733, 7/12/00. MASSACHUSETTS: 9 
puparia, Worchester Co., Worchester, east- 
ern bluebird nest #4129, 7/16/00. MICHI- 
GAN: 35 adults, 80 puparia, Hanover, east- 
ern bluebird nest #4397. 8/10/00; 15 adults 


Table I . Extended. 




Fnms Head 
Frons Ocellar Span 
Parafacial Frons 
Parafacidl Ocellar Span 
Paiafacicil Vibrissal Interval 
Cheek Eye 

Fnms 3rd Antennal Segment 
Parafacial 3rd Antennal 

0.26 (0.23-0.28) 
2.97 (2.83-3.18) 
0.56 (0.51-0.63) 
1.67 (1.58-1.83) 
1.09 (1.05-1.25) 
0.46 (0.42-0.5) 
3.12 (2.91-3.36) 

1.74 (1.64-LS3) 

0.25 (0.22-0.28) 
3.13 (2.82-3.27) 
0.56 (0.5-0.61) 
1.73 (1.5-1.91) 
1.12 (0.9-1.24) 
0.45 (0.39-0.49) 
3.18 (2.75-3.4) 

1.77 (1,64-1.91) 

0,265 (0.24-0.29) 
3.31 (3.06-3.73) 
0.555 (0.52-0.62) 
1 .84 (1 .65-20.0) 
1.12 (1.0-1.32) 
0.44 (0.42-0.46) 

and 1 10 puparia, same location and host as 
above, nest #4399. MINNESOTA: Four 
eastern bluebird nests from St. Paul; 150 
adults, 210 puparia, nest #3886, 7/15/00: 45 
adults. 50 puparia, nest #4241. 8/7/00; 10 
adults. 110 puparia. nest #4240. 8/6/00; 2 
puparia, nest #4239, 8/6/00: 40 adults, 50 
puparia, Sartell, eastern bluebird nest 
#5198. 7/21/01. NEW JERSEY: 7 adults. 6 
puparia, Newtonville, house wren nest 
#3978, 6/30/00; 8 puparia, Medford, eastern 
bluebird nest #2717, 8/3/99. NEW YORK: 
70 adults and 70 puparia, mixed with 5 P. 
hennetii, Cattaraugus Co.. Franklinville. 
tree swallow nest #4073, 7/25/00; 110 pu- 
paria. same location as above, kestrel nest 
#4076. 7/25/00; 5 puparia, Gloversville, 
eastern bluebird nest #4465, 5/25/00. 
OHIO; 75 adults, 110 puparia, Richland 
Co., eastern bluebird nest #2713, 7/15/99; 
19 puparia. Stark Co., Louisville, tree swal- 
low nest #4385, 7/24/00. PENNSYLVA- 
NIA: 75 adults. 85 puparia. Harrisburg. tree 
swallow nest #2638, 6/28/99. RHODE IS- 
LAND: 23 puparia. North Kingston, tree 
swallow nest #4896, 6/10/01. TENNES- 
SEE: 32 puparia. Knoxville. eastern blue- 
bird nest #5399. 8/4/01. VIRGINIA: 3 east- 
ern bluebird nests from Roanoke Co., Ro- 
anoke: 35 adults, 40 puparia, nest #3338. 5/ 
15/00; 50 puparia. nest #4146. 7/25/00; 160 
puparia, nest #4147, 7/25/00. VERMONT: 
9 adults, 55 puparia. Franklin, tree swallow 
nest #4350. 7/12/00. WEST VIRGINIA: 1 1 

adults. 45 puparia. Marion Co., Fairview, 
eastern bluebird nest #3386, 6/10/00; 15 
adults, 43 puparia, Lewisburg, tree swallow 
nest #3616. 6/15/00. WISCONSIN: 15 pu- 
paria. Dunn Co., Colfax, eastern bluebird 
nest #4942, 6/5/01; eight eastern bluebird 
nests, Dane Co.. Black Earth: 35 adults, 39 
puparia, nest #3295, 5/10/00; 3 adults. 15 
puparia, nest #3296, 5/10/00; 45 adults, 50 
puparia, nest #4279, 5/25/00; 1 5 adults, 28 
puparia, nest #4282, 8/7/00; 2 adults, 25 pu- 
paria, nest #4290, 6/29/00; 45 adults, 56 pu- 
paria, nest #4302, 5/25/00; 25 adults, 35 pu- 
paria, nest #4320, 5/25/00; 30 adults, 40 pu- 
paria, nest #4326, 6/6/00. 

Specimens examined. — 847 adults, 1988 

Distribution. — Alaska, Connecticut, Illi- 
nois, Indiana. Iowa, Kentucky. Maine, 
Maryland, Massachusetts, Michigan, Min- 
nesota, New Hampshire, New Jersey, New 
York, North Carolina, Ohio, Pennsylvania, 
Rhode Island, Tennessee, Virginia, Ver- 
mont, West Virginia, Wisconsin, Ontario 
and Saskatchewan, Canada (Fig. 1). These 
are the first records of this species from In- 
diana. Illinois. Kentucky, Minnesota, Rhode 
Island. Tennessee, and Wisconsin. Sabrosky 
et al. ( 1989) recorded this species from Del- 
aware, New Brunswick, Northwest Terri- 
tories, Ontario, South Dakota and Georgia. 

Hosts. — Eastern bluebird, great-crested 
flycatcher, kestrel, purple martin, northern 
mockingbird, house sparrow, barn swallow. 



° °° ? !fi l?i ? 

— O — (N VD O 

o i i i ^ 

S = 

(X c I I 2 5) 

ft > Q CQ < c^ 

tree swallow, tufted titmouse, house wren. 
The mockingbird and tufted titmouse hosts 
are new records for P. sialia. 

Protocalliphora (P.) occidentalis 
Whitworth, new species 

(Figs. 2c, d, 3b. d) 

Protocalliphora {P.) sialia: Sabrosky et al. 
1989 (in part). 

Diagnosis. — Proiocalliphora occidental- 
is adults are very similar to those of P. sia- 
lia, however male P. occidentalis have a 
narrower frons (0.08 vs. 0.10) (Table 1 ) and 
a smaller polished preocellar area (Fig. 3b 
vs. 3a); female P. occidentalis usually have 
a dull preocellar area (Fig. 3d) while P. sia- 
lia have a polished preocellar area (Fig. 3c). 
The stigmatal region of puparia in P. oc- 
cidentalis usually has folds faint or absent 
(Fig. 2c) vs. distinct folds in P. sialia (Fig. 
2a); ventral band ratios in P. occidentalis 
average 0.72 (0.57-0.85) vs. 0.83 (0.71- 
0.89) in P. sialia (Table 2). 

Male. — Mean ratios for P. occidentalis 
compared to P. sialia given in Table I . 
Frons usually nanower than P. sialia, preo- 
cellar area triangular, polished and 1-1.5 
times depth of median ocellus compared to 
2-2.5 times in P. sialia. 

Female. — Mean ratios for head compared 
in Table 1. Preocellar area usually dull mi- 
crotomentose. while in P. sialia usually 
large and polished (Figs. 3c. d). 

Puparium. — Mean ratios for puparial 
characters compared in Table 2. Compared 
to P. sialia, P. occidentalis with smaller 
ventral band ratio, ridges in hyperstigmatal, 
mesostigmatal, and hypostigmatal areas 
faint or absent vs. moderate to prominent, 
mesostigmatal and hypostigmatal area 
spines sparse, short, and often reduced to 
tubercles vs. spines dense to 15(x long. 

Types. — Holotype c?, allotype and 42 
paratypes (26 6 16 2) with matched pu- 
paria from Washington State, Kittitas Co.. 
Umtanum Ridge (road), 6/15/99. mountain 
bluebird nest #2612. Holotype and allotype 
in the National Museum of Natural History, 


Fig. 2. Puparium of Protocalliphoni sialia. a. Stignialal view. b. Dorsum. Pupariiim of P. occidenlali: 
Stigmatal view. d. Prothoracic fringe. 

Smithsonian Institution, Washington DC. 
Paratypes there and at Utah State Univer- 
sity, Logan. Utah, and Washington State 
University, Pullman, Washington. 

Additional paratypes.— CANADA, BRIT- 
ISH COLUMBIA: 7 adults, 40 puparia, Oso- 
yoos, western bluebird nest #3708, 7/1/00; 20 
adults, 1 1 5 puparia, Osoyoos, tree swallow 
nest #3714, 7/1/00; 3 tree swallow nests. 
Dykes area 30km south of Prince George, 
Summer 2001. Russ Dawson collector: 7 pu- 
paiia, with 9 P. riigosa. and 3 P. hennetti. 
nest #5793; 3 puparia with 28 P. rii(;osa. nest 
#5795; 10 puparia, nest #5807. 

western bluebird nests, Fredonia County, 
Mt. Trumbull: 10 adults. 110 puparia, nest 
#4015. 7/14/00; 35 adults, 1 15 puparia nest 
#4029, 6/10/00; 5 adults, 23 puparia, nest 
#4275, 8/7/00. CALIFORNIA: 1 1 puparia. 
El Dorado Co.. Mt. Akum Road, western 
bluebird, nest #2815, 7/10/00; 10 puparia. 
Rescue, violet-green swallow nest #3690, 7/ 

7/00; 110 puparia, Amador Co., Somerset, 
western bluebird nest #3849, 5/6/00. COL- 
ORADO: 130 adults, 175 puparia, Duran- 
go, ash-throated flycatcher nest #3620, 6/ 
25/00; 2 adults, 35 puparia. Grand Junction, 
ash-throated flycatcher nest #3686, 7/4/00; 
30 adults, 1 10 puparia, Durango, ash- 
throated flycatcher nest #4101, 7/15/00; 49 
adults, 1 1 5 puparia, Durango, ash-throated 
flycatcher nest #4108, 5/13/00; 85 puparia, 
Lyons, tree swallow nest #4142, 7/25/00. 
IDAHO: 2 puparia, Bonner Co., Athol, tree 
swallow nest #4039. 7/5/00; 36 adults and 
over 150 puparia, Kootenai Co., Hoodoo 
Valley, tree swallow nest #4267. 7/25/00. 
MONTANA: 48 adults, 75 puparia. Granite 
Co., Philipsburg, mountain bluebird nest 
#3497, 6/18/00; 31 adults, 65 puparia. 
Granite Co., Philipsburg, mountain bluebird 
nest #3501, 6/16/00. NEW MEXICO: 9 
adults, 92 puparia, Anoyo Seco, western 
bluebird nest #4392, 7/15/00; 9 adults, 12 



Fig. 3. Preocellar area of frons. a, Male Froroca/li/ihora 
Female P. sialici. d. Female P. occidentalis. 

b. Male Protocallipho 

pupaiia. AiToyo Seco. violet-green swallow 
nest #4395, 7/29/00. OREGON: 3 nests 
from Springfield; 15 adults and 32 puparia. 
western bluebird nest #3739, 6/16/00: 1 
adult, 9 puparia, tree swallow nest #3740, 
7/2/00: 12 adults, 27 puparia, tree swallow 
nest #5550, 7/14/01. UTAH: 1 adult, 35 pu- 
paria. Cache Co., Franklin Basin, tree swal- 
low nest #359, 7/18/70: 1 puparium mixed 
with 5 puparia of P. hinindo. Rich Co., 
Walton Canyon, cliff swallow nest #420, 7/ 
22/10: 5 puparia mixed with 45 puparia of 
P. parorum. Box Elder Co., Mantua, house 
wren nest #569, 7/8/69: 15 puparia. Cache 
Co., Logan Canyon, house wren nest 
#1050, 7/5/71: 12 puparia. Cache Co., Lo- 
gan, house sparrow nest #1302, 8/14/71; 2 
sialia puparia mixed with 16 P. rognesi 
Thompson and Pont [P. chrysorrhoea (Mei- 
gen)| puparia. Cache Co., Wellsville, bank 
swallow nest #1906, 6/26/74: 22 adults, 1 15 
puparia, Ogden. European starling nest 
#3764, 7/15/00. NEVADA: 6 puparia. Eu- 
reka, mountain bluebird nest #2926, 9/3/99: 
2 puparia. Eureka, mountain bluebird nest 
#2927. 9/10/99. WASHINGTON: 34 adults 
with matched puparia. Kittitas Co., Umtan- 

um Ridge, mountain bluebird nest #2611, 
6/15/99: 20 adults with matched puparia, 
Kittitas Co., Umtanum Ridge, bluebird spp. 
(mountain or western bluebird) nest #2609, 
6/15/99; 24 adult.s. 54 puparia, Kittitas Co., 
Umtanum Ridge, mountain bluebird nest 
#4226, 7/2/00; 22 adults, 35 puparia, Kit- 
titas Co., Umtanum Ridge, western bluebird 
nest #4228, 7/10/00. Additional paratypes 
to the same locations as the type series, also 
to the Spencer Entomological Museum, 
University of British Columbia, University 
of California, Berkeley, California Acade- 
my of Science, and my personal collection. 

Specimens examined. — 533 adults, 1878 

Distribution. — Arizona, California, Col- 
orado, Idaho, Montana, Nevada, New Mex- 
ico, Oregon, Utah, Washington, Wyoming, 
British Columbia, Canada and probably 
Mexico (Fig. I ). This is the first record of 
P. occidentalis from Nevada. 

Hosts. — Mountain and western bluebird, 
mountain chickadee, ash-throated flycatch- 
er, kestrel, pygmy nuthatch, red-breasted 
nuthatch, white-breasted nuthatch, house 
sparrow. European starling, barn swallow. 



tree swallow, violet-green swallow, oak tit- 
mouse, Bewick's wren, house wren. The 
red-breasted nuthatch and Bewick's wren 
hosts are new records for P. occidentalis. 
nests were contributed by Don Dahlsten, 
University of California, Berkeley. 

Etymology. — The name is taken from the 
Latin, occidental or "western" which re- 
flects the U.S. distribution of this species. 


Variation. — Individual adults of both 
species are difficult to distinguish from P. 
hennetti, P. hirundo. P. riigosa, and P. 
shannoni. See Whitworth (2002) for a key 
to these species. Lone females of P. occi- 
dentalis are virtually indistinguishable from 
female P. ruf^osa. Males of both species in 
good condition are usually distinctive, but 
dirty, teneral, or undersized specimens are 
a problem. The most distinctive feature of 
both species is the long prothoracic fringe 
in the puparia, which is usually 500|jl in di- 
ameter or more. Only P. paroniin Sabrosky 
et al. has a fringe that comes close, with 
some specimens to 450(x, and averaging 
425 |x. Occasionally specimens of P. sialia 
with undersized prothoracic fringe are 
found. The only Alaska specimens collect- 
ed were puparia from two tree swallow 
nests in Fairbanks, and all had a smaller 
than usual prothoracic fringe ranging from 
400-475[x. Based on the material available, 
1 have concluded they are P. sialia. but 
characters are somewhat intermediate and 
until more material is examined, including 
adults, this is a tentative identification. The 
fringe of occasional specimens of P. sialia 
in the eastern U.S. were similarly reduced, 
but this was unusual. Some specimens off. 
occidentalis from southern British Colum- 
bia, Washington, and Utah had an unusually 
long prothoracic fringe to 800|jl. Sabrosky 
et al. (1989) noted apparent normal body 
length for P. sialia (which included the 
eastern and western forms) was 8.5-9.5 
mm. However I reared many P. sialia that 
measured up to 10 mm, while P. occiden- 
talis usually were 8.5-9.0 mm (rarely to 9.5 

mm). This suggests that P. sialia tends to 
be longer, but size is not a good species 
character because nestlings often fledge be- 
fore all larvae mature, producing some un- 
dersized adults. 

The puparia of both species sometimes 
exhibit significant spine reduction in the 
mesostigmatal area. This is shared only 
with P. parorum among specimens with 
pronounced dorsal ridges. This character is 
common in P. occidentalis but also occa- 
sionally occurs in P. sialia. In P. occiden- 
talis the stigmatal area of puparia usually 
has no folds, P. sialia usually has pro- 
nounced to moderate folds. Scattered spec- 
imens of P. sialia from Ohio, Wisconsin, 
and Virginia also had reduced folds. 

Distribution. — The apparent wide sepa- 
ration of eastern and western populations 
by the Great Plains (Fig. 1 ) may prevent 
interbreeding and result in speciation. Spec- 
imens reared from nests in the extreme east- 
ern or western U.S. are usually very differ- 
ent, but midwestern puparia often have 
some intermediate characters that suggest 
interbreeding is occuning. Few nests have 
been examined from the Great Plains so the 
absence of specimens from that area may 
be due in part to the lack of collection ef- 
fort. However, P. sialia prefers the nests of 
birds in cavities and the lack of trees in the 
Great Plains could prevent populations 
from crossing the plains. With the expan- 
sion of farms and towns across the Great 
Plains in the past 150 years, the two pop- 
ulations of P. sialia recently may have had 
more opportunity for contact. Manmade 
structures often provide nest cavities and 
have shade trees or rows of trees for wind- 
breaks, which also provide nest cavities. 
The introduction of the European starling, 
over 100 years ago, could have helped the 
two species meet across the Great Plains 
since it is a favored host of both species 
and often nests in and around manmade 
structures (Bennett and Whitweirth 1992). 
More recently, many homeowners have be- 
gun installing nest boxes in places with few 
or no natural cavities to encourage blue- 


birds and other cavity-nesting birds, which 
may attract and disperse both species in ar- 
eas without trees. 

Some regions seemed to be totally de- 
void of both species, despite the fact that 
favored hosts like the eastern bluebird were 
abundant. The following nests and sites had 
neither species: sixty eastern bluebird nests 
in eastern Texas, 99 eastern bluebird nests 
in southeastern Oklahoma, 16 eastern blue- 
bird nests in southeastern Nebraska. 24 
eastern bluebird and purple martin nests in 
northern Missouri. 25 eastern bluebird nests 
in Arkansas, and 35 eastern bluebird and 
tree swallow nests in south central Iowa. 
Protocalliphora sialiu or P. ocvidentcdis 
has never been recorded from Arkansas, 
Missouri, Nebraska, Oklahoma, or Texas al- 
though P. sialia was found in northern 
Iowa. Protocalliphorci deceptor was found 
in each of the above areas and occasionally 
infested nests which would usually be in- 
fested by P. sialia. With the absence of 
their usual parasite, eastern bluebird nests 
in these areas were rarely infested. In areas 
where these parasites were common, infes- 
tation rates in favored host nests often ap- 
proached 100%. Neither species was found 
in North or South Dakota, or eastern Mon- 
tana, but few nests were examined from 
those areas, so further study is needed be- 
fore assuming they don't occur there. Sa- 
brosky et al. ( 1989) recorded P. sialia from 
an unknown location in South Dakota. Pro- 
tocallipliora sialia was never found in any 
of the southeastern states south of Tennes- 
see including South Carolina. Georgia. 
Florida, Alabama, Mississippi, and Louisi- 
ana, although Sabrosky et al. (1989) re- 
corded it from northern Georgia. In fact, 
Protocalliphora appears to be rare in much 
of the south, although I have recently ex- 
amined a series of 8 nests (wrens, chicka- 
dees, and warblers) from Smith County, 
Texas, that were all infested with P. decep- 
tor. several nests had over 100 puparia. 
This 100% infestation rate is in shaip con- 
trast to infestation rates of \%-5% by P. 

deceptor in over 200 nests examined from 
other areas of Texas. 


This study was made possible because of 
the cooperation of bird enthusiasts who sent 
me nests from all over the United States, 
Canada, and Alaska. I am especially in- 
debted to Tina Phillips of the Cornell Bird- 
house Network who publicized my request 
for bird nests. I also thank ornithologists 
Don Dahlsten, University of California. 
Berkeley, Russ Dawson, University of 
Northern British Columbia, Steve Matsuo- 
ka. United States Geology Survey, Anchor- 
age, Alaska, and Darrell Pogue, University 
of Texas at Tyler who sent many nests and 
specimens. Thanks to geneticist Emman- 
uelle Baudry. University Pierre et Marie 
Curie, Paris, France for her advice on ge- 
netic relationships. Thanks also to Rich 
Zack and Art Antonelli for their review 
comments. Also, thanks to Dawn Nelson of 
Seattle for the sketches and Patrick Craig 
of Monte Rio, California, for the slide pho- 
tos. Finally, thanks to my assistant Dana 
Hobbs, whose typing and computer skills 
were essential to the completion of this 

Literature Cited 

American Ornithologist's Union. 2000. Forty-second 
supplement to the American Ornithologist's Union 
Check-list of North American Ends. Auk 117: 

Bennett, G. F and T L. Whitworth. 1W2. Host, nest, 
and ecological relationships of species of Proto- 
calliplitiru (Diptera: Calliphoridac). Canadian 
Journal of Zoology 70: 51-61. 

Hall. David G. 1948. The blowflies on North America. 
Thomas Say Foundation (Entomological -Society 
of America), Lafayette. Ind.. 477 pp., 51 plates. 

Sabrosky. C. W.. G. F Bennett, and T L. Whitworth. 
1989. Bird blowflies (Protocalliphoia) in North 
America (Diptera: Calliphoridae). with notes on 
Palearctic species. Smithsonian Institution Press, 
Washington DC, 312 pp. 

Shannon. R. C. and I. D. Dobroscky. 1924. The North 
American bird parasites of the genus Proiocalli- 
plwra (Calliphoridae. Diptera). Journal of the 
Washington Academy of Sciences 14: 247-253. 

Whitworth, T. L. 2002. Two new species of North 
American Protocallipliora Hough (Diptera: Cal- 
liphoridae) from bird nests. Proceedings of the En- 
tomolomcal Society of Washington 105: 801-81 I. 

105(3). 2003. pp. 674-678 


Roger A. Burks and Richard A. Redak 

Entomology Department, University of California, River.side, CA 92521, U.S.A. 

Abstract. — Phera lacerta Fowler, combination restored, is transferred back to its orig- 
inal genus. Homalodisca liturata Ball, validity restored, is removed from synonymy with 
P. lacerta. Recent literature referring to P. lacerta as the smoke tree sharpshooter and a 
close relative of Homalodisca coagulata (Say), the glassy-winged sharpshooter, should 
actually be taken to refer to H. liturata. 

Key Words: sharpshooter, Cicadellidae, Homalodisca. leafhopper 

Phera lacerta Fowler, combination re- 
stored, was described in 1899 from Chil- 
pancingo, Guerrero, in southwestern Mexi- 
co (Fowler 1899b). It was transferred to 
Homalodisca Stal by Young ( 1968) as a se- 
nior synonym of Homalodisca liturata Ball 
validity restored, which had been de- 
scribed from Phoenix, AZ. Yuma. CA, and 
Comundu, Baja California Sur, Mexico, in 
1901. The synonymy was made without 
comment or explanation, although he had 
reviewed both P. lacerta (Young 1965) and 
H. liturata (Young 1958) previously in oth- 
er studies. That synonymy is reversed here, 
as P. lacerta and H. liturata represent dis- 
tinctly different entities belonging to differ- 
ent genera. 

Homalodisca liturata (as Homalodisca 
lacerta) has been the subject of much dis- 
cussion in recent literature (e.g., Nielsen 
1968, Powers 1973, Gill 1994, Sorensen 
and Gill 1996, Blua et al. 1999, Costa et al. 
2000, Bethke et al. 2001, Rakitov and Die- 
trich 200 1 ) as a close relative of the glassy- 
winged sharpshooter, Homalodisca coagu- 
lata (Say), a major pest of citrus, grapes, 
and ornamentals. Both H. liturata and H. 
coagulata are vectors of various strains of 

the phytopathogenic bacterium Xylella fas- 
tidiosa, which cause the diseases known as 
Pierce's disease in grapes, oleander leaf 
scorch, almond leaf scorch, phony peach 
disease, alfalfa dwarf, and citrus variegated 
chlorosis depending upon the crop species 
infected and bacterial strain involved (Blua 
et al. 1999). Homalodisca coagulata, a spe- 
cies from the eastern United States acciden- 
tally introduced to the western United 
States and northern Mexico, is the most im- 
portant vector of this disease in North 
American agricultural systems at this time, 
but H. liturata is also of interest as a closely 
related vector apparently native to south- 
western North America. 

Materials and Methods 

The type and other holdings of Homal- 
odisca liturata. in the National Museum of 
Natural History, Smithsonian Institution 
(USNM), were examined, as well as nu- 
merous specimens (over 100) of that spe- 
cies at the University of California. River- 
side, Entomology Research Museum 
(UCRC). Photographs (Figs. 1-2) of the 
lectotype (female) of Phera lacerta were 
obtained from The Natural History Muse- 


urn (M. Webb, BMNH), which led to the 
location of an additional male at UCRC that 
we assign to P. lacerta. collected from Bra- 
zil (BRAZIL. Sao Paulo: Piracicaba. ES- 
ALQ, 5-IV-1996. Screen, sweep wooden 
area, J. Pinto). All other photographs are of 
UCRC specimens, taken using Auto-Mon- 
tage software (version 4.00.0359 BETA, 
Synoptics. Ltd., UK 1997, 2001) through a 
3-CCD digital videocamera attached to a 


Pliera lacerta is distinguishable from 
both H. liturata and H. coagiilata by its 
longer, more robust proepimeron (Figs. 2, 
4) with a fiangelike ventral depressed mar- 
gin, the yellow longitudinal band along the 
side of its head and thoracic pleuron, and 
its pronotum with a longitudinal pale band 
flanked by darker stripes. The linear mark- 
ings on the vertex of P. lacerta (Figs. 1, 3) 
are also distinctive, but this character is 
widely variable (but not overlapping with 
P. lacerta) in H. liturata (Fig. 9), and we 
hesitate to treat it as a reliable character un- 
til more is known about its variation in P. 
lacerta. The yellow band along the thoracic 
pleuron (Figs. 2, 4) is an easily assessed 
character of the genus Phera Stal, but it is 
subject to variation in shape and color 
across several species, and may not be re- 
liable as a generic character. Young (1968) 
used the proepimeral shape and form to dis- 
tinguish Phera from Homalodisca. and this 
character is treated here to distinguish the 
two genera as it appears to be the most re- 
liable and unambiguous morphological 
character known to date. The central area 
of the proepimeron is longer than tall in 
Phera (Figs. 2, 4), but not in Homalodisca 
(Fig. 10). Phera species also have a large, 
depressed, flangelike ventral proepimeral 
margin, but a minute, shorter flangelike 
margin is present anteriorly (normally hid- 
den by the procoxa) in Homalodisca. Nev- 
ertheless, both proepimeral shape and the 
extent of its ventral depressed area appear 
to be reliable characters for distinguishing 

Phera from Homalodisca. except for Hom- 
alodisca insolita (Walker), which was re- 
garded by Young (1968) as an unusual spe- 
cies of dubious generic placement. On 
strength of proepimeral shape and form, P. 
lacerta certainly belongs in Phera. and H. 
liturata in Homalodisca. The differences 
between P. lacerta and H. liturata are very 
striking and we are unable to explain why 
Young (1968) synonymized the two. 

In addition to external differences, the 
genitalia of the male that we have identified 
as P. lacerta from Brazil (Figs. 5-8) differ 
substantially froin those of H. liturata. 
which were accurately figured by Young 
(1958). The forms of the pygofers. aedea- 
gus, connective, and styles of P. lacerta are 
very similar to those of Phera carhonaria 
(Melichar), as figured by Young (1968). In 
P. lacerta and P. carbouaria. the basal pro- 
cesses of the aedeagus are long and arching 
anteriad (Fig. 7). as opposed to those of H. 
liturata, which are always straight. Also, 
there is no projection from the atrium be- 
tween the bases of the processes in P. lac- 
erta. as there is in H. liturata. The pygofer 
of P. lacerta (Fig. 5) is also much longer 
than in H. liturata, greatly exceeding the 
length of the subgenital plates. Lastly, the 
subgenital plates of P. lacerta have lateral 
denticles not present in H. liturata, but 
which are found in several species of 

Ball ( 1901 ) was apparently already aware 
of Fowler's Central American and southern 
Mexican work at the time he described H. 
liturata, as he cited Fowler's discussion and 
figure of H. insolita in that publication. 
Phera lacerta was figured on the same plate 
as H. in.wlita (Fowler 1899a: plate XIV) 
and described only two months afterward 
(Fowler 1899b). Therefore, Ball was aware 
of at least the figure, and probably also the 
description, of P. lacerta at the time H. li- 
turata was described, and probably was 
well aware that the two species were dif- 
ferent. Melichar (1924) also correctly treat- 
ed H. liturata and P. lacerta as two distinct 
species in his monograph of the Cicadelli- 


Figs. 1-8. Phera lacerta. 1, Head and pronotum (dorsal view) of lectotype female. 2, Head and niesopleuron 
(lateral view) of lectotype female. 3. Head and pronotum (dorsal view) of male specimen. 4. Mesopleuron 
(lateral view) of male specimen. Arrow indicates ventral flange: pern = proepimeron. 5. Pygofer with subgenital 
plate. 6. Genitalia, posterior view. 7. Aedeagus, lateral view. 8. Connective and right style, dorsal view. 

nae, although he treated Homalodisca and 
Phera as synonymous genera. 

Literature records of Hoiiniloclisca lac- 
ertii from northern Mexico and the United 

States should be taken to refer to H. liltir- 
ata. which remains distinguishable from the 
closely related H. coagiilata using charac- 
ters given bv Younu ( 1958). Aside from its 


Figs. 9-10. Hoinalddisca lininiki. 9. Head and pniruUum (dursal \icw). 10. Mcsopleuroii (laloral \iew. proles 

Diiginal description, P. lacertci was correct- 
ly characterized only by Melichar (1924). 
and obviously remains very poorly known 
in terms of intraspecific variation. 

Knowledge of the true identity of H. li- 
uirata. the probable sister species of H. 
coagulata (Rakitov and Dietrich 2001), is 
important in planning biological control ef- 
forts of H. cocii>iilata and in studying the 
history of its distribution. Pliera lacerta is 
apparently a tropical entity, known from 
southern Mexico to southern Brazil. Con- 
versely, H. litiinita is known from south- 
western United States to central Mexico. 
The strategy of finding effective biological 
control agents in the home range of H. coa- 
^iilata by studying the distribution of its ap- 
parent sister species needs to be adjusted in 
light of this new information. 


We thank Mick Webb for providing pho- 
tographs and information about the lecto- 
type of Phera lacerta. Stuart McKamey for 
allowing examination of the lectotype of 
Hoinalodisca litiirata. and both for valuable 
and insightful comments and advice con- 
cerning this taxonomic problem. John Pinto 
and John Heraty allowed use of their mi- 
crophotography equipment for the images 
published here, and Matthew Buffington 
provided assistance in use of this equip- 
ment. Jutta Burger provided valuable ad- 
vice and technical assistance, and reviewed 
the manuscript, hi addition, Doug Yanega 
made available specimens for examination 

and photography, and provided helpful dis- 
cussion and comments. Finally, Jim Bethke, 
Jennifer Charles, Matthew J. Blua, and 
Thomas Prentice reviewed the manuscript 
and provided helpful comments and advice. 

Literature Cited 

Ball. E. D. 1901. A review of the Tettigonidae of North 
America north of Mexico. Proceedings of the 
Iowa Academy of Sciences 8: 35-75. 

Bethke. J. A.. K. A. Cainpbell, M. J. Blua, R. A. Re- 
dak, and D. A. Yanega. 2001. Range extension of 
Pseiieo punctatiis Fox and notes on predation of 
an introduced sharpshooter, Homalodisca coagii- 
lata (Say). Pan-Pacific 77( 1 ): 54- 

Blua. M. J.. R A. Phillips, and R. A. Redak. 1999. A 
new sharpshooter threatens both crops and orna- 
mentals. California Agriculture 53(2): 22-25. 

Costa. H. S.. M. J. Blua. J. A. Bethke. and R. A. Re- 
dak. 2000. Transmission of Xylellu fa^tidiosn to 
oleander by the glassywinged sharpshooter Hom- 
cdocliscii coai^idala. HortScience 35(7): 1265- 

Fowler. W. W. 1899a. Order Rhynchota. Suborder He- 
miptera-Homoptera (continued). Biologia Centra- 
li-Americana 2( 1, section 28): 217-224. 

. 1899b. Order Rhynchota. Suborder Hemip- 

tera-Homoptera (continued). Biologia Centrali- 
Americana 2(1. section 29): 225-232. 

Gill. R. J. 1994. New state records — glassy-winged 
sharpshooter California Plant Pest & Disease Re- 
port 13(1-2): 8-1 I. 

Melichar. L. 1924. Monographic der Cicadellinen I. 
Annates Musei Nationalis Hungarici 21; 195-243. 

Nielson. M. W. 1968. The leafhopper vectors of phy- 
topathogenic viruses (Homoptera. Cicadellidae) 
taxonomy, biology, and virus transmission. United 
States Department of Agriculture. Technical Bul- 
letin 1382. 

Powers. N. R. 1973. The biology and host plant rela- 



tions of Hoinalodisca lacerta (Fowler) in southern 
California. M.S. Thesis. California State Univer- 
sity. San Diego. 

Rakitov, R. and C. Dietrich. 2001. Evolution and his- 
torical ecology of the Proconiini sharpshooters. 
Proceedings of the Pierce's Disease Research 
Symposium, pp. 139-140. California Department 
of Food and Agriculture. 

Sorenson. J. T and R. J. Gill. 1996. A range extension 
oi Hoinalodisca cuagulata (Say) (Hemiptera; Cly- 
peorrhyncha: Cicadellidae) to southern California. 
Pan-PaciHc Entomologist 72(3): 160-161. 

Young, D. A. Jr 1958. A synopsis of the species of 
Hiiinaloilisca in the United States. Bulletin of the 
Brooklyn Entomological Society 53(1): 7-13. 

. 1965. Cicadelline types in the British Museum 

(Natural History) (Homoptera: Cicadellidae). Bul- 
letin of the British Museum (Natural History). En- 
tomology Series 17(4): 163-199. 

. 1968. Taxonomic study of the Cicadellinae 

(Homoptera, Cicadellidae). Part 1. Proconiini. 
United States National Museum Bulletin 261, 287 

105(3). 2003. pp. 679-684 




Donald G. Manley 

Department of Entomology. Clemson University, Pee Dee Research and Education Cen- 
ter. 2200 Pocket Road. Florence. SC 29506-9706. U.S.A. (e-mail: 

Abstract. — Dasymiitilla Jalisco, a new species of velvet ant (Hymenoptera: Mutillidae) 
from Mexico, is described from both sexes. The female is similar in appearance to Dci- 
syiniitilla caniiia (Smith), and the male is similar in appearance to Dasynnitilla intennixta 
Mickel. A host for D. Jalisco is the bee. Diadasia knabiaini Cockerell (Hymenoptera: 
Apidae). Spliaerophtlialma philinna Cameron and Mutilla (Ephuta) variicolor Andre are 
new synonyms of D. caniiia (Smith). 

Key Words: 

Dasynuitilla Jalisco, Mutillidae. new species. Dasytinililla caiiiiia, new syn- 

A recognized problem in working with 
velvet ants (Hymenoptera: Mutillidae) is 
that strong sexual dimorphism makes sex 
associations extremely difficult. Color pat- 
terns and often overall body size of the two 
sexes can be quite different (Brothers 
1989). Of more than 150 species of Dasy- 
iniitilla Ashmead. only about one-third are 
known from both sexes (Krombein 1979. 
Nonveiller 1990). 

Due to extreme sexual dimoiphism. sex 
associations can be made only by catching 
pairs //; copula (a very rare occurrence), 
through the use of caged females, or host 
data (Manley 1999). Although it is known 
that Dasymutilla species are parasitoids of 
other ground-nesting insects, primarily Hy- 
menoptera. Mickel (1928) noted that host 
relationships are known for only a few spe- 
cies. The situation has not changed much 
since that time. This may be further com- 
plicated by the fact that mutillid males and 
females may develop on different hosts 
(Matthews 1997). 

It is extremely rare to find evidence sup- 

porting both sex association and host rela- 
tionship within the same specimens, as is 
the case here. The specimens described here 
include seven females and three males, as 
well as two host bees. 

Materials and Methods 
Specimens used in this study were in- 
cluded with material sent to me for identi- 
fication by E. G. Riley from the Texas A & 
M University (TAMU) Insect Collection. 
All of these specimens were collected by 
William Godwin, and all were taken from 
cells of the same host in Jalisco. Mexico. 
The specimens have been compared with 
all known types of Dasymutilla, both males 
and females, and have been determined to 
be a new species. The two host bees were 
sent to Terry Griswold. Bee Biology and 
Systematics Laboratory. USDA. Utah State 
University, for identification. 

Dasymutilla Jalisco Manley, new species 

(Fig. lA) 

Diagnosis. — The female has the integu- 
ment entirely black, with contrasting yel- 



Fig. \. Dorsum of females of Dasymiitillu Jalisco and D. 
D. jaliscii. B, D. ciinina. [Scale lines = 0.8 mm.] 

showing patterns of black pubescence. A, 

lowish/golden and black pubescence. The 
thorax is as broad or broader than long, and 
is lacking a scutellar scale. The antennal 
scrobes are distinctly, although feebly, car- 
inate. A genal carina is lacking. The pygid- 
ium is rugose. The male has the integument 
of the head, thorax and legs entirely black. 
Integument of the abdomen is reddish ex- 
cept the terminal segment(s) black. Pubes- 
cence of the head and thorax is pale, con- 
trasted with black. The abdominal tergites 
are clothed with brilliant red pubescence, 
except the terminal segment(s) with black. 
Sternite II has a small, oval pit filled with 
pale setae. The antennal scrobes are dis- 

tinctly carinate. The last tergite lacks an 
apical fringe of setae. 

Description. — Female: Length, 13.5-16 
mm. Head black, vertex with dense, recum- 
bent, yellowish/golden pubescence, remain- 
der of head with paler pubescence; mandi- 
ble acute at tip, each with conspicuous in- 
ner tooth about one-quarter distance from 
tip: clypeus truncate on anterior margin, but 
concealed by long, pale setae; scape weakly 
carinate, the carina concealed by short, ap- 
pressed, pale pubescence; flagellomere I 
long, about as long as II and III united, re- 
maining flagellomeres subequal in length to 
second; antennal scrobe distinctly, although 


feebly, carinate; front and vertex coarsely 
punctate, but with dense yellowish/golden 
pubescence concealing sculpture; gena not 
as coarsely punctate as front and vertex, 
lacking genal carina, covered with ap- 
pressed pale pubescence; head almost as 
broad as thorax (3.3 mm x 3.5 mm). 

Thorax black; dorsum of thorax approx- 
imately 1.1 X as long as broad ( — 3.5 mm 
X 3.3 mm); scutellar scale absent; cephalic 
margin of pronotum evenly rounded, not 
emarginate medially; posterior face at sharp 
(—90°) angle to dorsal surface; dorsum of 
thorax with posterior V-shaped area of 
dense, appressed, yellowish/golden pubes- 
cence, concolorous with front and vertex, 
and anterior triangle of dense, appressed, 
black pubescence; posterior face of propo- 
deum, pleura, and sternum with appressed, 
pale yellow pubescence. 

Abdomen black; tergite I, including api- 
cal fringe, with appressed, pale yellow pu- 
bescence; tergite II with inverted V-shaped 
area of dense, appressed, black pubescence 
on anterior half; remainder of tergite II with 
dense, appressed, yellowish/golden pubes- 
cence, concolorous with front and vertex, 
except median half of apical fringe black; 
felt lines about 0.4 X length of segment, 
consisting of appressed, pale yellow setae; 
tergites III-IV largely with dense, ap- 
pressed, black pubescence, except lateral 
fringes of both tergites and apical fringe of 
tergite IV pale yellow; tergites V-VI en- 
tirely pale yellow pubescent; sternum, in- 
cluding apical fringes, pale yellow pubes- 
cent, except last sternite with long, black 
setae; sternite I with short, blunt carina, 
about 0.4X length of segment; pygidium 
coarsely rugose. 

Legs black, clothed with dense, pale yel- 
low setae. 

All specimens (7) similar in size and col- 

Male: Length, 13-14 mm. Head black, 
with long, appressed, white pubescence, 
dense on front and vertex, sparser else- 
where; mandible edentate, but each with 
strong dorsal carina ending about 0. 1 X 

length from tip; clypeus flat, bidentate me- 
dially on anterior margin; scape bicarinate 
beneath, lower carina stronger than the up- 
per, thinly clothed with white pubescence; 
flagellomeres subequal in length; antennal 
scrobe conspicuously carinate; eye normal, 
distance behind eye about equal to its great- 
est diameter; ocelli small, ocellocular dis- 
tance at least 3X greatest diameter of ocelli; 
head about as broad as thorax. 

Thorax black, pronotum and scutellum 
clothed with dense, appressed, white pubes- 
cence, mesonotum with dense, appressed, 
black pubescence, remainder of thorax with 
long, sparse, erect, white setae; cephalic 
margin of pronotum not emarginate medi- 
ally; tegula glabrous, impunctate, with 
sparse, appressed, black setae originating 

Abdomen reddish dorsally. fenuginous 
ventrally, except first and last segments 
black; tergites II-VI with dense, appressed, 
red pubescence; first segment with long, 
sparse, white setae; last segment with long, 
sparse, black setae; felt line long, about 
0.6 X length of second segment, comprised 
of appressed. white setae; sternite I with 
median, longitudinal carina, not produced 
on either end into tooth; sternite II approx- 
imately 3 mm long, with small, round pit 
densely filled with appressed white setae, 
situated about one-third distance from an- 
terior margin; sternites clothed with long, 
sparse, erect, white setae except as other- 
wise noted; pygidium finely rugose; pygid- 
ium lacking apical fringe of setae. 

Legs black, with white setae; apices of 
middle and hind femora rounded, neither 
squarely truncate nor sulcate. 

Apical half of forewing black, basal half 
infuscated but translucent; hindwing entire- 
ly translucent. 

Holotype.— 9. MEXICO, Jalisco, Mpio. 
LaHuerta, Chamela Biol. Sta.. VIl-26-1996, 
Wm. Godwin, collector, deposited in the 
Texas A & M University (TAMU) Insect 

Allotype. — c5 , same data and deposition 
as holotype. 



Paratypes. — 6 9.2 6, same data as ho- 
lotype, 2 9 and 1 6 in the author's collec- 
tion, remainder in TAMU collection. 

Host. — Diadasia knabiana Cockerell 
(Apidae), same data as holotype, deter- 
mined by Terry Griswold. Bee Biology and 
Systematics Laboratory. USDA. Utah State 
University, deposited in the TAMU collec- 

Etymology. — Named for the Mexican 
state in which the specimens were collect- 


Although no additional biological infor- 
mation is known, the fact that these 12 
specimens ( 10 mutillids and two host bees) 
were all collected from the same host cell 
complex provides substantial evidence of 
both the conspecificity of the mutillids and 
the host relationship. Diadasia Patton is a 
known host for some species of Dasyinulil- 
la (Krombein 1979). 

As is often the case with mutillids, the 
male and female differ in color pattern, al- 
though they are approximately the same 
size. The female goes to couplet 1 1 in 
Mickel's (1928, 1936) keys. It can be dis- 
tinguished easily from all Nearctic Dasy- 
mutilla strictly by the color pattern. 

The taxonomy of Neotropical Dasymiitil- 
la is clearly in a state of confusion. Dasy- 
mutilla Jalisco is very similar both visually 
and morphologically to D. canina (Smith) 
(Fig. IB). In D. Jalisco, the black pubescent 
pattern of the thorax is more nearly trian- 
gular in shape, the inverted V-shaped black 
pattern on tergum II extends only to about 
the midline of the segment, and the pale 
pubescence of the abdominal tergum is yel- 
lowish/golden, concolorous with the pale 
pubescence of the head and thorax (Fig. 
lA). In D. canina. the black pubescent pat- 
tern of the thorax extends anterolaterally to 
the humeral angles, thus giving more of a 
T-shaped appearance, the inverted V-shaped 
black pattern on tergum II extends past the 
midline, sometimes almost joining the 
black pubescence on the apical margin, and 

the pale pubescence of the abdominal ter- 
gum is almost white, much lighter than that 
of the head and thorax (Fig. IB). Consid- 
ering the within-species color variation in 
mutillids, it is possible that the females are 
color variants of the same species. How- 
ever, there is no evidence to support that at 
this time. 

Dasymutilla canina was described as 
Mntilla canina by Smith (1855) from the 
female only. The holotype is in The Natural 
History Museum (London). Saussure 
(1867) described both the male and female 
of M. sumichrasti. Mickel apparently could 
find neither in 1930-31 and designated a 
lectotype. The female lectotype designated 
by Mickel is in the Musee d"Histoire Na- 
turelle (Geneve). Mickel synonymized the 
two (1964) without explanation. I have ex- 
amined both female types and am in agree- 
ment with Mickel. No male type specimens 
were found by me in 1991. 

Dasyiniirilla philinna (Cameron) was de- 
scribed in 1895 from the female only. The 
holotype is in The Natural History Museum 
(London). Mntilla (Eplnaa) variicolor was 
described by Andre (1898) from the female 
only. The lectotype designated by Mickel 
in 1930-31 is in the Museum National 
d'Histoire Naturelle (Paris). Mickel synon- 
ymized the two (1964), again without ex- 
planation. I have examined both of these 
types and again am in agreement with 
Mickel. Having examined all four female 
types, it is apparent that these are all the 
same species. New synonymy follows at the 
end of this paper. 

I have examined male specimens identi- 
fied as D. sumichrasti. I have not, however, 
come across any male types. Males of D. 
sumichrasti that I have examined are clearly 
(morphologically) different from the male 
of D. Jalisco. Dasymutilla sumichrasti, as 
described by Saussure, appears to be a con- 
glomerate of at least four different species, 
those being D. personata (Cameron), D. 
terminata (Smith), D. deyrollesi Mickel, 
and D. tlialia (Cameron). It appears to be, 
in part, the male of D. canina. D. person- 


cita. and D. tenninala. Dasymiitilla tluilia is 
synonymous with D. inreriiii.xta Mickel, and 
is the male of D. sichelicma (Saussure) 
(Manley and Radke. 2002). Dasynnifilla 
dcyroUesi is clearly distinct. 

Males of all of the above species, as well 
as D. Jalisco, are very similar (almost iden- 
tical) in appearance. They can, however, be 
distinguished morphologically. Dasynuttilla 
personata can be distinguished easily from 
the others as it lacks a sternal pit filled with 
hairs. Dasynuitilla cleyrollesi can be distin- 
guished from the remaining species by the 
presence of an apical fringe of setae on the 
last tergite. Dcisyniiirilla tenniiuita and D. 
tluilia are the most difficult to distinguish. 
However. D. tenniiiata has the sides of the 
propodeum smooth and polished, while in 
D. tlialia the sides are punctate. 

The male of D. Jalisco keys to D. iiiter- 
mixta Mickel in Mickel's (1928. 1936) 
keys. It is distinguished easily from D. iii- 
lemiixta as well as the above species by its 
broad head, somewhat truncate on the pos- 
terior margin, by lacking an anterior emar- 
gination on the pronotum. by the anterior 
position of the pit on sternum II, and by the 
sculpture of the pygidium. Dasymutilla in- 
tennixta has the head nanowed posteriorly, 
has a median emargination on the prono- 
tum, has the sternal pit more median in po- 
sition, and has the pygidium glabrous and 

Although subsequent molecular and bi- 
ological studies may change the taxonomic 
relationships of some (or all) of the species 
mentioned, it is clear that the specimens de- 
scribed herein represent inale and female of 
the same species, and that the bee genus 
Diadasia serves as a host for this species. 

Dcisymiitilla caiiina (Smith), 
new combination 

Miitilla ccinina Smith 1855: 58. Holotype 9 
in The Natural History Museum (Lon- 

Miitilla sumichrasti Saussure 1867: 357. 
Lectotype 9 in the Musee d'Histoire Na- 
turelle (Geneve). 

Miitilla (Sphaeropthalma) sumichrasti: 
Blake 1871: 236. 9 (nee 6). 

Spluieroplithalma philiwia Cameron 1895: 
354. Holotype 9 in The Natural History 
Museum (London). New synonym. 

Spliaeroplithatnia sumichrasti: Cameron 
1895: 359, 9 (nee c5 ). 

Miitilla traiismarina: Dalla Torre 1897: 92. 
9 (nee 6). 

Miitilla (Ephiita) variicolor Andre 1898: 
56. Lectotype 9 in The Museum Nation- 
al d'Histoire Naturelle (Paris). New syn- 


I thank CI. Besuchet of the Musee 
d'Histoire Naturelle (Geneve). Tom Hud- 
leston. Suzanne Lewis, and Christine Taylor 
of The Natural History Museum (London), 
Janine Casevitz-Weulersse of the Museum 
National d'Histoire Naturelle (Paris). An- 
tonio Rolando of the University of Torino 
(Italy), and Phil Clausen of the University 
of Minnesota for access to their collections, 
including type specimens; and I thank Terry 
Griswold, Bee Biology and Systematics 
Laboratory. USDA. Utah State University, 
for identification of the host bees. This is 
technical contribution number 4773 of the 
South Carolina Agricultural Experiment 
Station. Clemson University. 

Literature Cited 

Andre. E. IS98. tilude sur les Mutillides du Museum 
de Paris. Annales de la Sociele Entomologique de 
France 67: l-7y. 

Blake. C. A. 1871. Synopsis of the Mutillidae of North 
America. Transactions of the American Entomo- 
logical Society 3: 217-265. 

Brothers. D. J. 1989. Alternative life-history styles of 
mutillid wasps (Insecta. Hymenoptera), pp. 279- 
291. In Bruton. M. N., ed. Alternative Life-His- 
tory Styles o\ Animals. Kluwer Academic Pub- 
lishers. Dordrecht. 

Cameron. R 1894-1896. Mutillidae. pp. 259-395. In 
E D. Godman and O. Salvin, eds. Biologia Cen- 
trali-Americana. Hymenoptera Vol. 2. 

Dalla Torre, K. W. V. 1897. Catalogus Hymenoplero- 
rum hucusque descriptorum systematicus et syn- 
onimicus. 8 (Fossores): 1-99. 

Krombein. K. V. 1979. Mutillidae. pp. 1276-1314. In 
Krombein. K. V.. R D. Hurd. Jr., D. R. Smith, and 



B. D. Burks, eds. Catalog of Hymenoptera in 
America North of Mexico, Vol. 2. Apocrita (Acu- 
leata). Smithsonian Institution Press. Washington. 
DC. pp. llW-2209. 

Manley. D. G. 1999. Synonymy of Dasynmulla iioc- 
tiirnii Mickel (Hymenoptera: Mutillidae). Pan-Pa- 
cific Entomologist 75: 18-22. 

Manley. D. G. and W. R. Radke. 2002. Synonymy of 
Dasyiniiulla sicheliaiui (Saussure) (Hymenoptera: 
Mutillidae). Pan-Pacific Entomologist 78:2-'<0- 

Mallhcvvs. R. W. 1997. Unusual sex allocation in a 
solitary parasitoid wasp. Splnieroprhalina pci>syl- 
vaiiicti (Hymenoptera: Mutillidae). Great Lakes 
Entomologist .^0: .51-34. 

Mickel. C. E. 1928. Biological and taxonomic inves- 
tigations on the Mutillid wasps. United States Na- 
tional Museum Bulletin 14.^: 1-3.51. 

. 1936. New species and records of nearctic 

Mutillid wasps of the genus DasymuiilUi. Annals 
of the Entonioloaical Society of America 29: 29- 

1964. Synonymical notes on neotropical Mu- 

tillidae (Hymenoptera). Proceedings of the Ento- 
mological Society of London 33: 163-171. 

Nonveiller, G. 1990. Catalog of the Mutillidae, Myr- 
mosidae and Bradynobaenidae of the Neotropical 
Region including Mexico. Hymcnoptcrorum Ca- 
lalogus. Pars 18: 1-150. 

dc Saussure, H. 1867. Mutillarum novaruni species al- 
iquot. Annales de la Societe Entomologique de 
France 7: 351-364. 

Smith. E 1855. Catalogue of Hymenopterous insects 
in the collection of the British Museum. Pt. Ill, 
Mutillidae and Pompilidae. London. 


105(3). 2003. pp. 685-697 


Robert D. Gordon and Scott McCleve 

(RDG) Systematic Entomology Laboratory. PSI, Agricultural Research Service, U.S. 
Department of Agriculture, c/o National Museum of Natural History. Smithsonian Insti- 
tution. Washington. DC 20.'>60-0168. U.S.A. Current address: Northern Plains Entomol- 
ogy. RO. Box 65. Willow City. ND 58384. U.S.A. (e-mail:; (SM) 
2210 13th Street. Douelas. AZ 85607. U.S.A. (e-mail: 

Abstract. — Five previously undescribed species of Eii/uirixia Brown. E. boliviaiia from 
Bolivia, E. campbelli from Costa Rica, E. isthmia and E. panamaensis from Panama, 
and E. mexicana from Mexico are described and integrated into the existing classihcation. 
A revised key to all known species is presented. 

Key Words: taxonomy, new species, ant nests 

Species of Eiiparixiu Brown (1927) are 
highly modified, very distinctive beetles 
that have evolved for an existence in ant 
nests. Woodruff and Cartwright (1967) 
summarized the known biological infor- 
mation consisting of three documented host 
associations and several suspected associa- 
tions. Enparixia formica Hinton, E. bruiieri 
Chapin, and E. moseri Woodruff and Cart- 
wright were collected in nests of leafcutting 
ants of the genus Atta Fabricius. Eiiparixia 
campbelli. n. sp.. was taken from an Atta 
detritus cavity in Guatemala (J. M. Camp- 
bell, label data). Most specimens are found 
at lights; rarely is a species excavated from 
a host nest. 

Discovery of a cryptic species closely 
similar to E. diiiicani Brown led to an in- 
vestigation of the Eiiparixia holdings in the 
National Museum of Natural History, 
Washington, DC. This comparison of spec- 
imens revealed an additional three new spe- 
cies described here and still another unde- 
scribed species from Bolivia was discov- 
ered by Paul Skelley. Florida State Collec- 
tion of Arthropods, Gainesville, FL. 

Acronyms for collections mentioned in 
the text are as follows: (ASM) Scott 
McCleve, Douglas, AZ; (CNC) Canadian 
National Collection, Ottawa; (FSCA) Flor- 
ida State Collection of Arthropods, Gaines- 
ville, FL; (MHN) Museo de Historia Nat- 
ural "Noel Kempff Mercado," Santa Cruz, 
Bolivia; and (USNM) National Museum of 
Natural History, Smithsonian Institution, 
Washington, DC. 


Genus Eiiparixia Brown 

Eiiparixia Brown 1927: 288; Woodruff and 
Carwright 1967: 6. Type species: Eiipa- 
rixia diincani Brown. By monotypy. 

Diagnosis. — Eupariini with inflexed 
clypeal margins; explanate pronotal mar- 
gins; basally constricted pronotal sides; ely- 
tral intervals usually cariniform; epipleurae 
covering episternum, epimeron and apices 
of middle coxae; middle coxae widely sep- 
arated; mesosternum separated from meta- 
sternum by transverse carina; middle and 
posterior tarsi long, slender. 

Remarks. — Woodruff and Cartwrisht 


(1967) reviewed the taxonomy of Eiiparix- 
ia. described one new species, and rede- 
scribed all known species. No subsequent 
studies have been published and Woodruff 
and Cartwright should be consulted for a 
summary of all previously known infor- 
mation concerning Eiiparixia. 

Examination of all presently known spe- 
cies revealed a strong gradient of moipho- 
logical adaptation from comparatively un- 
modified to highly modified pronota. Eii- 
parixia paiianuiensis, n. sp.. and E. costar- 
icensis Hinton possess almost rectangular 
pronota with only slight modifications of 
the lateral margin (Fig. 1). Euparixia for- 
mica Hinton and E. isthmia, n. sp., repre- 
sent an intermediate stage in pronotal mod- 
ification (Fig. 3). and the remaining species 
have highly modified pronota typified by an 
extreme constriction in the basal half (Figs. 
5, 7, 9. 11) (Woodruff and Carwright 1967: 
21, fig. 5). 

Revised Key to Species of Euparixia 

\. Anterior clypeal apex with small tooth on 

each side (Figs. 1. -^) 2 

- Anterior clypeal apex slightly angulate on 
each side (Figs. 5. 7. 9. 1 1) 3 

2( 1 ). Pronotum nearly rectangular, constricted in 
basal '/„. lateral margin gradually, nearly 
evenly rounded anterior to constriction (Fig. 
1 ) panamaensis, n. sp. 

- Pronotum not rectangular, lateral margin 
more or less sinuate (Fig. 3) . . isthmia. n. sp. 

3( 1 ). Elytral intervals moderately convex, not 
cariniform. except E. boliviana with inter- 
vals 7-9 weakly carinate (Fig. 10) 4 

- Elytral intervals distinctly cariniform (Figs. 

6. 8. 12) 5 

4(3). Pronotum narrowed only in basal V3 

costaricensis Hinton 

- Pronotum narrowed in basal Vi (Fig. 9) . . . 

boliviana. n. sp. 

5(3). Pronotum constricted only in basal W. pos- 
terior angles prominent and acute (Figs. 1. 
5. 7) 6 

- Pronotum constricted in basal % or more 
(Fig. 1 1 ), posterior angles obsolete or nearly 

so (Fig. II) 8 

6(5). Pronotum narrowed only in basal Vi. poste- 
rior angles prominent and acute (Figs. 5. 7) 

- Pronotum narowed at middle, posterior angles 
weakly defined, rounded fonnica Hinton 

7(6). Pronotal disc with deep punctures separated 
by thin, high ""walls" presenting a honey- 
comb appearance (Fig. 7) ... duncani Brown 

- Pronotal disc with shallow punctures sepa- 
rated by flat pronotal surface (Fig. 5) 

mexicana. n. sp. 

8(5). Basal W of pronotum with deep punctures 
narrowly separated by high ""walls"" pre- 
senting a honeycomb appearance (Fig. I 1); 
metasternum densely, coarsely punctured 

camphelli, n. sp. 

Basal V3 of pronotum with punctures not 
deep, separated by low ridges or convex 
pronotal surface; metasternum essentially 
impunctate 9 

9(8). Pronotum with lateral explanation nearly 
impunctate. lacking tubercles, pronotal sur- 
face slightly convex between punctures; 

(Louisiana and Cuba) 

nioseri Woodruff and Cartwight 

- Pronotum with lateral explanation weakly 
punctate, with some fine tubercles, pronotal 
surface with punctures separated by low 
ridges (Cuba) hnineri Chapin 

Euparixia panamaensis Gordon and 
McCleve, new species 

(Figs. 1, 2) 

Description. — Length 4.6 mm. width 2.1 
m. Elongate, convex, shiny, dark red. 

Head: Clypeus widely emarginate be- 
tween toothed angles, sides arcuate to fee- 
ble genal sulcus. Margin not reflected dor- 
sally at center, ventrally inflexed at center; 
genal angle obtuse. Surface with moderate, 
dense, nearly contiguous punctures across 
occiput: punctures smaller, sparser between 
eyes, becoming elongated anteriorly, pro- 
gressively smaller, changing to minute 
punctures near anterior margin. Frontal su- 
ture not evident. Eye not visible in repose. 
Lateral margin of gena with narrow sulcus 
from margin to deep pore at anterior edge 
of eye, sulcus continued above eye, with 
short, sharp carina at margin adjacent to 

Pronotum: Broader than long (1.8 mm 
X 1.2 mm), moderately convex. Basal mar- 
gin with single row of large, contiguous 
punctures; basal Vi of disc with few large, 
confused punctures anterior to basal row, 
then scattered, large punctures 1-2 diame- 
ters apart with tiny puncture or granule in 


Figs. 1—4. Pronota and elytra. I. Eupi 
pronotuni. 4. E. isthinhi. elytra. 

rixia piinamaeusis, prnnotutn. 2. E. panamaensis. elytra. ?>. E. islhinia. 

bottom of each, large punctures becoming 
smaller anteriorly. Scattered among large 
punctures are tiny punctures remaining un- 
changed to anterior margin where they be- 
come indistinguishable from reduced larger 
punctures. Sides explanate, weakly punc- 
tured, dull, with scattered small setae. Base 
with weak longitudinal groove in basal Vi. 

larger punctures with small punctures in 
bottom often contiguous in and near 
groove. Oblique depressions moderately 
well defined, about 10 punctures long and 
2-3 punctures wide. Anterior angles broad- 
ly rounded, sides slightly diverging, almost 
parallel for -A of length of explanation to 
almost imperceptible angle, then slightly 


converging for remaining y, of explanation: 
posterior angles obtuse, angles wider than 
elytra across base, basal constriction very 
short, only '/j of pronotum (Fig. 1). 

Elytron: Elongate-oval (right elyron 
missing), widest at apical Vs; intervals flat 
with scattered minute granules, each with 
an extremely minute seta, granules some- 
limes in single row, sometimes in weak, 
widely spaced, offset double rows. Striae 
with punctures moderately deep, small, 
sides of striae uneven and with indentation 
behind many punctures on outer side of 
stria, floor of stria with tiny transverse lip 
behind each puncture associated with each 
indentation of outer side of stria; mesal side 
of each stria nearly straight (Fig. 2). Scu- 
tellum with shallow, irregular, rugose, 
punctate depression at base. 

Sternum: Prosternal process broadly ar- 
cuate at base, margins raised, shiny, thick 
at base, carinate laterally, disc depressed in 
a "V" with apex forward. Middle coxae 
separated by 1 Vi length of middle trochan- 
ter, middle depression clearly defined, de- 
pressed anteriorly, with 2 naiTow isosceles 
triangular raised areas, 1 on each side, with 
base of each triangle near anterior margin 
of depression and apex pointing posteriorly; 
microsculpture of depression not evident at 
5()X. Meso- and metasterna separated by 
transverse carina with projections extending 
onto metasternum only, merging with shiny 
disc of metasternum. Center of meta- 
sternum with deep, longitudinal sulcus. Sul- 
cus behind middle coxae deep anteriorly, 
moderately punctate, shallow, with large, 
confused punctures posteriorly. Disc of 
metasternum shiny with minute punctures, 
sides with few large punctures each with 
minute punctures in bottom. Depression be- 
hind middle coxa deep, shaiply defined, 
shiny, rugulose, punctate in bottom. Side of 
metasternum laterally and anteriorly with 
small to moderate punctures. 

Abdomen: First sternum with closely 
spaced, longitudinal carinae forming quad- 
rate cells on posterior margin (fluting). Ster- 
na 2-5 each depressed on anterior margin. 

depressions with closely spaced, long cari- 
nae forming quadrate cells, cells on second 
sternum about !4 length of sternum, V^ 
length of sternum on third sternum, more 
than Vi length of sternum on fourth and fifth 
sterna at center. Posterior margin of sterna 
2-3 with small Vi punctures, larger laterally, 
giving scalloped effect to margins. Poste- 
rior margin of fourth sternum with row of 
minute, light colored setae. Disc of sterna 
2-3 shiny, with scattered, minute, setose 
punctures, punctures larger laterally. Ster- 
num 4 with disc apparently impunctate at 
center, punctures evident laterally. Disc of 
fifth sternum shiny, with small setose punc- 
tures. Pygidum with disc somewhat shiny, 
with short, longitudinal carina at basal cen- 
ter and numerous, scattered, small tuberlces 
on each side. Basal Vi of pygidium with 
strong longitudinal sulcus at center. 

Leiis: Anterior tibia tridentate, middle 
tooth longest, apical tooth bent outward at 
obtuse, nearly right, angle. Apical spur 
elongate, nanow, slightly curved. Anterior 
trochanter prominent, shiny dorsally, dull 
ventrally, inner margin with few setae, 
crenulations vaguely evident. One long seta 
on femur near trochanter. Anterior femoral 
groove and about Vi of dorsal surface with 
appressed golden setae; ventral surface 
completely covered with moderate punc- 
tures separated by their own diameter. Mid- 
dle femoral marginal groove complete from 
apex to near trochanter; posterior femoral 
marginal groove evident from apex to less 
than Vi distance to trochanter. Middle and 
posterior femora shiny, with minute punc- 
tures and associated setae, punctures sepa- 
rated by 1-3 a diameter. Posterior coxa be- 
tween trochanters shiny with few moderate 
to large punctures on mesal Vi. 

Type material. — Holotype 6: Panama, 
Canal Zone, Gamboa, 24-V-53 (USNM). 

Distribution. — Known only from the 
type locality. 

Material examined. — The holotype male. 

Etymology. — The species is named for 
the country where the type specimen was 


Remarks. — The nearly rectangular out- 
line of the pronotum with expianate mar- 
gins continued almost to the base will serve 
to separate this species from those known. 
Other unique features include the large 
punctures with a tiny puncture in the bot- 
tom of each on the pronotum and meta- 
sternum. flat elytral carinae with scattered, 
minute granulations, scutellum with only a 
shallow, rugose depression at the base, and 
the unusual mesosternum with two trian- 
gular raised areas. 

The holotype bears a handwritten label 
identical to that of two paratypes of E. isth- 
luki except dates VI instead of V. Both spe- 
cies were likely collected from the same 
light trap, and this demonstrates for the first 
time that two species of EitparLxia are sym- 
patric. The holotype is missing the right el- 
ytron, right hind leg, and all, or parts of all, 

Eiiparixia isthmia Gordon and McCleve, 
new species 

(Figs. 3, 4) 

Description. — Length 4.4 mm, width 2.0 
mm Elongate, convex, dull, dark brown. 

Head: Clypeus broadly arcuate between 
toothed angles, sides arcuate to posterior 
margin without genal notch, anterior mar- 
gin not reflected dorsally at center, ventrally 
feebly recurved at center; gena prominent, 
right angled. Surface densely, moderately 
punctate across base, punctures separated 
by less than a diameter, becoming much 
smaller and sparser anteriorly and laterally, 
changing to minute rounded denticles. Pos- 
terior margin of gena with narrow, shiny, 
impunctate sulcus extending almost to an- 
terior pronotal margin, with small, deep 
pore above eye, and with short, sharp carina 
adjacent to eye. Frontal suture lacking. 

Pronotum: Broader than long (2.0 mm 
X 1.2 mm), moderately convex (Fig. 3). 
Basal margin with single row of large, 
deep, contiguous punctures, punctures an- 
terior to row confusedly placed and of var- 
ious sizes, some contiguous or overlapping, 
others separated by a diameter, most smaller 

punctures bearing a short, light colored seta 
near anterior margin. Pronotum in basal Vi 
with deep, longitudinal, punctate groove 2 
large punctures wide, and with deep, broad, 
oblique depression on each side of disc, de- 
pressions irregularly punctate with large 
punctures. Surface anterior to oblique de- 
pressions finely, regularly punctate, punc- 
tures becoming minute near anterior mar- 
gin. Expianate sides dull, opaque, impunc- 
tate or with weak punctures or denticles and 
scattered short setae. Anterior angles shal- 
lowly, broadly rounded, forming nearly 
right angle with lateral margin. Expianate 
margins with angle at apical V3. constricted 
posterior to angle, then 2nd angle present 
at median Vi before basal constriction. Basal 
constriction short, basal margin about as 
wide as elytra at base (Fig. 3). 

Elytron: Elongate oval, widest at apical 
1/3, intervals on disc flat or only weakly car- 
inate (Fig. 4), with single row of minute 
granules separated by about a diameter, 
each granule bearing a minute seta; inter- 
vals more carinate laterally, especially 9th 
interval, and apically. Each stria with row 
of indistinctly sculptured, elongate, shallow 
punctures separated by less than length of 
puncture. Scutellum narrow, shiny, without 
basal depression. 

Sternum: Prosternal process broadly 
rounded at base, margins raised, slightly 
shiny, disc raised, irregularly, rugosely 
sculptured, with alutaceous sulcus on each 
side between disc and lateral margins. Me- 
sosternum with coxae separated basally by 
\V2 length of middle trochanter; median de- 
pressed area clearly margined, depression 
deep, anterior % with sharp, central, longi- 
tudinal carina flanked on each side by deep 
groove, then series of 5-6 more or less par- 
allel, anastamozing carinae; carinae and 
grooves between alutaceous, basal portion 
of mesosternum slightly shiny, with micro- 
reticulation barely evident at 50X. Meso- 
and metasterna separated by indistinct, 
transverse carina, longitudinal extension of 
carina onto meso- and metasterna lacking. 
Metasternum with center of disc depressed. 


with deep, coarse sulcus appearing to have 
tiny, longitudinal carina at bottom for near- 
ly entire length; carina behind middle coxae 
strong, adjacent sulcus indistinct, weekly 
punctate; remainder of disc moderately 
shiny, minutely, moderately punctate, each 
puncture with minute, broad, whitish seta; 
depressed area in front of posterior coxa 
dull, deep; sides feebly shining, moderately, 
indistinctly punctate. 

Abdomen: Basal sternum depressed 
along posterior margin, depression with 
longitudinal carinae forming rectangular 
cells. Second sternum with single row of 
short, weak, parallel, longitudinal carinae in 
anterior !/„,, carinae not forming quadrate 
punctures; punctures on posterior margin 
enlarged, with posterior V2 of each effaced, 
forming scalloped margin; third sternum 
with single row of short, shaiply distinct 
carinae about V,, length of sternum, not 
forming quadrate cells along anterior mar- 
gin, posterior margin with single row of en- 
larged V2 punctures forming scalloped mar- 
gin; fourth sternum with anterior V2 de- 
pressed, depression with longitudinal cari- 
nae. forming quadrate cells, posterior 
margin weakly crenulate; fifth sternum with 
anterior V2 depressed, depression with close- 
ly spaced, longitudinal carinae forming 
cells about 4 times longer than wide; pos- 
terior V2 of sternum shiny; disc of sterna 2- 
5 weakly punctate, punctures small, mod- 
erately spaced, with tiny setae more evident 
than punctures. Pygidum with narrow, dull, 
apical margin; disc depressed, dull, with 
fine median carina at basal center and sev- 
eral small tubercles laterally. Basal Vi of py- 
gidium alutaceous. with weak, longitudinal 

Legs: Anterior tibia tridentate, middle 
tooth longer than others, apical tooth ob- 
tuse, bent outward at less than right angle; 
apical spur slender, curved. Anterior tro- 
chanter large, inner margin with 3—4 setae 
in apparent punctures causing coarse, weak 
crenulation of margin, shiny and impunc- 
tate dorsally, ventrally dull, rugose, punc- 
tate. Anterior femur with single seta on 

each side near trochanter. Anterior femoral 
groove and about V3 of dorsal surface of fe- 
mur covered with fine, sparse setae; poste- 
rior surface dull, with moderate punctures 
separated by about a diameter or less, some 
punctures indistinct. Middle femur with 
marginal groove strongly impressed from 
trochanter to apex. Posterior femur with 
marginal groove extending length of femur, 
except weak or absent near trochanter. Pos- 
terior coxa between trochanters alutaceous 
on mesal V2, shiny on lateral V2, without ap- 
parent punctures or setae. 

Variation. — Length of male paratype 4.9 
mm. and female paratype 4.5 mm. Both 
paratypes have either no setae or only 1 seta 
on anterior femur near trochanter. The fe- 
male is lacking both elytra, one anterior leg, 
and all. or parts of all, tarsi except for one 
complete anterior tarsus. 

Type material. — Holotype 6: Panama, 
Canal Zone, Ft. Click, Bldg. 708, 17 May 
"65, light (USNM). Paratypes, 2; I cJ, 1 9, 
Panama. Canal Zone. Gamboa, 24-VI-.'r3. 

Distribution. — Known only from the Ca- 
nal Zone, Panama. 

Material examined. — The three type 

Etymology. — The specific epithet is from 
the Latin isthmus, meaning neck or narrow 
passage, and refers to the Isthmus of Pan- 

Remarks. — This species has the inter- 
mediate type of pronotum (see generic dis- 
cussion), a character shared only with E. 
formica Hinton. The mesosternal sculpture 
appears to be unique in the genus. The lon- 
gitiudinal carinae are quite striking, some- 
what like a portion of a woodcut as used in 
printing illustrations with the carinae and 
grooves seemingly carved into the surface. 
The lack of an anterior scutellar depression 
is also unusual. 

Euparixia mexicana Gordon and 
McCleve, new species 

(Figs. 5. 6) 
Description. — Length 5.4 mm. width 2.4 
mm Elongate, convex, feebly shiny, dark 
reddish brown. 


Figs. 5-8. Pronota and elytra. 5, Euparixia iiiexicaiu 
pronotiim. 8, E. cliiiuani. elytra. 

proiioluin. 6, E. 

elytra. 7. E. Jul 

Head: Clypeus widely truncate between 
weak angles, sides feebly arcuate to very 
slight genal notch, margin dorsally, feebly 
recurved at center and ventrally inflcxed at 
center; genae prominent, right angled. Sur- 
face finely and densely punctate between 
eyes, punctures separated by a diameter or 
less, finer at sides and anteriorly, changing 
to minute granules in anterior VS. granules 

continuing to anterior margin. Frontal su- 
ture not visible. Eyes not visible when head 
in repose. Lateral margin of gena with a 
weak, punctate sulcus beginning near an- 
terior margin and deepened gradually near 
eye into a deep pore, and with short carina 
at margin of gena adjacent to eye. 

Proiiotnin: Broader than long (2.1 mm 
X 1.6 mm, very convex (Fig. 5). Basal mat- 


gin with single distinct row of contiguous 
large punctures. Basal V2 of disc with large 
punctures a diameter or less apart, becom- 
ing abruptly smaller and sparser anteriorly; 
much smaller, shallow, punctures present 
between large punctures of basal Vi and 
continuing anteriorly, becoming denser as 
larger punctures disappear. Explanate sides 
impunctate or with indistinct small punc- 
tures. Oblique depressions on pronotum 
very weak, filled with about 20 large punc- 
tures in about 2 rows. Anterior angles 
broadly rounded. Posterior angles distinct, 
right angled. Explanate margins with an an- 
gle at anterior %, then narrowed slightly to 
2nd prominent angle at median Vs, then nar- 
rowed sharply to short basal constriction. 
Pronotum at base much narrower than ely- 
tra at base. 

Elytron: Elongate oval, widest at apical 
Vr. intervals convex, weakly carinate (Fig. 
6), each interval bears row of minute gran- 
ules barely visible at SOX, separated by 
their own diameter. All 10 striae with series 
of deep punctures; superimposed above 
deep punctures is a chain of larger and shal- 
lower punctures (the stria itself) formed by 
toothlike points projecting from each side 
of each raised interval; points extend to- 
ward each other from each adjacent interval 
toward or partially over each deeper punc- 
ture, giving complex appearance. Scutellum 
naiTow, with sharply defined depression at 

Sternum: Prosternal process broadly 
rounded, almost truncate posteriorly, disc 
strongly margined at base, less so laterally. 
Mesosternum with coxae separated by 1 V2 
length of middle trochanter: median depres- 
sion clearly margined, depression in basal 
% somewhat shiny, microsculpture evident 
at SOX, anterior Vs of depression slightly 
raised nearly to level of margin and arcu- 
ately delimited behind on a line beginning 
opposite anterior Vi of middle coxa, curving 
anteriorly toward prosternal process and 
continued to middle of opposite coxa: 
raised anterior Vi of depression irregularly 
shiny, rugose, punctate. Meso- and metata- 

stema divided by transverse carina with fine 
carina projecting both anteriorly into me- 
sosternal depression and posteriorly onto 
metasternum. Latter carina gives way to 
long, coarse, deep median sulcus. Carina 
bordering posterior portion of middle coxa 
large, separated from disc of metasternum 
by deep, variably punctate sulcus, remain- 
der of disc with fine to moderate punctures 
except for shiny impunctate area between 
posterior end of median sulcus and hind 
coxae; depressed area anterior to hind coxa 
deep, sharply bordered, confusedly rugu- 
lose, narrowly smooth and shiny adjacent 
to posterior coxa; anterior and lateral mar- 
gins between middle coxa and epipleuron 
with punctures larger than on disc, surface 
shiny between punctures. 

Abdomen: First sternum with in^egularly 
spaced, longitudinal carinae on posterior 
margin forming quadrate cells open behind. 
Sterna 2-S each depressed on anterior mar- 
gin, depression with irregularly spaced, lon- 
gitudinal carinae, depression longer on each 
successive sternum. Posterior % of sterna 2- 
3 with coarse punctures, larger laterally, 
followed by single row of larger punctures 
open behind, giving a scalloped effect to 
posterior margin. Sternum 4 with smaller 
punctures and with series of extremely mi- 
nute, light colored setae at posterior margin. 
Fifth sternum impunctate at center of pos- 
terior margin. Pygidium with raised, shiny, 
apical margin thickened at middle; disc de- 
pressed, shiny, with short median carina at 
basal center and several small tubercles on 
each side near basal margin; Basal V2 of py- 
gidium above ridge strongly alutaceous, 
with a strong, longitudinal sulcus. 

Legs: Anterior tibia tridentate; apical 
tooth bent outward at nearly right angle. 
Apical spur elongate, narrow, slightly 
curved. Anterior trochanter prominent, 
shiny dorsally, rugose, punctate ventrally, 
inner margin with about 12 extremely short, 
light colored setae giving margin minute 
crenulations. Both anterior femora with 3 
setae near trochanter. Anterior femoral 
groove and about % of dorsal surface cov- 



ered with short golden setae. Middle fem- 
oral marginal groove complete from apex 
to apex of trochanter. Posterior femoral 
marginal groove extending from apex to 
apex of trochanter. Posterior femoral mar- 
ginal groove extended from apex less than 
Vi distance to trochanter, feebly impressed. 
Middle and posterior femora with moderate 
punctures, each with short, conspicuous 
seta. Posterior coxa between trochanters 
punctate, punctures moderate, distinct, mi- 
nutely setose. 

Variation. — Length 4.8 to 5.7 mm. The 
single female from Tomatlan (4.9 mm) is 
essentially the same size as the male with 
same data (4.8 mm). Some specimens show 
a vague, short, longitudinal groove at the 
pronotal base. Some specimens bear minute 
setae on the pronotum. elytral intervals, and 
metasternum. One Ajijic specimen has the 
posterior angle of the pronotal explanation 
more developed than any other; the same 
specimen has the basal constriction of the 
pronotum longer than the others, although 
it also varies in length among the others. 
The number of setae near the trochanter 
varies considerably among specimens as 
follows, with the first number of each pair 
representing the left femur: 2-3, 5-2. 3-2. 
3-4, 3-2, 3-3. 

Type material. — Holotype 6: Mexico. 
Sonora. 17 km sw Moctezuma, vii-21, 22- 
80, 944 m, at light, S. McCleve, P Jump 
(FSCA). Paratypes, 6. 1 6. Mexico. 
JAL(Jalisco), 3 mi. N. Barra de Navidad, 
Bahia de Coastecomate, 17 August, 1964, 
WLNutting, It. trap, thorn forest clearing: 2 
6, Mexico, JAL., Ajijic, 21 June. 1964, 16 
July, 1964, 5140'. WLNutting, UV. It. trap: 
1 6.1 9, JA( Jalisco), hwy 200, 15 mi S 
Tomatlan, vii- 11-84, UVL, S. McCleve, R 
Jump: 1 6. Mexico, Sin(Sinaloa), Veno- 
dio(Venadillo?), C.17, Kusche "18. donor 
BP Clark (ASM, USNM). 

Distribution. — Mexico (northern Sonora, 
Sinaloa, and Jalisco). 

Material examined. — The seven type 

Etymology. — The species name refers to 

the country where all type specimens were 

Remarks. — This species is most similar 
to E. duncani Brown. The pronotal outlines 
are very similar, but E. nw.xicaiia has nar- 
rower lateral explanations (Figs. 5, 7). The 
elytral intervals of E. mexicana are less car- 
inate, bearing only a row of separate, mi- 
nute granules, whereas in E. duncani the 
granules are crowded together onto the ca- 
rinae (Figs. 6, 8). Both meso- and meta- 
sterna are sculpted differently in each spe- 
cies. Because of the similarity in appear- 
ance, this species has been misidentified as 
E. duncani Brown. The two Ajijic para- 
types and the single Venodio, Jalisco, para- 
type were listed among specimens of E. 
duncani identified by Woodruff and Cart- 
wright (1967). It is likely that other speci- 
mens of E. mexicana are confused with E. 
duncani in other collections. To our knowl- 
edge, no specimens of E. duncani have 
been collected in Mexico, although it al- 
most certainly occurs at least in northern 
Sonora, having been collected at Douglas, 
Arizona, on the Mexican border. The host 
of E. mexicana is likely Atta mexicana 
(Smith) (which is common at the type lo- 
cality in Sonora): therefore it also could oc- 
cur in Arizona where A. mexicana exists in 
a small section of Organ Pipe National 

Euparixia boliviano Ciordon and 
McCleve, new species 

(Figs. 9, 10) 

Description. — Length 4,7 mm, width 2.1 
mm. Elongate, convex, feebly shiny, dark 
reddish brown. 

Head: Clypeus distinctly emarginate 
with angles rounded, sides straight to very 
slight genal notch, apical margin dorsally, 
feebly recurved at center and ventrally in- 
dexed at center: genae feebly produced, 
rounded. Surface finely and densely punc- 
tate between eyes, punctures separated by a 
diameter or less, finer at sides and anteri- 
orly, changing to minute granules in ante- 
rior '/3, granules continuing to anterior mar- 


Figs. 9-12. Pronota and elytra. 9, Euparixia holiviami. proiidtum. 10. fi. holiviaua. elytra \\. E. cdinphi'lli. 
pronotum. 12. E. camphclli. elytra. 

gin. Frontal suture barely perceptible. Eyes 
not visible when head in repose. Lateral 
margin of gena with a weak, punctate sul- 
cus beginning near anterior margin and 
deepened gradually near eye into a deep 

pore, and with short carina at margin of 
gena adjacent to eye. 

Pronotum: Broader than long ( 1 .8 X 1.2 
lum), very convex (Fig. 9). Basal margin 
with single distinct row of contiguous large 


punctures. Basal Vi of disc with large, near- 
ly contiguous punctures becoming gradu- 
ally smaller and sparser anteriorly, small 
punctures absent. Explanate sides impunc- 
tate. Oblique depressions on pronotum dis- 
tinct, filled with about 20 large punctures in 
about 2 rows. Anterior angle abruptly 
broadly rounded. Posterior angles obsolete. 
Explanate margins with angle at anterior ¥3, 
then strongly narrowed to base. Pronotum 
at base much narrower than elytral base. 

Elytron: Elongate oval, widest at apical 
Vr, intervals convex, not carinate except in- 
tervals 7-9 weakly carinate (Fig. 10). each 
interval with row of minute granules barely 
visible at 50X. separated by their own di- 
ameter. All 10 striae with series of deep 
punctures separated by less than diameter 
of a puncture. Scutellum narrow, with 
sharply defined depression at base. 

Sternum: Prosternal process broadly 
rounded posteriorly, disc strongly margined 
at base, less so laterally. Mesosternum with 
coxae separated by 1 Vi length of middle 
trochanter; median depression clearly mar- 
gined, with strong median carina, depres- 
sion in basal % somewhat shiny, micro- 
sculpture evident at 50x, anterior Vi of de- 
pression slightly raised nearly to level of 
margin and arcuately delimited behind on a 
line beginning opposite anterior Vi of mid- 
dle coxa, curving anteriorly toward proster- 
nal process and continued to middle of op- 
posite coxa; raised anterior Vi of depression 
irregularly shiny, rugose, punctate. Meso- 
and metastema divided by transverse cari- 
na, carina smooth, without fine carinae. Lat- 
ter carina gives way to long, coarse, deep 
median sulcus. Carina bordering posterior 
portion of middle coxae large, separated 
from disc of metasternum by deep, variably 
punctate sulcus, remainder of disc with fine 
to moderate punctures except for shiny im- 
punctate area between posterior end of me- 
dian sulcus and posterior coxae; depressed 
area anterior to posterior coxa deep, sharply 
bordered, confusedly rugulose, narrowly 
smooth and shiny adjacent to middle coxa; 
anterior and lateral marains between me- 

socoxa and epipleuron with punctures fine, 
surface shiny between punctures. 

Abdomen: First sternum with irregularly 
spaced, longitudinal carinae on posterior 
margin forming very small quadrate cells 
open behind. Sterna 2-3 not depressed on 
anterior margin, with narrow row of fine, 
dense, nearly contiguous cells; sterna 4-5 
with large, elongate, narrow cells on ante- 
rior margin separated by narrow, longitu- 
dinal carinae. Posterior margin of sterna 4 
with row of small, irregular tubercles. Fifth 
sternum impunctate medially. Pygidium 
with raised, shiny, apical margin thickened 
at middle; disc depressed, shiny, with short 
median carina at basal center and several 
small tubercles on each side near basal mar- 
gin; basal Vi of pygidium above ridge 
strongly alutaceous. with a strong, longitu- 
dinal sulcus. 

Legs: Anterior tibia tridentate; apical 
tooth evenly curved outward. Apical spur 
elongate, narrow, slightly curved. Anterior 
trochanter prominent, shiny dorsally, ru- 
gose, punctate ventrally. Both anterior fem- 
ora with 1 seta near trochanter. Anterior 
femoral groove and about Vi of dorsal sur- 
face covered with short golden setae. Mid- 
dle femoral marginal groove complete from 
apex to apex of trochanter. Posterior fem- 
oral marginal groove extending from apex 
to apex of trochanter. Middle and posterior 
femora with small punctures throughout, 
punctures separated by less than to 3 times 
diameter of a puncture, each puncture with 
very short, stout seta. Posterior coxa be- 
tween trochanters smooth with weak, sparse 

Type material. — Holotype i: Bolivia, 
Santa Cruz, 3.7 km SSE. Buena Vista. Ho- 
tel Flora & Fauna, 430 m. 5-15-XI-2001, 
M.C. Thomas and B.K. Dozier, tropical 
transition forest, BLT. (MHN). 

Distribution. — Known only from the 
type locality. 

Material examined. — The holotype. 

Etymology. — The species name refers to 
the country where the holotype specimen 
was collected. 


Remarks. — This species is in the group 
with basally constricted pronota abruptly 
narrowed in the basal half, a group typified 
by E. campbelli. The noncarinate discal in- 
tervals are similar only to those of E. cos- 
taricensis which has a broad pronotum con- 
stricted only in the basal third. 

Eiiparixia campbelli Gordon and 
McCleve, new species 

(Figs. 11. 12) 

Description. — Length 5.2 mm, width 2.2 
mm. Elongate, convex, dull, dark brown. 

Head: Clypeus widely truncate between 
weak angles, sides obliquely truncate to 
weak genal notch, anterior margin barely 
reflected dorsally at center, inflexed ven- 
trally at middle. Surface densely, deeply, 
coarsely and regularly punctate between 
eyes, some punctures elongate, punctures 
sharing margins in reticulate pattern becom- 
ing smaller in transverse, complete band be- 
tween anterior margins of eyes, then larger 
again until gradually becoming smaller, giv- 
ing way to minute granulations anteriorly 
on clypeus. Each puncture with short, mi- 
nute, erect golden seta barely visible at 
50 X. Deep sulcus present along lateral mar- 
gin of gena with deep pore at anterior inner 
corner of eye, continued weakly above eye 
with short, sharp carina adjacent to eye. 

Pronotum: Broader than long (2.0 X 1 .4 
mm), very strongly convex. Almost entire 
dorsal surface covered with deep, coarse, 
regular, adjacent punctures much like a 
honeycomb, punctures becoming smaller 
and shallower near anterior margin, con- 
tinuing to anterior margin. Minute, short, 
semierect, posteriorly projecting, golden 
seta present on anterior rim of each punc- 
ture. Punctures on lateral explanations be- 
coming shallower but larger, then indistinct, 
giving way to irregular, minute granula- 
tions. Oblique depressions not evident ex- 
cept as a group of about 10 slightly shal- 
lower punctures where depressions should 
be; longitudinal groove at base lacking. An- 
terior angles narrowly rounded, acute. Ex- 
planate margins short, terminated with 

sharp right angle at anterior Vi. followed by 
long basal constriction (Fig. 1 1 ); smooth, 
shiny sulcus present along lateral margin of 
basal constriction, merging with reticulate 
punctures near base. 

Elytron: Elongate oval, widest at apical 
Vs, intervals strongly carinate, at 50X show- 
ing single row of minute adjacent granules 
each with extremely minute seta visible in 
profile at 50X; sutural interval with prom- 
inent, distinct, single row of discrete punc- 
tures visible at 25 X. Striae deep, punctures 
deep, small, sharply defined, a small, acute 
projection from carina on each side ap- 
proaching each puncture; strial punctures 
separated by 1 to 2 their own diameter, area 
between shiny (Fig. 12). Scutellum narrow, 
elongate, with deep, porelike depression at 

Sternum: Prosternal process broadly ar- 
cuate at base, all margins raised and shiny, 
disc depressed. Mesosternum with coxae 
separated basally by more than length of 
middle trochanter; median depressed area 
smooth, shiny, with microreticulation evi- 
dent at 50 X, becoming irregular, dull near 
prosternum. Meso- and metasterna separat- 
ed by fine, transverse carina having short, 
longitudinal extensions; carina on meta- 
sternum succeeded by long, coarse, broad, 
deep sulcus irregularly and minutely rugose 
at bottom. Sulcus bordering middle coxa 
broad, deep, punctate at bottom. Remainder 
of disc shiny, covered with moderate to 
large punctures, most punctures except 
largest bearing 1 minute, light colored seta 
each; sides with punctures larger, less well 
defined than on disc; depressed area in front 
of posterior coxa weakly shiny, with shal- 
low punctures. 

Abdomen: First sternum with widely 
spaced, short carinae in posterior Vi. not 
forming distinct quadrate cells. Sterna 2-5 
depressed on anterior Vi. depression with 
widely, inegularly spaced longitudinal ridg- 
es, depressed area more extensive on each 
subsequent sternum. Disc of sterna 2—3 
with variably sized, moderate to large punc- 
tures; posterior margin of same sterna with 


moderate partial punctures givins: margin 
scalloped effect. 

Legs: Anterior tibia tridentate. apical 
tooth slender, longer than other 2, bent out- 
ward at right angle to tibia; apical spur 
elongate, slender; anterior trochanter large, 
prominent, shiny, impunctate dorsally, con- 
cave, rugose ventrally; inner margin with 
numerous minute setae, not clearly crenu- 
late. Anterior femur with 3 setae near tro- 
chanter on left side, 2 on right side. Ante- 
rior femoral groove and about Vi dorsal sur- 
face of femur covered with fine, semiap- 
pressed golden setae; posterior surface of 
femur dull, covered with large, overlapping 
punctures. Middle femoral marginal groove 
complete from apex to near trochanter; pos- 
terior femur with marginal groove extended 
only Vi distance from apex to trochanter, 
strongly incised. 

Variation. — The female paratype is 4.1 
mm long and has 3 setae on each anterior 
femur near trochanter 

Type material. — Holotype 6 : Guatemala. 
Finca San Rafael Olimpo, Cuyatenango, 
Such.(Suchitepequez), 1.21.1966, J. M. 
Campbell, ex. large detritus cavity of Atta 
sp., 6-8 ft. deep (CNC). Paratype, 1 9, 
same data as holotype (CNC). 

Distribution. — Known only from the 
type locality. 

Material examined. — The holotype and 
one paratype. 

Etymology. — This species is named for 
the collector of the type series, J. M. Camp- 

bell, in recognition of his many contribu- 
tions to the study of Coleoptera. 

Remarks. — The shape of the pronotum. 
with only a single strong angle on the ex- 
planate margin, is similar to Eiiparixia hrii- 
neri and E. moseri. However E. camphelli 
lacks any indication of a weak angle behind 
the prominent angle of the lateral margin. 
The pronotal punctation of E. camphelli is 
also unique in being of uniform size and 
completely covering all but the explanate 
margins in a crowded honeycomb pattern. 

For loan of specimens, we thank Gloria 
House (USNM) and an unknown curator 
(CNC). A special thanks to Paul Skelley 
(FSCA) for photographs of surface sculp- 
ture of all species discussed, and for the E. 
bolivana specimen for study. The SEMs 
were taken by Paul Skelley at the Univer- 
sity of Florida, ICBR, Electron Microscopy 
Core Laboratory, Gainesville, PL. For man- 
uscript review we are indebted to Paul Skel- 
ley (FSCA), and Alma Solis and Allen 
Norrbom, Systematic Entomology Labora- 
tory. USDA. Washington. DC. 

Literature Cited 

Brown. W. J. 1927. Two new North American genera 
111 the tribe Eupariini (Coleoptera). Canadian En- 
tomologist 59: 288-289. 

Woodruff. R. E. and O. L. Cartwright. 1967. A review 
of the genus Eiiparixia with description.s of a new 
species from nests of leaf-cutting ants in Louisi- 
ana (Coleoptera: Scarabaeidae). Proceedings of 
the United States National Museum 123: 1-21. 

105(3). 2003. pp. 698-707 





Paui M. Marsh and Scotp R. Shaw 

(PMM) Cooperating Scientist, Systematic Entomology Laboratory, PSI, Agricultural 
Research Service. U.S. Department of Agriculture, c/o National Museum of Natural His- 
tory, Smithsonian Institution, Washington, DC 2().'i6()-()160, U.S.A. (correspondence ad- 
dress: P.O. Box 384, North Newton, KS 671 17. U.S.A.) (e-mail:; 
(SRS) University of Wyoming Insect Museum, Department of Renewable Resources, 
University of Wyoming, Laramie. WY 82071-3354. U.S.A. 

Ahstnut. — Six North American species in four species-groups of the genus Aleiodes 
are treated. The coiiipre.'isor (Herrich-Schaeffer) species-group includes A. /yaliiuilus (Wal- 
ley), new combination and A. pahnatoides. new species; the iifei ( Walley) species-group 
includes only A. iifci (Walley). new combination; the gressitti (Muesebeck) species-group 
includes A. lissos, new species; and (he procerus species-group includes A. granulatus 
(DeGant). new combination, and A. (iiifiustipciniis, new species. Other new combinations 
proposed are: A. (ilii;li<ircii\ls (Quadri). new combination; A. compressor (Herrich- 
Schaeffer), new combination; A. i>rcssitti (Muesebeck), new combination; and A. iicir- 
(iimdc (Rohwer). new combination. 

Key Words: Hymcnoptera, Uraconidae. parasitoids 

The rogadine braconid genus Aleiodes 
Wesmael is worldwide in distribution but is 
particularly species-rich in the Holarclic 
Region. Aleiodes is well diversified in 
North America, with at least 90 species in 
the United States and Canada (S. Shaw et 
al. 1997). This study is the seventh paper 
in a series on Aleiodes species-groups, in- 
iciuied to provide a complete revision of the 
genus for North America (see S. Shaw et 
al. 1997, 1998a. 1998b; Marsh and S. Shaw 
1998, 1999, 2001). The four groups cov- 
ered here are small, with only 13 included 
Holarctic species, six of which occur in 
North America. This paper treats several 
minor and unrelated groups that are not fre- 
quently encountered, but nevertheless must 
be treated to complete our revisit)n of North 

American species. For convenience, these 
are being consolidated into one paper. Be- 
cause our intent is to provide a revision of 
the North American species, species treat- 
ments are limited to the Nearctic fauna. 

Aleiodes species are koinobiont endopar- 
asitoids of lepidopterous larvae, especially 
macrolepidoptera of the superfamilies Noc- 
tuoidea and Geometroidea. and to a lesser 
extent. Arctioidea. Sphingoidea. and Papi- 
lionoidea (S. Shaw et al. 1997). The method 
of parasitism, unique to the tribe Rogadini, 
is noteworthy: the Aleiodes larva completes 
its feeding and pupates within the shrunken 
and mummified remains of the host cater- 
pillar. In all known cases, the form of ihe 
numnny caused by a particular Aleiodes 
species is characteristic for liial host and 


paiasitoid, so mummified remains are of 
considerable diagnostic value and should be 
retained with the parasitoid when reared. 
For a more complete discussion of Aleiodes 
biology, readers may refer to M. Shaw 
(1983, 1994), M. Shaw and Huddleston 
( 1991), S. Shaw (1995) and S. Shaw et al. 
( 1997). Very little is know about the biol- 
ogy of the species of the groups included 
in this paper, but the few records indicate 
parasitism of noctuid and arctiid larvae. 


Species covered in this paper can be 
identified as members of the subfamily Ro- 
gadinae using the keys of S. Shaw (1995), 
M. Shaw and Huddleston (1991), or Shar- 
key (1997). Our definition of Aleiocles fol- 
lows that of S. Shaw (1993), S. Shaw et al. 
(1997) and van Achterberg (1991). Speci- 
mens can be determined as Aleiocles using 
the key of Shaw ( 1997). The species-groups 
of North American Aleiodes can be identi- 
fied using the key provided in S. Shaw et 
al. (1997). Fortier (1997) and Fortier and 
Shaw (1999) provided a cladistic analysis 
of the Aleiodes species-groups. 

Terminology follows that used for Aleio- 
des by S. Shaw et al. (1997), S. Shaw 
(1993), and Marsh (1989). Microsculpture 
terminology follows that of Harris (1979). 
Wing vein terminology agrees with the sys- 
tem adopted by Wharton et al. (1997). A 
labeled diagram of wing veins was provid- 
ed by S. Shaw et al. (1997). 

Acronyms for collections where type ma- 
terial is deposited are as follows: CNC (Ca- 
nadian National Collection, Ottawa, Ontar- 
io, Canada), CUI (Cornell University, Ith- 
aca. New York), FSCA (Florida State Col- 
lection of Arthropods, Gainesville, FL), 
MSU (Michigan State University, East Lan- 
sing, MI), NCDA (North Carolina Depart- 
ment of Agriculture, Raleigh, NC): RMSEL 
(Rocky Mountain Systematic Entomology 
Laboratory, University of Wytiming, Lara- 
mie, WY), UCD (University of California, 
Davis, CA), UCR (University of California. 
Riverside. CA), UK (University of Kansas. 

Lawrence. KS), USNM (National Museum 
of Natural History, Smithsonian histitution. 
Washington, DC). 

Aleiodes compressor Species-Group 

Included species. — A. com/yressor (Her- 
rich-Schaeffer 1838), new combination 
(Europe); ,4. cdii>ltareiisi (Quadri 1933), 
new combination (India, Spain); /\. /nil- 
imitiis (Walley 1941), new combination 
(North America); A. pahiuitoides. new spe- 

Diagnostic characters. — First metasomal 
tergum (petiole) parallel-sided (Fig. 8) or 
nearly so, apex of metasoma laterally com- 
pressed in some females (Fig. 6); hind wing 
vein RS sinuate, marginal cell narrowest in 
middle (Fig. 2). 

Comments. — Species of this group are 
easily distinguished by the parallel-sided 
petiole, compressed metasoma, and form a 
monophyletic group within Aleiodes (For- 
tier and S. Shaw 1999). These species have 
been previously placed in the genus Petal- 
odes Wesmael; however, van Achterberg 
(1991) synonymized this genus with Aleio- 
des. Aleiodes ufei (Wallley) was also in- 
cluded in Petcdodes based upon the nearly 
parallel-sided petit)le, but it has distinctly 
different venation in the hind wing and is 
placed in its own species-group (see Fortier 
and S. Shaw 1999). 

M. Shaw ( 1994) stated that the European 
species, A. compressor, "has a blade-like 
gaster and parasitizes a common geometrid 
that lives concealed in spun leaves in its 
spring generation, and later in the summer 
a notodontid living in similar spun leaf 
packets." If the compressed metasoma 
(gaster) is a useful adaptation for getting at 
concealed hosts, then this may explain the 
convergent evolution of this lealure in the 
ufei group. 

Key to the North American Species of 
THE Compressor Spe-cies-Grolu' 

I. Apex i)t metasomal tcrga ^-(^ brown; lateral 

me.sonolal lobes usually brown 

IHiliiuiioidcs. new speeies 


Figs. 1-4. Wings oi Ateiodes species. 1, A. ufei. 2, A. palmcitoides. 3, A. Hssos. 4. A. 

- Metasomal lerga 3-6 entirely orange; lateral 
niesonotal lobes orange palmatux (Walleyl 

Aleiodes palmatoides Marsh and Shaw, 
new species 

(Fig. 2) 

Female. — Body color: head including 
antenna and mouth parts honey yellow, 
ocellar triangle black: mesosoma honey yel- 
low, with lateral mesonotal lobes and pron- 
otum, mesopleuron and propodeum dorsal- 
ly brown; metasoma honey yellow, first and 
second terga brown laterally, terga 3-5 
brown apically, venter yellow; legs yellow, 
hind feinur light brown apically; wings hy- 
aline, veins brown, stigma bicolored brown 
with yellow at apex and base, tegula yel- 
low. Body length: 4.0-5.5 mm; fore wing 
length, 3.0-3.5 mm. Head: eyes and ocelli 
large, 33-35 antennomeres, flagellomeres 
slightly longer than wide; malar space 
short, less than basal width of mandible and 
about % eye height; temple narrow, about 
Vi eye width; occipital carina weak or ab- 
sent on vertex, meeting hypostomal carina; 
oral space small and circular, diameter less 

than basal width of mandible and about Vi 
face height; clypeus weakly swollen; ocelli 
large, ocellocular distance about % diameter 
of lateral ocellus; head entirely coriaceous; 
palpi not swollen; mandibles small, tips not 
crossing when closed. Mesosoma: prono- 
tum rugulose laterally; mesonotum and scu- 
tellum coriaceous, notauli weakly scrobic- 
ulate. meeting in small triangular rugose 
area before scutellum; mesopleuron coria- 
ceous, often weakly rugulose medially, sub- 
alar sulcus rugose, sternaulus absent; pro- 
podeum rugose coriaceous dorsally, coria- 
ceous laterally, median carina complete. 
Legs: tarsal claws not pectinate; hind coxa 
finely coriaceous dorsally. Wings (Fig. 2): 
fore wing with vein r V3 length of 3RSa and 
Vi length of m-cu, vein Icu-a beyond IM 
by distance slightly greater than length of 
Icu-a, vein ICUa Vi length of ICUb; hind 
wing with vein RS slightly sinuate, margin- 
al cell nanowest in middle, vein r-m slight- 
ly shorter than IM, vein IM about % length 
of M+CU, vein m-cu weakly indicated. 
Metasoma: first tergum costate. apical 


width only slightly greater than basal width, 
median carina complete; second terguni 
costate, apical width equal to or less than 
basal width, median carina complete; third 
tergum costate on basal %, smooth on apical 
Va. median carina absent or weakly present 
on basal half; remainder of terga smooth, 
fourth tergum occasionally with few costae 
at base; ovipositor about Vi length of hind 

Male. — Essentially as in female; 38-39 
antennomeres, flagellomeres 3 times longer 
than wide; fourth metasomal tergum strong- 
ly costate. 

Holotype. — Female: VIRGINIA. Louisa 
Co.. 4 mi. S. Cuckoo. July 4-18. 1989. J. 
Kloke and D. R. Smith. Malaise trap. De- 
posited in USNM. 

Paratypes. — VIRGINIA: 5 9,3 6. same 
data as holotype, dates of July 4-October 
6. 1989; 2 9. Essex Co., 1 mi. S.E. Dunns- 
ville. September 17-October 10, 1991 and 
May 25-June 5. 1991. Malaise trap, D. R. 
Smith. NORTH CAROLINA: 3 9, Orange 
Co., Chapel Hill. September 9, 1975 and 
August 1. 1976. Malaise trap. Deposited in 

Distribution. — Known only from the 
type localities in Virginia and North Caro- 

Biology. — Unknown. 

Comments. — This species differs from 
other North American species by the nearly 
parallel-sided first metasomal tergum, 
which is similar to A. palinatiis from which 
is is distinguished by the brown markings 
on the mesonotum and metasomal terga. 

Etymology. — The specific name refers to 
the similarity of this species to A. pciliiuitiis. 

Aleiodes palmatiis (Walley). 
new combination 

(Figs. 6. 8) 

Petalodes pahuatiis Walley 1941: 214. 

Diagnosis. — Body unicolored honey yel- 
low, stigma bicolored; body length, 6.0-7.0 
mm; 33-35 antennomeres; malar space 
shorter than basal width of mandible; face 

rugo-coriaceous, frons, vertex and temple 
coriaceous; oral opening small and circular, 
diameter about equal to basal width of man- 
dible; pronotum rugose; mesonotum and 
scutellum coriaceous, notauli weakly scro- 
biculate anteriorly, meeting in weak ob- 
scure rugulose area before scutellum; me- 
sonotum coriaceous, subalar sulcus weakly 
rugulose, sternaulus absent; propodeum ru- 
gose coriaceous dorsally, coriaceous later- 
ally, median carina complete; first metaso- 
mal tergum (Fig. 8) costate, basal width 
about equal to apical width, sides parallel, 
length nearly twice apical width, median 
carina complete although weak apically; 
second tergum costate, longer than wide, 
apical width slightly less than that of first 
tergum, median carina not complete; third 
tergum costate on basal half, coriaceous on 
apical half, median carina absent; remainder 
of terga coriaceous, laterally compressed 
from apical half of second tergum to tip of 
metasoma (Fig. 6); forewing with vein Icu- 
a beyond vein IM by distance greater than 
length of Icu-a; hind wing with vein RS 
sinuate, marginal cell naiTowest in middle, 
vein Ir-m about equal to IM, vein m-cu 
weak, only short stub present; tarsal claws 
with 3-4 thin spines at base. 

Type material examined. — Peuilodes 
paliuatiis Walley. holotype female, BRIT- 
ISH COLUMBIA. Canim Lake [CNC]. 

Distribution. — Ontario west to British 
Columbia, south to Wisconsin, Minnesota, 
Utah and California. 

Biology. — Recorded as a solitary para- 
sitoid of Nycteola cinereana Neumoegen 
and Dyar (Noctuidae). Also reared from an 
undetermined microlepidopteran on Popii- 
liis treinuloides Michx. 

Comments. — Aleiodes palmatus is distin- 
guished from A. palmatoides by the honey 
yellow mesonotum and metasoma (in A. 
palnuitoides the mesonotal lobes and apical 
borders of metasomal terga 3-6 are brown). 

Aleiodes ufei Species-Group 

Included species. — A. ufei (Walley) 
1941, new combination (North America). 


Diagnostic characters. — First metasomal 
tergum (petiole) parallel-sided, apex of me- 
tasoma laterally compressed in females; 
hind wing vein RS straight, marginal cell 
naiTowest at base, widening gradually to 
apex (Fig. 1). 

Comments. — Only one species is includ- 
ed in this species-group. It also has the 
nearly parallel-sided petiole as in the com- 
pressor species-group but is distinguished 
by the straight hind wing vein RS and the 
gradually widening marginal cell and by its 
bicolored black and orange body. Cladistic 
analysis by Fortier and S. Shaw (1999) in- 
dicated that this species is not closely re- 
lated to members of the compressor group, 
despite superficial resemblance in the form 
of the metasoma. 

Aleiodes iifei (Walley). new combination 

(Fig. 1) 

Petalodes iifei Walley 1941: 215. 

Diagnosis. — Body bicolored black and 
orange, head, antenna and mesosoma black, 
first and second metasomal terga orange, re- 
mainder of terga black, third tergum occa- 
sionally orange on basal Vi, legs orange, 
apex of hind tibia and tarsus black, wings 
hyaline, veins brown, tegula black: body 
length, 8.0-9.0 mm; 49-55 antennomeres; 
malar space longer than basal width of 
mandible; head entirely punctate and shin- 
ing; ocellocular distance slightly longer 
than diameter of lateral ocellus; pronotum 
rugose, mesonotum and scutellum minutely 
punctate and shining, mesopleuron punctate 
and shining, subalar sulcus and sternaulus 
rugose; propodeum rugose dorsally, smooth 
laterally, median carina complete; first me- 
tasomal tergum parallel sided, apical width 
equal to basal width, first and second terga 
costate rugose, median carina complete, 
third tergum costate on basal V3, remainder 
of terga smooth, apical metasomal seg- 
ments laterally compressed; vein Icu-a of 
fore wing beyond IM by distance slightly 
greater than length of Icu-a; marginal cell 
of hind wing gradually widening to apex. 

vein RS straight, vein m-cu present (Fig. 1); 
tarsal claws not pectinate. 

Type material examined. — Petalodes iifei 
Walley, holotype female. Lake Almanor, 
California [USNM]. 

Distribution. — Specimens have been ex- 
amined from California, Nevada, and Brit- 
ish Columbia. 

Biology. — The type series was reared 
from Ufeus plicatiis Grote (Noctuidae). 

Comments. — This species is superficially 
similar in coloration to other bicolored 
black and orange species, such as A. ter- 
iniiudis Cresson and A. cibdomiiuilis Cres- 
son, but is distinguished by the parallel-sid- 
ed first metasomal segment and the laterally 
compressed apical metasomal segments. 

Aleiodes gressitti Species-Group 

Included species. — A. f;rcssitti (Muese- 
beck 1964), new combination (Campbell 
Islands); A. lissos new species. 

Diagnostic characters. — Metasomal terga 
very finely coriaceous and shining, appear- 
ing smooth (Fig. 7); fore wing vein Icu-a 
beyond IM by 1.5 times length of Icu-a; 
hind wing vein RS slightly sinuate, margin- 
al cell narrowest in middle (Fig. 3). 

Comments. — This is a monophyletic 
group (Fortier and Shaw 1999) of only two 
species at present which is defined by the 
nearly smooth and shining metasomal terga, 
which does not occur in any other species 
we have studied. 

Aleiodes lissos Marsh and Shaw, 
new species 

(Figs. 3, 7) 

Female. — Body color: entire body in- 
cluding legs honey yellow, flagellum grad- 
ually turning brown apically, wing veins 
light brown, stigma and vein C-I-Sc-I-R yel- 
low. Body length: 5-6 mm; fore wing 
length. 5-6 mm. Head: 38 antennomeres, 
first flagellomere slightly longer than sec- 
ond, all flagellomeres longer than wide; ma- 
lar space slightly longer than basal width of 
mandible and V3 eye height; temple Vi eye 
width; occipital carina meeting hypostomal 


Figs. .3-8. Morphology of Aleiodes species, 5. Pronotum. lateral view, A. angustipeimis. 6, Metasoma. lateral 
I'iew. A. pulnianis. 7. Metasoma. dorsal view. .4. lis.sos. 8. First metasomal tergum. dorsal view. A. palmcitus. 

carina; oral space small and circular, width 
slightly less than malar space and about Vi 
face height; clypeus swollen; ocelli mod- 
erate size, ocellocular distance equal tii di- 

ameter of lateral ocellus; face coriaceous 
with few rugae below antennae, median 
ridge between antennal sockets; frons cori- 
aceous, rugose behind antennae; verte.x and 



temples coriaceous: maxillary palpus not 
swollen; mandibles small. Mesosomo: pron- 
otum coriaceous, weakly rugose medially; 
mesonotum and scutellum coriaceous; no- 
tauli scrobiculate, meeting in wide rugose 
area; prescutellar furrow wide, with one 
cross carina; mesopleuron finely coria- 
ceous, subalar sulcus weakly rugose; ster- 
naulus absent; propodeum coriaceous, with 
only short median carina at base. Legs: tar- 
sal claws not pectinate but with 3-4 large 
spines at base; inner spur of hind tibia Vi 
length of hind basitarsus; hind coxa coria- 
ceous dorsally. Wings (Fig. 3); fore wing 
with vein r % length of 3RSa and % length 
of m-cu, vein Icu-a beyond IM by 1.5 
times length of Icu-a, vein ICUa slightly 
less than '2 ICUb; hind wing with vein RS 
\ery slightly arched in middle, marginal 
cell naiTowest in middle, vein fvH-CU 
slightly longer than IM, vein m-cu weak. 
Metasoina (Fig. 7); all terga very finely co- 
riaceous and shining; first tergum slightly 
wider at apex than length; median carina 
absent on all terga; ovipositor about % 
length of hind basitarsus. 

Male. — Essentially as in female. 

Holotype.— Female: CALIFORNIA. 
Monterey Co., Monterey Peninsula, August 
9, 1978, C. P. Ohmart coll., reared from 
Halisidota consohrina? Deposited in 

Paratypes.— CALIFORNIA; \ 9. I 6. 
same data as holotype; 1 9 , Berkeley, April 
20, 1935; I 6. Marin Co,, parasitic on larva 
of Halisidota harhsii (label not clear on 
species name); 19.4 mi. W. Stanton Ran. 
HQ, Sta. Cruz Is., September 14, 1964, M. 
Irwin; 1 S. Monterey, August 10, 1938, R. 
I, Sailer. COLORADO: 1 9, Davenport 
Camp, 36 mi. S. Florence, Custer Co., July 
8, 1967, 8500 ft., F, R & M. Rindge. NEW 
MEXICO: I 9, 14 mi. N Silver City, Grant 
Co., July 8, 1961. G. C. Eickwort. ARI- 
ZONA: I 9, Mt. Lemmon, 36 mi. N Tuc- 
son, 4300 ft., July 30, 1965, R H. Freytag, 
L. P. Gibson collectors, blacklight trap. De- 
posited in USNM, RMSEL. MSU, CUI, 

Distribution. — Known only from Califor- 
nia, Arizona, New Mexico, and Colorado. 

Biology. — Three specimens of the type 
series are labeled as questionably reared 
from "Halisidota consobrina" and one 
from Halysidota harrisii Walsh (Arctiidae), 

Comments. — This species is distin- 
guished from all other North American spe- 
cies by the nearly smooth body, particularly 
the metasoma. 

Etymology. — The specific name is from 
the Greek lissos meaning smooth in refer- 
ence to the smooth metasomal terga. 

Aleiodes procerus Species-Group 

Included species. — A. procerus Wesmael, 
1838 (Europe, Japan): A. narangae (Roh- 
wer 1934), new combination (China); A. 
graniilatiis (De Gant 1936), new combi- 
nation (North America); A. crassiptes Te- 
lenga, 1969 (Mongolia); A. angitstipennis 
new species. 

Diagnostic characters. — Pronotum (Fig. 
5) with median length greater than ocell- 
ocular distance; fore wings narrow, width 
about 'A length (Fig. 4): fourth inetasomal 
tergum sculptured. 

Comments. — The procerus species- 
group will run to the dispar species-group 
in the key to species-groups in S. Shaw et 
al. (1997). Subsequent to the publishing of 
that key, Fortier (1997) and Fortier and S. 
Shaw (1999) split the procerus group from 
the dispar group, and we have followed that 
system. The procerus species-group is dis- 
tinguished by the long pronotum, the sculp- 
tured fourth metasomal terguin, and the nar- 
row wings. 

Key to the North American Species of 
THE Procerus Species-Grocp 

1. Fore wing vein Icu-a beyond IM by distance 
less than length of Icu-a. vein ICUa shorter 
than Icu-a: head, propleuron and apical meta- 
somal terga black, rest of body usually orange 
graniilarus (DeGant) 

- Fore wing vein Icu-a beyond IM by distance 
equal to or greater than length of Icu-a, vein 
ICUa longer than Icu-a: body entirely honey 
\ellovv iingiistipennis. new species 


Aleiodes aiigustipeiinis Marsh and 
Shaw, new species 

(Figs. 4. 5) 

Female. — Body color: entirely honey 
yellow, coxae, trochanters and of fem- 
ora lighter yellow, apical Vi of flagellum 
brown: wings slightly fuscous, veins 
brown, stigma and vein C + Sc + R yellow. 
Body length: 7.0 mm; fore wing length, 4.5 
mm. Head: eyes and ocelli small, not cov- 
ering most of head; 49-50 antennomeres, 
all flagellomeres longer than wide; malar 
space long, longer than basal width of man- 
dible and % eye height: temple broad, about 
% eye width: occipital carina complete, 
meeting hypostomal carina; oral opening 
small and circular, diameter about equal to 
basal width of mandible and -/=, face height: 
ocelli small, diameter of lateral ocellus 2/3 
ocellocular distance; head entirely coria- 
ceous; maxillary palpus not swollen: man- 
dibles small, tips not crossing when closed. 
Mesosonia: pronotum (Fig. 5) distinctly 
lengthened, median length about twice 
ocellocular distance and length of first fla- 
gellomere, nearly on same plane as slightly 
declivous mesonotum, costate coriaceous; 
mesonotum and scutellum coriaceous, no- 
tauli weakly scrobiculate, meeting in shal- 
low rugose area; mesopleuron coriaceous, 
subalar sulcus shallow and weakly costate, 
sternaulus weakly indicated by costate area; 
propodeum flat and not declivous apically, 
entirely costate coriaceous, median carina 
weak but complete. Legs: tarsal claws not 
pectinate; hind coxa coriaceous dorsally. 
Wings (Fig. 4): narrow, width of fore wing 
about 'A length; fore wing with vein r V3 
length of 3RSa and % length of m-cu, vein 
Icu-a beyond 1 M by distance slight greater 
than length of Icu-a, vein ICUa Va length 
of ICUb; hind wing with vein RS slightly 
sinuate, marginal cell narrowest in middle, 
vein Ir-m about V2 length of IM, vein IM 
y, length of M + CU, vein m-cu absent. Me- 
tasonui: first tergum costate coriaceous, 
longer than apical width, median carina 
complete; terga 2-4 costate coriaceous, me- 

dian carina complete on terga 2 and 3: re- 
mainder of terga finely coriaceous; ovipos- 
itor about % length of hind basitarsus. 

Male. — Essentially as in female. 

Holotype. — Female: FLORIDA. Marion 
Co., 9 mi SSW Ocala, Kings Land Country 
Estates, September 10-19, 1975, J. Wiley. 
Deposited in FSCA. 

Paratypes. — FLORIDA: 1 V. Putnam 
Co.. 2 mi NW Orange Springs. August 27- 
September 10, 1975, J. Wiley, Malaise trap; 
1 9, Gainesville, Alachua Co., July 2. 
1976, W. H. Pierce: 1 9, Highlands Co., 
Archbold Biol. Sta., April 4, 1979, H. V. 
Weems, Jr. and Sylvia Halkin. insect flight 
trap: 1 cJ, Suwannee Co., Houston, April 
28, 1924, T. H. Hubbell. Deposited in 

Distribution. — Known only from the 
type localities in Florida. 

Biology. — Unknown. 

Comments. — This species looks some- 
what similar to A. aciciilaiiis Cresson (in 
the coxalis species-group) because of the 
costate metasoma and the lengthened pron- 
otum, but it is distinct from ackulaliis by 
its narrow wings, longer pronotum and en- 
tirely yellow stigma. It is distinguished 
from granulatiis by fore wing vein Icu-a 
being farther from vein IM and by the en- 
tirely honey yellow body. 

Etymology. — The specific name is from 
the Latin angustus meaning nanow and 
penna meaning wing in reference to the 
narrow wings. 

Aleiodes graniilatus (DeGant). 
new combination 

Rogas granitlata DeGant 1930: 163-164. 

Diagnosis. — Body bicolored. head black, 
antenna and mouth parts yellow, mesosoma 
sometimes entirely black, pronotum and 
mesonotum always black, mesopleuron and 
propodeum sometimes orange, metasoma 
with first tergum orange or black, second 
and third terga orange, remainder of terga 
black, legs orange, apical half of hind femur 
and tibia black, wings hyaline, veins brown. 


Stigma bicolored brown with yellow at apex 
and base; body length 4.5 mm; 45-30 an- 
tennomeres; malar space longer than basal 
width of mandible; head entirely coria- 
ceous, face somewhat rugose; oral opening 
small and circular, diameter slightly less 
than basal width of mandible; ocelli small, 
diameter of lateral ocellus less than ocel- 
locular distance; pronotum rugose coria- 
ceous; mesonotum coriaceous; mesopleuron 
coriaceous, sternaulus absent: propodeum 
rugose coriaceous, median carina complete; 
metasomal terga 1-4 costate, median carina 
complete on terga 1-3; tarsal claws not pec- 
tinate; fore wing with vein Icu-a beyond 
vein IM by distance less than length of 
Icu-a; hind wing with vein RS arcuate, 
marginal cell narrowest medially. 

Type material examined. — Rogas gran- 
ulata DeGant. holotype female, Cleveland. 
Ohio [USNM]. 

Distribution. — Michigan, Wisconsin. 
Ohio. Connecticut. New York, North Car- 
olina, Ontario, New Brunswick. 

Biology. — Associated with cabbage in- 
fested with "Aiitographa brassiccw." 
(probably Trichoplusia ni (Hiibner) (Noc- 

Comments. — Aleiodes granuUitits is not 
a particularly common species, considering 
that its presumed host is a pest of cabbages. 
This species is apparently closely related to 
angiistipennis but differs by fore wing vein 
Icu-a being closer to vein IM and by the 
bicolored black and orange body. 

Literature Cited 

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the Afrotropical and W. Palaearctical Rogadinae 
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DeGant. E D. 1 930. Two new species of parasitic H\ - 
menoptera (Braconidae) from Ohio. Proceedings 
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menoptera: Braconidae). Journal of Hymenoptera 
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105(3), 2003, pp. 708-714 


Stephen A. Marshall and Jindrich Rohacek 

(SAM) Department of Environmental Biology. University of Guelph, Guelph, Ontario, 
Canada NIG 2W1 (e-mail;; (JR) Slezske Zemske Miizeum, 
Opava. Tyrsova 1. CZ-746 46 Opava, Czech Republic (e-mail: 

Abstract. — Podiomitra ostracotarsata, new genus and species, is described in the rare 
and poorly known subfamily Homalomitrinae (Sphaeroceridae). This fully-winged, but 
apparently flightless, species was collected in a Malaise trap in Costa Rica, probably in 
association with a winged army ant. 

Key Words: Sphaeroceridae, Homalomitrinae, amy ants. Poclioiuitra 

The subfamily Homalomitrinae was de- 
scribed by Rohacek and Marshall (1998) 
for a group of five extremely rare, bizarre- 
looking species with reduced wing vena- 
tion, an enlarged and flattened head, no 
ocelli, reduced chaetotaxy, and greatly en- 
larged legs with all tarsomeres swollen and 
with the apical tarsomere deeply excavated. 
We here report the discovery of yet another 
highly distinctive genus in this enigmatic, 
isolated subfamily. 

Podiomitra Marshall and Rohacek, 
new genus 

Type species, — Podiomitra ostracotar- 
sata Marshall and Rohacek, new species. 

Generic diagnosis. — The following char- 
acters distinguish Podiomitra from other 
homalomitrines: Antennae (Figs. 5. 6) dis- 
tinctly separated, scape small, pedicel 
markedly tapered basally. Frons strongly 
differentiated into a prominent anterior por- 
tion and a medially flat upper portion (Fig. 
5). Prosternum with setulae. Mesoscutum 
short, with only small setulae other than 
prescutellar dorsocentral setae; scutellum 
with 2 pairs of marginal scutellar setae (and 
a pair of minute setulae on disc), the apical 

pair long (Fig. 5). Anepisternum with only 
uniformly small setulae. katepisternal setae 
also reduced. Metapleuron with a single 
seta between base of halter and hind coxa 
(Fig. 1 ). Wings (Figs, 3, 4) with no venation 
beyond basal fifth; venation in basal fifth 
relatively complete but all veins very short- 
ened (including C and R veins); cell br ab- 
sent; M basally weakly sclerotized and 
fused to R4-I-5, apically tapered and re- 
duced to fold beyond dm-cu; cell dm short, 
with rounded posterior apical corner; cross- 
vein r-m fused to basal part of M, Legs 
stout and long (Fig. 1), distal tarsomeres 
distally divided into anterior and posterior 
valves, claws arising in a dorsal position 
deep between the valves (Fig, 2), Abdomen 
with poorly sclerotized terga and sterna, 
and with very long and dense setae on ter- 
ga, sterna and wrinkled pleural membrane 
(Figs, 8, 9), T7 expanded onto ventral side, 
posteriorly tapered, with a number of cup- 
like sensilla at anterior margin; S7 narrow; 
T8 and S8 short, the former transverse, the 
later posteromedially incised, both covered 
by micropubescence. 

Etymology, — The generic name Podio- 
mitra refers to the podium like frons. 


Podiomitra ostracotarsata Marshall and 
Rohacek, new species 

(Figs. 1-9) 

Species diagnosis. — Head brown, body 
otherwise pale yellow. Head as broad as 
thorax, lower frons strongly elevated and 
projecting anteriorly, upper frons depressed, 
broad and flat, ocellar triangle absent. Pal- 
pus barely visible in type, but apparently 
well developed and setose. Mesoscutum 
with uniformly small setulae except for a 
pair of posterior dorsocentral setae lying 
Hat against upper surface of scutellum. Ab- 
domen long-setose. Wing veins compressed 
into basal fifth of wing but relatively com- 
plete, R4-F5 and costa forming a loop, r-m 
reduced, and dm-cu present. Costagial seta 
absent, but costa densely setose, distal setae 
conspicuously long. 

Description. — Holotype female: Body 
length 1 .6 mm. Body and legs pale, head 
brown. Head (Figs. 5, 6) as broad as thorax, 
frons and upper part of face strongly pro- 
jecting anteriorly; eye restricted to posterior 
half of head, with relatively few (about 70), 
large facets, finely setulose. Gena broad. 
Hat on lower half, with two, thin subequal 
setae on vibrissal angle. Postgena with ex- 
panded posteroventral corner (Fig. I ). Ep- 
istoma large but not depressed or delineated 
by a shelf. Antennae narrowly but distinctly 
separated, scape short but almost contigu- 
ous and apparently fused with intervening 
Junule (Fig. 6); pedicel very narrow basally: 
lirst Hagellomere densely long setulose dis- 
tally; arista long haired, white, slightly lon- 
ger than head width. Frons (Fig. 5) with 
narrow, densely setulose orbital strips: a 
broad, flat upper frons surrounded by a sil- 
very micropubescent stripe; and a promi- 
nent, shelflike lower frons with 6-7 small, 
inclinate interfrontal setae and a flat, dense- 
ly microsetulose upper surface. Upper back 
of head setulose, outer vertical setae slight- 
ly longer than surrounding setulae, other se- 
tae indistinguishable. Ocelli and ocellar tri- 
angle absent. 

Prosterniuii membranous, with four small 

setae. Mesoscutum (Fig. 5) with small se- 
tulae, uniformly distributed laterally and 
posteriorly, medially mainly restricted to 3 
darker pigmented longitudinal stripes, and 
with a single pair of dorsocentral setae ly- 
ing flat over the scutellum. Scutellum with 
large apical setae, minute lateral setae and 
a single pair of small discal setae. Anepis- 
ternum (Fig. 1 ) with only small setulae but 
densely setulose on anterior half; metapleu- 
ron with a single seta between base of halter 
and base of leg. Wing (Figs. 3. 4) with C 
restricted to basal fifth of wing. C (espe- 
cially distally) and terminal portion of Rl 
with long setae. Sc almost invisible; R2-f-3 
and R44-5 very short; R4-I-3 curved back 
to join costa in a continuous loop bisected 
by R2 + 3. Cell br absent (reduced because 
M is basally closely attached or fused to 
R4-I-5) and M apically tapered and reduced 
to foldlike structure beyond dm-cu; cell dm 
distinct though short, with rounded poste- 
rior outer corner; crossvein r-m attached to 
M, dm-cu and CuAl distinct; Al short, ba- 
sally darkly pigmented, apically tapered. 
Anterior margin of wing beyond costa 
somewhat secondarily thickened and finely 
pubescent in contrast to posterior margin, 
which has a fringe of long hairs. Alula re- 
duced, inconspicuous. Halter long, white; 
knob very large (collapsed and Hat on 

Legs (Fig. 1) long and thick, not flat- 
tened, uniformly covered with short setulae, 
only mid tibia with a small apicoventral 
bristle; all tarsomeres thickened, first, sec- 
ond and fifth tarsomeres of hind leg equal 
in length, third and fourth shorter and each 
with a small, dark microsetulose patch ap- 
icoventrally; apex of all tarsi valvelike, 
claws displaced to dorsal position and in- 
serted between lateral valves, pulvulli fac- 
ing each other within valvelike tarsal apex 
(Fig. 2). 

Abdomen with very weakly sclerotized 
terga and sterna, all whitish yellow, densely 
and long setose, with longest setae inserted 
on darkened patches (Figs. 8, 9). Tl-l-2 
long but dorsomedially basally desclero- 


Fig. \. Piulioiiiirm osti-acolarsalii. female holotype. Wing reconstructed according to slide mounted right 
wins. Scale bar = 0.5 mm. 

tized and with setae concentrated along 
posterior margin. T3-T6 slightly transverse, 
with very long setae, particularly at poste- 
rior margin; long dense setae also inserted 
on wrinkled dilatable pleural membrane. 
S2— S6 narrower than associated terga, 
about as long as broad, with long setae at 
lateral margins, medial setae short (Fig. 9). 
Postabdomen (Figs 8, 9). T6 and S6 sim- 

ilar to those of preceding segment, includ- 
ing chaetotaxy. T7 dissimilar to all other 
homalomitrines, large, expanded laterov- 
entrally forming an incomplete posteriorly 
conical ring, with long setae inserted in the 
middle of lateral margins and with peculiar 
dish-shaped sensilla clustered at anterior 
margin. S7 rounded trapezoidal, as broad as 
S6, with 2 pairs of long setae posteriorly. 


Figs. 2-6. Podii>mitra ostracotarsiita. female hololype. 2. Mid tarsus dorsally. 3. Delail of wing venation. 
4. Right wing. 5. Head and Mesoscutum dorsally. 6. Head frontally. Scale bars: Fig. 4 = 0.5 mm. others = 0.2 
mm. Abbreviations: Al — anal vein. CuAl — cubitus, dm — discal medial cell, dm-cu — discal medial-cubital ( = 
posterior) cross-vein, M — media. Rl, R24-3. R4-I-5 — 1st. 2nd. 3rd branch of radius: r-m — radial-medial cross- 

T8 short, transverse, with small anterome- with distinct posteromedial incisit>n. hence 

dial incision and moderate setae at posterior posterior margin bilobate. all micropubes- 

margin: completely micropubescent as is cent, with fine setae posterolaterally. Sper- 

posterior half of T7. S8 narrower than T8. mathecae (Fig. 7) 1 + 1 blackish, with cup- 


Figs. 7-9. Podiomitra ostracotarsata, female holotype. 7, Spermathecae. 8. Postabdomen dorsally. 4. po 
(abdomen (plus abdominal .segments 4-5) ventrally. Scale bars: Fig. 7 = 0.05 mm, others = 0.2 nun, Abhn 
viations: T — tergum, S — sternum. 

like body proximally covered by a number 
of tubercles and completely fused (no 
boundary visible) with sclerotized part of 
spermathecal duct (as in Sphaeroinitni). 
TIO subtriangular, almost unpigmented, 
bare except for a pair of small medial setae. 
SIO larger than TIO, subcircular. with short 
marginal setae, the posteromedial pair lon- 
gest. Cercus (like TIO and SIO) without mi- 

cropubescence, with long, moderately sin- 
uate apical, dorsopreapical and lateral 
preapical setae (Fig. 8). 

Type material. — Holotype 9: COSTA 
RICA, Prov. Cartago, La Represa, Torre del 
I.C.E., entre Porras y Villegas. [hydro-elec- 
tric tower between Ponas and Villegas] 
I800m. ABR 1997. R. Delgado. Malaise 
L_N.l 86130.560 1 00. INBio collection 


#46783. Left wing with broken apical half, 
right (complete) wing removed and mount- 
ed on a microslide pinned below speciinen. 
Abdomen detached, cleared and put in tube 
with glycerine pinned below specimen. 
An unidentified mite is attached between 
the thorax and the ventral side of the ab- 

Etymology. — The specific epithet ostra- 
colarsala refers to the shell-like formation 
of the tarsal apex. 

Relationships. — Although there is no 
doubt about the inclusion of Podioinitni os- 
tracotarsata in the Homalomitrinae, it is 
not obviously more closely related to either 
one of the described genera in the subfam- 
ily, nor do the other two genera form a 
well-supported clade excluding Podioinitni. 

Podioinitra ostracotaisata superficially 
resembles Sphaeromitra Rohacek and Mar- 
shall in the size and shape of the eye, the 
short scape, and the extremely reduced 
wing venation, but only one of the eight 
putative synapomoiphies listed by Rohacek 
and Marshall (1998) for the genus Sphae- 
romitra (reduction of mesonotal chaeto- 
taxy) could apply to Podioinitra, and this 
character differs in detail between the two 
genera. Sphaeromitra has only small setu- 
lae on both the scutum and scutellum; Po- 
diomitra has distinct prescutellar dorsocen- 
tral setae and well-developed scutellar se- 
tae. Other possible synapomorphies of 
Sphaeromitra and Podioinitra include the 
postgena with expanded posteroventral cor- 
ner, discal scutellar setulae (although there 
is only 1 pair in Podiomitra). shortened 
R2-I-3. and female abdominal sternite 7 
smaller than tergite 7. 

Podiomitra is superficially very different 
from Homalomitra Morgues-Schurter in 
wing development, head shape and chae- 
totaxy, but both genera share the plesio- 
moiphic retention of crossvein dm-cu. Of 
the eight putative generic apomorphies list- 
ed by Rohacek and Marshall (1998) for 
Homalomitra. two might be shared with 
Podiomitra. Both genera have a long fringe 
of fine setae along the posterior margin of 

the hind wing, and both genera have the 
antennae nanowly separated. The latter 
character, however, is more marked in 
Homalomitra and is accentuated by the 
greatly enlarged scape, as a result of which 
the Homalomitra looks very different from 
the other genera. 

Possible synapomoiphies linking the oth- 
er homalomitrine genera (Sphaeromitra and 
Homalomitra) to the exclusion of Podio- 
mitra include the strongly flattened legs and 
enlarged scape. 

Although more putative synapomorphies 
(5) support a sister group relationship be- 
tween Sphaeromitra and Podioinitra than 
other resolutions to this three taxon set (2 
synapomorphies per alternative clade), 
some of the characters are of uncertain ho- 
mology and polarity, and more material in- 
cluding male specimens is needed to re- 
solve the generic relationships of the Ho- 

Discussion. — This remarkable subfamily 
is known from a very few specimens col- 
lected at scattered localities from through- 
out the Neotropical Region. Although most 
specimens are from Malaise traps, the mod- 
ified legs, reduced thorax, and weakened 
wings of Sphaeromitra and Homalomitra 
led Rohacek and Marshall (1998) to spec- 
ulate that they might get around by phoresy 
rather than flight. Some specimens have 
been collected in association with ecitonine 
ants, so our best guess is that homalomitri- 
nes develop in ant kitchen middens, and 
move from nest to nest by phoresy. The dis- 
covery of the new species described here, 
in which the wing is reduced to a thin, vein- 
less membrane over its distal tj. further 
suggests that these flies are phoretic. All 
homalomitrines have a pocket or cleft at the 
apex of the tarsus, but this feature is espe- 
cially conspicuous in Podiomitra ostraco- 
taisata. in which the apex of the tarsus is 
split into anterior and posterior halves, each 
half lined by a pulvillus that has rotated 90 
degrees from its usual position, one facing 
forward and one facing backward. The 
claws are reduced and in a dorsal position 


deep in the cleft. The apical structure of Marshall). We thank Manuel Soils (INBio) 

each tarsomerc is thus clearly modified for for bringing our attention to this and other 

grasping something between its valvelike valuable specimens selected from Costa Ri- 

halves. can malaise trap samples. 


This study was partly supported by Grant 
No. 206/00/0236 of the Grant Agency of 
the Czech Republic (J. Rohacek) and partly 
supported by an NSERC operating grant (S. 

Literature Cited 

Rohacek, J. and S. A. Marshall. 1998. A review of the 
Honialoinitrinae, new subfamily (Sphaeroceridae), 
with the description of the new genus and three 
new species. European Journal of Entomology 
9.<i(3): 455-491. 


105(3). 2003. pp. 715-732 




Katherine Schick. Zhiwei Lri. and Paul Goldstein 

(KS) Essig Museum of Entomology, 201 Wellman Hall 31 12. University of California, 
Berkeley. CA 94720-3112, U.S.A. (e-mail:; (ZL, PG) Di- 
vision of Insects, Department of Zoology, Field Museum of Natural History, Chicago, IL 
60605, U.S.A.; ZL, cuixent address and address for correspondence: Department of Con- 
servation, Arizona-Sonora Desert Museum, 2021 N. Kinney Road, Tucson, AZ 85743, 
U.S.A. (e-mail: zliuOOl ("'email. 

Abstract. — Phylogenetic relationships among 14 species of 3 Holarctic gall wasp gen- 
era, Diastroplnis Hartig, Gonaspis Ashmead, and Xestophanes Forster (Hymenoptera: Cy- 
nipidae: Cynipinae: Aylacini), are inferred from 43 adult morphological characters. The 
incorporation of four characters associated with gall morphology is explored, although 
only a single parsimony-informative gall character is obtained. All analyses retrieve Dias- 
troplnis as paraphyletic with respect to the monotypic genus Gonaspis. These two genera 
share several morphological synapomorphies as well as related host plant associations. 
The genus Gonaspis is thus a new junior synonym of Diastrophus. and Gonaspis po- 
tentillae is herein transfened to Diastrophus as D. potentillae Bassett. In addition, D. 
kincaidii var. aiistrior Kinsey is formally elevated to specific status, as Diastrophus aiis- 
trior Kinsey (new status). Association of Diastrophus species with Rubus. as well as 
with woody host plants, could have originated once, but the possibility of more than one 
origination cannot be excluded. In either case, at least two reversal events have contributed 
to the patterns of association of Diastrophus with Potentilla, and with herbaceous host 
plants. Reconstruction of the historical biogeography of the group using a dispersal and 
vicariance analysis model suggests that the ancestor of Diastrophus was either East Ne- 
arctic (EN) or Holarctic. The two West Palearctic (WP) species resulted from either two 
independent dispersals from EN followed by subsequent vicariance, or from a basal vi- 
cariance between WP and EN and a subsequent dispersal from EN to WP followed by a 
subsequent vicariance between the two regions. The West Nearctic species resulted from 
dispersals from EN and are relatively recent. 

Key Words: biogeography. Diastrophus, Gonaspis. host shifting, host plant growth form, 
morphology, phylogeny 

Extant gall-forming members of the fam- cini are apparently monophyletic and ap- 

ily Cynipidae (Hymenoptera) comprise five pear to have relatively conserved host plant 

of the six tribes in the subfamily Cynipinae, associations with host plants usually be- 

i.e., Aylacini, Cynipini, Eschatocerini, Pe- longing to a single genus or a few closely 

diaspidini, and Diplolepidini (Ronquist related genera (Liljeblad and Ronquist 

1999). All the cynipid tribes except Ayla- 1998). In contrast, the tribe Aylacini, com- 


prising 2 I genera, is apparently an assem- 
hiage of paiaphyletic and polyphyletic 
ciades (Ronquist 1994, Liljeblad and Ron- 
quist 1998, Ronquist 1999). and its mem- 
bers use a wide range of host plants be- 
longing to Apiaceae. Asteraceae, Brassica- 
ceae. Lamiaceae. Papaveraceae, Rosaceae. 
and Valerianaceae (Liljeblad and Ronquist 
1998, Ronquist and Liljeblad 2001 ). 

According to Liljeblad and Rt)nquist 
( 1998), all cynipid wasps inducing galls on 
woody rosid plants form a monophyletic 
clade. These rosid gallers include the tribes 
of Diplolepidini inducing galls on Rosa 
(Rosaceae), Eschatocerini on Acacia and 
Prosopis (Fabaceae), Pediaspidini on Acer 
(Aceraceae). and the species-rich Cynipini 
primarily on Qiiercits (Liljeblad and Ron- 
quist 1998, Ronquist 1999). Ronquist and 
Liljeblad (2001 ) explored various aspects of 
the host plant association t)f cynipid wasps 
on basis of the aforementioned phylogeny 
and concluded that cynipid gall wasps are 
"extremely conservative' in their associa- 
tion with plant growth form. They further 
suggested that the gall wasp ancestor is 
likely to have been an herb galler, and col- 
onization of woody hosts has occuired only 
three times, assuming that all the eight Rii- 
/?».v-galling species of Diastrophus Hartig 
form a monophyletic clade. 

The genus Diastrophus is rather unusual 
among cynipid gall wasps in two aspects of 
their host plant associations. Firstly. Dias- 
trophus, as well as Gonaspis Ashmead and 
Xestophanes Forster, is associated with ro- 
saceous hosts of the genera Rubiis. Poten- 
tilla. and Fragaria (with one species, Dias- 
trophus smilacis Ashmead inducing galls 
on the monocotyledonous Smilax [Smila- 
caceae]). in contrast to the other Aylacini 
genera which induce galls on relatively ad- 
vanced herbaceous host plants (Ronquist 
1994). Secondly, host plants of the genus 
include both woody (Ruhus and Smilax) 
and herbaceous {Potent ilia and Fragaria) 
plants, a deviation from the generally very 
conservative pattern of association with 
plant growth form in Cynipidae. Therefore, 

Other cynipids 

Other cynipids 







Fig. \. Phylogenetic relationship among Dicislro- 
pliiis. Gonaspis, and Xestoplumcs, (a) a.s .suggested by 
Nieves-Aldrey (1994). (b) as suggested by Liljeblad 
and Roni.|iiisl (1998). 

the evolution of host plant associations in 
Diastrophus could shed light on the evo- 
lution of host use in Cynipidae in general. 

Nieves-Aldrey (1994) suggested that 
Diastrophus, Gonaspis, and Xestophanes 
form a monophyletic clade (Fig. la) on the 
basis of their rosaceous host associations, 
and the presence of lobed claws and gla- 
brous sculpture on the vertex and mesotho- 
rax (Figs. 2a-c). On the basis of phyloge- 
netic analysis using morphological charac- 
ters, Liljeblad and Ronquist (1998) partially 
supported this view, but suggested that Xes- 
tophanes and the inquilinous tribe Syner- 
gini form the sister clade of (Diastro- 
phus -\- Gonaspis) (Fig. lb). Recent molec- 
ular work (Liu et al. unpublished data) sup- 
ports the view of Nieves-Aldrey ( 1994). 

Diastrophus is morphologically distinct 
among Aylacini genera by the combination 
of strongly lobed claws, subcosta and radius 
reaching to anterior margin of wing; and 
completely free third and fourth abdominal 
terga in both sexes (Fig. 2d, cf. Xestopha- 
nes poteniillae De Villers. Fig. 2e) (Nieves- 


Aldiey 1994). and thus obviously mono- 
phyletic. It is not clear whether Gonaspis. 
which comprises only one known species, 
should be included within Diastrophiis or is 
indeed a sister clade of Diastroplms. Ron- 
quist (1994) and Liljeblad and Ronquist 
( 1998) listed five synapomorphies for Dias- 
troplms + Gonaspis. However, these studies, 
for their respective puiposes. each included 
only one Diastroplms species and were not 
able to clarify the relationship among Gon- 
aspis and Diastroplms species. In fact, only 
a single morphological feature separates the 
monotypic Gonaspis from Diastroplms: the 
scutellum of Gonaspis conspicuously pro- 
jected posteriorly (Fig. 2f). This is an au- 
tapomorphic character, for which an inter- 
mediate form exists in D. ciisviitaeformis 
Osten Sacken (not illustrated). 

Biologically, the members of Diastro- 
plms. as well as those of Gonaspis and Xes- 
tophanes. are univoltine and lack the alter- 
nation of generations (heterogony) found in 
the more derived oak-galling cynipids. Like 
galls induced by other wasps of the tribe 
Aylacini, their galls are mostly simple 
swellings on stems, leaves, and roots, and 
lack complicated secondary external struc- 
tures. Cynipid wasps lumped in the tribe 
Aylacini invariably make galls not detach- 
able from the host plant. This is also true 
for Gonaspis. Xestoplianes, and most spe- 
cies of Diastroplms. regardless of host 
form, i.e., woody versus herbaceous. Only 
one species of Diastroplms. D. citscutaefor- 
luis. makes detachable galls, as do some 
Cynipini gallers of woody plants. In addi- 
tion, some members of Diastroplms asso- 
ciated with woody Rubus bushes are at- 
tacked by inquilines of the cynipid genus 
Sytiophromorpha Ashmead. Inquilines of 
cynipid gall wasps do not induce galls of 
their own. but instead live in and feed on 
galls induced by the other cynipid wasps. 
Only galls on woody host plants are known 
to be attacked by inquilines. 

While Gonaspis is a monotypic genus 
endemic to the Nearctic and Xestophanes 
includes only two species from the West 

Palearctic. the genus Diastroplms (Hyme- 
noptera: Cynipidae: Aylacini) comprises 
sixteen described species distributed 
throughout the Holarctic. Known species of 
Diastrophus include two from the West Pa- 
learctic (D. inayri Reinhard 1877 and D. 
nibi [Bouche 1834]) and one undescribed 
species from Japan in the East Palearctic 
identified on the basis of the presence of 
Diastroplms galls and rearing of the host- 
specific Synophromorpha Ashmead inqui- 
lines from these galls (Abe 1998. Abe per- 
sonal communication); one described nom- 
inate subspecies (D. k. kincaidii Gillette 
1893) and its heteronomic subspecies (D. 
kincaidii austrior Kinsey 1922) from the 
West Nearctic; and twelve East Nearctic 
species (D. bassetti Beutenmiiller 1892, D. 
cusciitaeformis Osten Sacken 1863. D. fra- 
gariae Beutenmiiller 1915, D. fusiformaiis 
Ashmead 1890, D. minimus Bassett 1900, 
D. nebulosus (Osten Sacken 1861), D. niger 
Bassett 1900, D. picens Provancher 1886, 
D. radicum Bassett 1870. D. smilacis Ash- 
mead 1896. D. tiimefactus Kinsey 1920, 
and D. tiirgidiis Bassett 1870). The two Eu- 
ropean species of Diastroplms Hartig 1840 
(Hymenoptera; Cynipidae) were recently 
reviewed (Nieves-Aldrey 1994). However, 
there has been no revisionary study of the 
Nearctic species of this genus since Beut- 
tenmueller (1909). although Burks (1979) 
compiled a list of all Nearctic species of the 
genus. Moreover, phylogenetic relation- 
ships among the Diastrophus species are 
not known and a phylogeny-based system- 
atic revision has never been attempted. 

Therefore, our goals of the present study 
are to reconstruct the phylogenetic relation- 
ships among Diastroplms species and Gon- 
aspis, based on adult morphology, and use 
the resulting cladogram to examine ( 1 ) the 
macroevolution of associations between 
wasps and their host plants and test the hy- 
pothetical monophyletic shifting from her- 
baceous host plants to the woody Rubus 
host plants, (2) the historical biogeography 
of the genus, and (3) the phylogenetic po- 
sition of Gcmaspis potentillcw in relation to 


Fig. 2. Dtiiitro/ihu.s and Xestopluiiies species, (ui hind cUm of D. nebiitosits: (b) head of D. nebiilosiis: (c) 
mesonotum of D. nebulosiis; (d) metasoma of D. nebiilosiis: (e) metasoma of X. poteiuillae: (f) mesosoma of 
D. polemillae. 

Diastrophus species, testing the phyloge- 
netic independence of the monotypic genus 
Gonaspis. as discussed by Liljeblad and 
Ronquist (1998). 

Materials and Methods 

Type specimens of 14 of the Nearctic 
species and the subspecies were examined. 

along with additional specimens compared 
to types for Nearctic and Palearctic species. 
The undescribed Japanese species is not in- 
cluded in the analysis because no speci- 
mens of this species were reared. Diastro- 
phus piceits is also excluded from the study 
because the only reported material of this 
species is its type, which we were unable 



Table 1. Taxa included in the study, their distribution and host plants. EN = East Nearctic. WN = West 
Nearctic. NA = Nearctic. and WP = West Palearctic. The undescribed species of Diaslrophus from Japan, 
reported on basis of presence of galls, is not included in the analysis. The eastern Canadian species. D. piceiis 
Provancher 1886. is not included in the .study because we were unable to locate the type. References listed are 
in addition to original references to each species, of which all have been consulted. 



Hosi PL.iii 


Duisii<ipln<\ iiiistniir 



Burks (1^74) 

D. hassetii 



Burks (1979) 

D. ciisciitaefiirmis 



Burks (1979) 

D. frageriae 


Fnigaria virginianu 

Burks (1979) 

D. fusifoniiaiis 



Burks (1979) 

D. kiiuaidii 


Riihiis parsiflonis 

Burks (1979) 

D. iiiayri 


Potenrilla argenleii 

Nieves-Aldrey (1994) 

D. minimus 


Potciuilhi caiiMlcusis 

Burks (1979) 

D. nclmlosiis 



Burks (1979) 

D. ni'^er 


Potenlilla canadensis 

Burks (1979) 

D. iculiLiim 

NA (mainly EN) 

Riibiis occidentalis 

Burks (1979) 

D. nibi 



Nieves-Aldrey (1994) 

D. smilacis 



Burks (1979) 

D. liimefactiis 



Burks (1979) 

D. liirgidii.s 


Rubus strigosus 

Burks (1979) 

Goinispis poli'iuilUw 



Weld (1950). Liu (held data) 

Xestopluiiies poteiuilliie 



Nieves-Aldrey 1 1994) 

to examine. We also noted that Beuten- 
miiller (1909) was unable to distinguish it 
from other Diastroplnis species based on 
Provancher's original description and sug- 
gested that this might not be a valid species. 
The two subspecies of D. kiiicciidii were in- 
cluded as separate operational taxonomic 
unit. Loaned material included specimens 
from six institutions and three private col- 
lections. A total of 395 specimens were ex- 
amined. Collections utilized for this study 
and the persons through whom the loans 
were kindly arranged are as follows: 

AMNH American Museum of Natural 
History. New York. NY. USA (J. 

CASC California Academy of Sciences. 
San Francisco. CA. USA (R. Zu- 

CNCI Canadian National Collection of 
Insects. Ottawa, ON. Canada (J. 

EMEC University of California, Berke- 
ley. CA. USA (C. BaiT). 

FMNH Field Museum of Natural Histo- 

ry. Chicago, IL. USA (P. Gold- 

MCZC Museum of Comparative Zoolo- 
gy, Cambridge, MA, USA (P 

UCDC University of California, Davis, 
CA. USA (S. Heydon). 

USNM National Museum of Natural His- 
tory, Smithsonian Institution, 
Washington. DC. USA (D. 

Other specimen contributors are: 

ZL Zhiwei Liu 
OP Olivier Plantard 
FR Fredrik Ronquist. 

Morphological characters for all species 
were examined and identified using a Leitz 
Wetzlar stereoinicroscope. SEM images 
were obtained for Gonaspis poteiitillae. 
Xesiophanes potentilloe. and Diastrophus 
nehttlosKs, using Hitachi S4700 Field Emis- 
sion Scanning Electron Microscope (FE- 
SEM) at the AMNH. Additional SEM im- 
ages were also taken for D. mayri, D. niger. 



and D. turgidus using an Amray 1 810 at the 
FMNH to assist character coding. Mounted 
specimens were first transferred to and kept 
in 70% ethanol for weeks, dissected in 70% 
ethanol, cleaned with kitchen ammonia, and 
washed in water and in a series of ethanol 
solutions of different concentration. Dis- 
sected body parts were then stored in 100% 
ethanol and air dried before being mounted 
to SEM stubs, and gold-coated. 

Terminology for skeletal features follows 
Richards (1977), Ronquist and Nordlander 
(1989), and Ronquist (1995). Terminology 
describing surface sculptures follows Haixis 
(1979). Phylogenetic relationships among 
the taxa were analyzed with gall morphol- 
ogy characters included and excluded, re- 
spectively, using parsimony algorithm as 
implemented in PAUP* version 4.0b8a 
(Swafford 2002). Heuristic search method 
was used with 5000 random addition se- 
quence replicates, followed by tree bisec- 
tion-reconnection branch swapping (other 
search options were default in PAUP). Cal- 
culations of Bremer support for each node 
(Bremer 1994) were programmed for batch 
processing using the same options as the 
tree searching procedure described above. 
All characters were unordered and un- 

Associations with host plant genera were 
investigated by optimization onto the re- 
sulting two best trees a posteriori using 
MacClade version 4.03 (Maddison and 
Maddison 2001). The number of host plant 
colonization events was evaluated assuming 
an ACCTRAN optimization. We evaluated 
the degree to which host associations were 
phylogenetically conserved by calculating 
their unit consistency indices. 

Historical biogeography was reconstruct- 
ed by using DIVA Version 1.2 (Ronquist 
2001) according to the dispersal-vicariance 
optimization method proposed by Ronquist 
(1997). Dispersal-vicariance analysis recon- 
structs the ancestral distribution in a given 
phylogeny without any prior assumptions 
about the form of area relationships. Spe- 
ciation is assumed to subdivide the ranges 

of widespread species into vicariant com- 
ponents: the optimal ancestral distributions 
are those that minimize the number of im- 
plied dispersal and extinction events (Ron- 
quist 1997). The DIVA method differs from 
cladistic biogeography in that it allows non- 
hierarchical area relationships, and is there- 
fore particularly useful when reconstructing 
the distribution history of groups occuiTing 
in areas that have a reticulate palaeogeo- 
graphic history, such as the Northern Hemi- 
sphere (Nordlander et al. 1996 and refer- 
ences therein). 


Character analysis. — We identified 43 
moiphological characters of adult wasps, 
and four characters of gall morphology 
(Appendix). Characters 8, 36, 42, and 45- 
47 were autapomorphic and thus parsimo- 
ny-uninformative. They were excluded 
from analysis, but included in the matrix for 
reference purposes. 

Cladistic analyses. — Parsimony analysis 
of all characters resulted in two equally most 
parsimonious trees (Length = 167; CI = 
0.36; RI = 0.44; Figs. 3a, 3b). The strict 
consensus tree is shown in Fig. 4. Excluding 
the single parsimony-infomiative gall char- 
acter resulted in the same two most parsi- 
monious trees with no change in tree topol- 
ogies, and only slight difference in tree sta- 
tistics (Length = 165, CI = 0.36, RI = 
0.44). Inclusion of gall morphology in anal- 
ysis resulted in better, though slight, support 
for three internodes (noticeably two basal in- 
ternodes) of the best trees (Fig. 4). The two 
most parsimonious trees differ only in the 
positions of D. niayri and D. ciiscutaeformis. 

As is shown in the strict consensus tree, 
the phylogenetic relationship among the 
Diastropluis species is mostly resolved. As 
we predicted, the monotypic genus Gonas- 
pis is nested within Diastrophus. The rela- 
tionship among D. niayri, D. cuscittaefor- 
mis, the clade (D. riibi. (D. fragariae, D. 
tumefactus)). and the rest of the genus at 
the base of the phylogenetic tree is not re- 
solved. The two subspecies of D. kiitcaidi, 











3 1 










1 4 





D. mayri 

D. cuscutaeformis 

D. rubi 

D. fragariae 

D. tumefactus 

D. bassettii 

D. nebulosus 


D. fusiformans 

D. austrior 

D. turgidus 

D. kincaidii 

D. smilacis 

D. radicum 

D. niger 

D. minimus 



D. rubi 

D. fragariae 

D. tumefactus 

D. cuscutaeformis 

D. mayri 

D. bassettii 

D. nebulosus 


D. fusiformans 

D. austrior 

D. turgidus 

D. kincaidii 

D. smilacis 

D. radicum 

D. niger 

D. minimus 


Fig. 3. Two equally most parsimonious trees tor DiasUopliiis. Gonaspis, and Xcslnphanes. with the number 
of apomorphic character changes for each branch indicated (tree length = 167. CI = 0.36, RI = 0.44). Program 
for phylogenetic analysis was PAUP 4.0 (beta 10) (Swofford 2002). Six characters that are parsimony-uninfor- 
mative were excluded from the analysis, but were included when calculating the number of apomorphic character 
changes on terminal internodes. 

i.e., D. k. kiucaidi and D. k. austrior. do not 
appear as sister species in the phylogeny, 
although they were shown to be closely re- 

Optimization of host plant association 
onto tree #1 (Fig. 5a) requires 6 steps (CI 
= 0.50) to explain the cun^ent pattern. It 
suggests that the basal species of Diastro- 



Table 2. Character matrix. Missing data and characters inapphcable to a particular taxon are represented by 
■■-". Polymorphism is by letters "p" (=01), "q" (=02), and "x" ( = 12). Based on result of phylogenetic 
analysis, Gonaspis is treated in the present study as a synonym of Diaslrophiis. and thus G. polenlillae = D. 
polenlillae, sensu lalo. 




















/). ansrrior 






n. bassettii 






O. cuscutaeformis 






D. frageriae 






D. fusiformans 






D. kincaidii 






D. mayri 






D. winmuis 






I), nclnilosus 






D. niiier 






D. nulicmn 






n. nihi 






D. sinilcuis 






D. nimefactiis 






D. ridsiiilKS 






pliiis was associated with Poteutilhi. which 
appears to have been re-colonized twice by 
more derived species, and that colonization 
of Riibiis has occurred once (having been 
reversed by the recolonizations of Poteiitil- 
la and one shift each to Fragaha and Smi- 
la.x). Optimization onto tree #2 (Fig. 5b) re- 
quires 7 changes (CI = 0.33) to explain the 
cuiTent pattern, an extra change over the 
previous scenario and suggests that the as- 
sociation with Riihiis is less conserved. The 
associations with the plant genera Fragaria. 
and particularly that with Sniilax. in both 
optimizations, are nested in the crown clade 
of each most parsimonious cladogram. As- 
sociations with both Potentilla and Riihiis 
are slightly more conserved in tree #1 (Fig. 
5a), in which the unit consistency indices 
of each association are 0.25 and 0.20, re- 
spectively, as compared with 0.20 and 0.17 
in tree #2 (Fig. 5b). 

Optimization of plant growth form as- 
sociations onto tree #1 (Fig. 6a) requires 
four changes (CI = 0.25) and suggests that 
the basal species of Diastrophus was asso- 
ciated with herbaceous host plants and col- 

onization of woody host plants occuired 
only once, followed by three subsequent re- 
versals to herbaceous hosts. In contrast, op- 
timization onto Tree #2 (Fig. 6b) requires 
five changes with lower consistency index 
(CI = 0.20), and is ambiguous about the 
ancestral state of plant growth form asso- 

DIVA analysis of the historical bioge- 
ography of Diastrophus based on the phy- 
logenetic relationship among species de- 
picted in Figs. 3a— b resulted in two equally 
optimal alternative reconstructions, each 
suggesting four dispersal events. Based on 
tree #1, the ancestor of Diastrophus was 
present in both West Palearctic and East 
Nearctic and a vicariance event split the 
West Palearctic D. mayri from the rest of 
the genus in East Nearctic (Fig. 7a). A clade 
later expanded its distribution to West Pa- 
learctic, followed by a vicariance event, 
which gave rise to D. ruhi in West Palearc- 
tic and D. fragariae + D. tumefactus in 
East Nearctic. The West Nearctic species D. 
aiistrior and D. kincaidii resulted from in- 
dependent dispersal events from East Ne- 




D. mayri 

D. cuscutaeformis 

D. rubi 

D. fragariae 

D. tumefactus 

D. bassettii 

D. nebulosus 


D. fusiformans 

D. austrior 

D. turgidus 

D. kincaidii 

D. smilacis 

D. radicum 

D. niger 

D. minimus a 

Fig. 4. Strict con.sensus tree of the two equally most parsimonious trees. Number above and beluw each 
branch are Bremer Support values resulting from constrained search with the single informative gall morphology 
character included and excluded. 

arctic followed by subsequent vicariance. 
The occunence of D. fusiformans is due to 
recent dispersal. Alternatively (Fig. 7b), the 
ancestor of Diastropluis was present in East 
Palearctic, and both West Palearctic species, 
i.e., D. mayri and D. rubi. are due to dis- 
persal events to West Palearctic followed by 
a vicariance event. Otherwise the two re- 
constructions do not differ (Figs. 7a-b). 


Phylogeny and systematics of Diastro- 
pluis. — The phylogenies should be viewed 
as working hypotheses about relationship of 
the group. Although resolved, the low con- 
sistency indices, retention indices, and Bre- 
mer values suggest that there is not enough 
character congruence in our data to render 
a robustly supported cladograni. External 

morphology in this group exhibits consid- 
erable homoplasy, possibly as a result of 
convergence associated with gall-forming 
life styles. An analysis based on molecular 
characters may be more informative. None- 
theless, our analysis does provide a provi- 
sional phylogeny and a preliminary basis 
for exploring the macroevolution of host 
plant associations and historical biogeog- 

The phylogenetic position of the mono- 
typic Gonaspis as nested within Diastro- 
pluis is confirmed by all analyses, thus the 
species Gonaspis potentillae (Bassett 1864) 
is hereby formally transferred to Diastro- 
pluis. where the species was originally 
placed. The genus Gonaspis was estab- 
lished by Ashmead (1897) based on Dias- 
tropluis scutcllaris Gillette 1891, which 








+^ 0- 0' 0- <5- <)■ ^'^ 0- 0- 0- 0- O' <:>• 0- 0- 0' <d- 



^o^ „ .c? . / 

^ 0- <:)■ <:)• ^'^ o- <) 0- o o- <d- o- o- q ^- 

"••^r ■••••^-, 

Rub us 



Fig. 5. Macroevolutioiiary pattern of host plant association in Diastrophiis. Go 
optimized using MaCIade 4.0 onto best tree #1 and best tree #2, respectively. 

and Xcsloplia 



was considered a junior synonym (Weld 
1952), and listed as a variety of, G. poten- 
tillae by Weld (1959). Weld's first treatment 
has become accepted (Burks 1979). There- 
fore, the genus Goiuispis Ashmead 1897 is 
a new junior synonym of Diastroplnis 
Hartig 1840. 

Diaslmphiis kincciiclii auslrior. originally 
described by Kinsey (1922) as a variety of 
/). kiiwaidii Gillette, 1893, was later treated 
as a subspecies of the latter by Weld ( 1959). 
Burks (1979) listed it as a synonym of D. 
kincciiclii. Since the two taxa do not appear 
as sister species in the cladogram, we thus 
establish D. kincciiclii van ciii.strior Kinsey 
1922 as a valid species, Dicistrophus aus- 
lrior Kinsey 1922. Both D. kiiicaiclii and D. 
tiiislrior induce galls on Riil^nx piirviflonts. 
however, D. kincciiclii is distributed in 
Washington and Oregon, and /). auslrior is 
restricted to California based on known col- 
lection records. 

Host plant associations and gall fea- 
tures. — Tree #1 suggests that an association 
with Poleniilla was the ancestral condition 
with respect for Diastroplnis. and was sec- 
ondarily derived in D. tninefactiis. and the 
respective common ancestors of the sister 
species [D. minimus + D. niger], and of |D. 
potenlillae (= Gonaspis potentlllcw) + D. 
fusiformans]. The position of the species as- 
sociated with Smila.y and Fragaria suggests 
relatively recent and unique host shifts. In 
contrast, tree #2 offers an ambiguous sce- 
nario for ancestral host plant associations of 
Diastrophus. This is a result of the shifted 
position of D. mayri. Otherwise the two 
scenarios (Figs. 5a— b) are similar. In either 
case, association with Potentilla appears to 
have been labile, with at least three inde- 
pendent origins, whereas association with 
Riihus may be more conserved. In neither 
scenario did we find a perfectly conserved 
association with any host plant genus for 
which there exist more than one associated 
Diastroplnis species (i.e., Potentilla and 

With regard to the macroevolutionary 
pattern of association with host plant 

growth form, tree #1 gives a more parsi- 
monious and straightforward reconstruction 
compared with tree #2, which requires one 
more extra step, has lower consistency, and 
is uncertain about the ancestral state of host 
plant growth form association. The first sce- 
nario suggests a single event of host shift- 
ing from herbaceous to woody hosts. In 
contrast, the second scenario suggests this 
only as a possibility, but also suggests an- 
other possibility, i.e., coloni/atitMi to woody 
hosts has occurred independently at least 
twice. In either case, reversal from woody 
hosts to herbaceous hosts has occurred at 
least twice, contrary to the general pattern 
in the clade of rosid gall makers, where col- 
onization of woody hosts occuned only 
once and no reversal were observed (Ron- 
quist and Liljeblad 2001). It is interesting 
to note the plasticity of the many aspects of 
Diatrophus biology. Most of Diastroplnis 
(sensii lata) and Xestopluines are the only 
taxa that induce galls on rosaceous host 
plants among the 156 known species in the 
tribe Aylacini; all others are associated with 
advanced herbaceous hosts. Diastroplnis is 
the only cynipid genus that has both mem- 
bers that are associated with herbaceous 
host plants and with woody hosts, contrary 
to the generally conservative association 
with plant growth forms in all other cynipid 
gall makers. As mentioned earlier, all basal 
cynipid genera are herb gallers, while the 
host plants of the rosid gallers are all 
woody. This conservative characteristic of 
association with plant growth form reaches 
its extreme in the rosid gallers comprising 
more than 1,000 species and with no single 
exception: Diplolepidini (63 species) are as- 
sociated with Rosa (Rosaceae), Eschatocer- 
ini (3 species) with Acacia and Prosopis 
(Fabaceae), Pediaspidini (2 species) with 
Acer (Aceraceae), and Cynipini (ca 1, ()()() 
spp) with Quercus (Fagaceae, with a few 
exceptions associated with other genera of 
the same plant family; Ronquist 1994, Lil- 
jeblad and Ronquist 1998, Ronquist 1999). 
Acct)rding to phylogenelic study based 
on both morphology and molecular data. 




■^ 0- 0- 0- 0- ^'^ 0- 0- Q- O- 0- 0- 0' 0- 0- 0- 






-V 0- ^- o- <:)• Q- ^^^ 0- 0- 0- 0- O' 0- 0- 0- 0- 


Fig. 6. Macroe\olutionar\ pattern of association iit' host plant growth forms in Ditislropluis. Goiuispis. and 
Xestophanes as optimized using MaClade 4.0 onto best tree #1 and best tree #2. respectively. 


the plant genera Fragaria. Potentilla. and 
Ruhus, associated with Diaslrophiis belong 
to a monophyletic clade with another genus 
Rosa in the plant family Rosaceae (Evans 
1999). Therefore, species of Diastropluis 
are still conservative in their host plant as- 
sociation, with the exception of D. smilacis. 
Once the ancestral species of the genus col- 
onized any of the genera Fragaria, Poten- 
tilla. or Ruhus. it would have become less 
restrictive for subsequent colonization of 
other genera in the same clade, from her- 
baceous to woody host or vice versa, lead- 
ing to repeated reversal events from woody 
liosts to herbaceous hosts (Figs. 6a-b) and 
thus the relatively high species diversity of 
the genus compared to most other herb- 
galling cynipid genera (cf., Ronquist and 
Liljeblad 2001). 

Like all other herb gallers, Diastropluis 
species mostly induce simple swellings on 
stems with multi-chambers. However, sev- 
eral species show derivations; D. cuscutae- 
forniis makes clusters of singular, detach- 
able, and seed-like galls, D. fragariae 
makes galls on leaf petioles of strawberry 
{Fragaria virginiana Duch.), and D. radi- 
cuiu and D. bassettii make underground 
galls on the roots of Ruhus. Indeed, the only 
parsimony-informative gall character we 
scored is the location of the galls, below or 
above ground, and that only by virtue of 
two species bearing one of two alternative 
states (underground gall-forming). Howev- 
er, this behavior appears convergent in each 
most parsimonious tree, considering that 
the galls of D. bassettii are stem galls and 
are not always below ground level (Beuten- 
mueller 1909), and hence was coded as 
polymorphic in our analysis. Our phyloge- 
netic analysis supported this by showing the 
D. bassettii and D. radicum are not sister 
species (Figs. 5a-b). 

Biogeography. — The biogeographical re- 
construction of speciation pattern based on 
tree #1 (Fig. 7a) suggested that the ancestral 
species of Diastropluis was distributed in 
both West Palearctic and East Nalearctic, 
with a basal vicariance event that separated 

D. mayri and the rest of the genus. In this 
scenerio, D. riibi originated from a vicari- 
ance event that followed dispersal of its 
common ancestral species with D. fragar- 
iae and D. cusciitaeformis, froin East Ne- 
arctic to West Palearctic. Alternatively, re- 
construction (Fig. 7b) based on the second 
best tree suggests that the common ancestor 
of Xestophanes and Diastropluis occurred 
in both West Palearctic and East Nearctic 
and had later become split through a vicar- 
iant event. Two subsequent dispersal events 
gave rise to the two West Palearctic Dias- 
tropluis species. 

The two biogeographical reconstructions 
do not differ otherwise, and all other spe- 
cies involved have an East Nearctic distri- 
bution, except D. austrior. D. kincaidii. and 
D. fusiformaus. Diastropluis austrior and 
D. kincaidii are only found in California. 
Oregon, and Washington, while D. fusifor- 
nians is also found in the East Nearctic. The 
host plant of the two species, Ruhus par- 
viflorus Nutt.), has a distribution in western 
North America and not beyond the Great 
Lake areas to the east. Diastropluis fusifor- 
nuins makes galls on Potentilla. and its 
wider distribution may be attributable to the 
distribution of its host plant. The terminal 
position of this clade indicates that coloni- 
zation of West Nearctic has been relatively 
recent event(s). 

In a forthcoming study, Engel et al. (pers. 
comm.) suggested that Aiilacidea succinea. 
a fossil species described from Baltic ainber 
(Eocene, ca 45 MYA) by Kin.sey (1937) 
and transferred therein to a newly erected 
genus, is closely related to the genera Xes- 
tophanes and Diastropluis (sensu lata). 
Therefore, the presence of Diastrophus pos- 
sibly dates as far back as 45 million years 
ago in the Eocene. Since the oldest known 
cynipid fossils with putative associations 
with Rosa is from the Oligocene (33-23 
MYA: Cockerel! 1921, Ronquist 1999) and 
the oldest fossils that are definitely cynipine 
galls are on Quercus from Middle Miocene 
(ca. 15-12.5 MYA; Waggoner and Poteet 
1996, Waggoner 1999), the species of Dias- 


a = West Palearctic 
b = East Neartic 
c = West Nearctic 




D. mayri 


D. cuscutaeformis 


D. rubi 


D. fragariae 


D. tumefactus 


D. bassettii 


D. nebulosus 




D. fusiformans 


D. austrior 


D. kincaidii 


D. turgidus 


D. smilacis 


D. radlcum 


D. niger 


D. minimus 


a = West Palearctic 
b = East Neartic 
c = West Nearctic 




D. cuscutaeformis 


D. rubi 


D. fragariae 


D. tumefactus 


D. mayri 


D. bassettii 


D. nebulosus 




D. fusiformans 


D. austrior 


D. I<incaidii 


D. turgidus 


D. smilacis 


D. radicum 


D. niger 


D. minimus 


Fig. 7. DIVA reconstruction of historical biogeography for Diastroplnis based on tree one and tree two, 
respectively. Search options were default in DIVA. Solid arrows are unambiguous dispersal events and dashed 
arrows indicate competing alternatives of dispersal events. 


irophiis associated with woody Rithiis are 
likely to represent the first pioneers in Cy- 
nipidae in the woody world, althiiiigh lead- 
ing to a different clade from the by far more 
speciose clade comprising Diplolepidini, 
Eschatocerini, Pediaspidini, and Cynipini 
(Liljeblad and Ronquist 1998). 

Definite evidence of the early appearance 
of Rosaceae in fossil records is not known 
before Middle Eocene (ca 42-50 MYA) 
(Stewart 1983), therefore the existence of 
the genus Diastrop/iiis much earlier is un- 
likely. Land connections between North 
America and Europe probably existed until 
the Oligocene (37 Ma) (Hallam 1981). The 
presence in Beringia of temperate plants 
such as members of Potentilla was possible 
until the latest Tertiary or even Quaternary 
when glaciations cut off dispersal of tem- 
perate plants through the Bering land bridge 
(Tiffney 1985). Thus, it is difficult to spec- 
ulate by which route. North America — Eu- 
rope land connection or Beringia, earlier 
species of Diaslrophiis expanded their dis- 
tribution. The collection of Synnphronior- 
plui from Japan as discussed earlier and 
from southern China (Liu. unpubl. data) in- 
dicates that Diastrophiis is very likely to be 
present there as well. Inclusion of those 
species in future analysis certainly will pre- 
sent a clearer picture of the historical bio- 
geography of the genus. 


A grant from the Scholarship Committee 
of the Field Museum Natural History made 
it possible for KS to visit the FMNH for 
data collection. ZL was supported by a 
Boyd postdoctoral fellowship at FMNH. ZL 
was funded by the Center for Insect Sci- 
ence, University of Arizona, through NIH 
Training Grant # 1 K12 Gm00708 during 
the final revisions of the manuscript. SEM 
images presented in the paper were taken at 
the AMNH by ZL while holding a Ka- 
Iblleisch postdoctoral fellowship there. The 
DIVA program was kindly provided by F. 
Ronquist. Betty Strack provided assistance 
with the SEM at FMNH. Comments from 

R. Brusca, J. Ott, and D. Smith have greatly 
improved the manuscript. 

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354 figures. 

List of Characters 
I. Structural characters 

1 . Head width compared to mesosoma 
width: (0) not or only slightly wid- 
er, (1) distinctly wider. 

2. Apparent number of female anten- 
nal segments: (0) 13, (1) 14. 

3. Length of third antennal segment in 
relation to the fourth: (0) distinctly 
longer, (1) subequal. 

4. Shape of face in frontal view: (0) 
high, with height about four-fifths 
width of face. ( 1 ) transverse, with 


height three quarters or less oF 
width of face. 

5. Shape of gena in frontal view below 
eye: (0) straight in upper three 
fourths and distinctly curved ven- 
trally. ( 1 ) smoothly curved. 

6. Ventral projection of clypeus: (0) 
not projected, { 1 ) distinctly project- 
ed ventrally, forming broad emar- 
gination with ventral margin of ma- 
lar space, (2) strongly projected 
ventrally, tonguelike. 

7. Height of pronotal plate: (0) height 
about one-half width, ( 1 ) height at 
least two-thirds width. 

8. Notauli: (0) incomplete: (1) com- 

9. Width of notauli: (0) narrow 
throughout, (1) conspicuously wid- 
ened posteriorly, (2) wide through- 

10. Distance between notauli at tlw 
posterior margin of mesoscutum: 
(0) S3 times width of notaulus at 
posterior margin, (1) not or slightly 
more than 2 times width of notaulus 
at posterior margin. 

1 1 . Distance between anterior ends of 
notauli or signal extensions of no- 
tauli relative to that between pos- 
terior ends of notauli: (0) about two 
times as wide anteriorly as posteri- 
orly, ( 1 ) three to four times as wide 
anteriorly as posteriorly. 

12. Anterior extension of median me- 
soscutal impression: (0) absent. ( 1 ) 

13. Parascutal carinae in dorsal view: 
(0) smoothly curved, not raised la- 
teroposteriorly, (1) expanded later- 
oposteriorly into a rounded angle, 
not raised posteriorly, (2) expanded 
lateroposteriorly into an acute, dis- 
tinctly raised posterior angle. 

14. Parapsidcd signa: (0) absent. ( I ) 

15. Anteroadmedian signi on mesoscu- 
tum: (0) present, (1) absent. 

16. Posterior projection of scutelluni: 

(0) absent, scutelluni gradually 
sloped and rounded posteriorly, ( 1 ) 
scutellum distinctly projected pos- 
teriorly into an apical, broadly trun- 
cate process, (2) scutellum conspic- 
uously projected posteriorly into an 
apical, nanowly truncate process 
(Fig. 2f). 

17. Slope of posterior projection of scu- 
tellum in lateral view: (0) distinctly 
curving downward toward posterior 
apex, ( 1) almost flat. 

1 8. Extension of longitudinal linear ca- 
rinae on dorsal surface of scutel- 
lum: (0) absent completely or ab- 
sent in posterior third of scutellum, 

( 1 ) extending to end of dorsal sur- 

19. Anterior connection of longitudinal 
linear carinae on dorsal surface of 
scutellar disk: (0) absent, (1) form- 
ing concentric semicircles at the an- 
terior end of scutellar disk. 

20. Median depression of scutellum ex- 
tending posteriorly from septum: 
(0) absent, (1) only over anterior 
portion of scutellar disk, (2) 
throughout scutellar disk. 

21. Width of lateral bars: (0) nan^ow 
(less than half length), (1) wide 
(greater than half length). 

22. Presence of areolet on forewing: 
(0) absent. (1) present, but not de- 
fined by tubular vein(s), (2) present, 
defined by three distinct veins. 

23. Shape of areolet in forewing: (0) 
present, less than 4 times width of 
anal vein, ( 1 ) present, equal to or 
wider than 4 times width of anal 

24. Length of bulla in forewing: (0) ab- 
sent, ( 1 ) present, nearly as wide as 
vein width and less than two times 
vein width, (2) present, at least 
twice as wide as vein width. 

25. Size of abdomincd tergum 3 in re- 
lation to post-petiolar metasoma in 
lateral view: (0) more than half of 
metasoma. ( 1 ) about half the meta- 


soma, (2) less than one-third meta- 
somal area. 

II. Surface sculpture 

26. Coarseness of radiating strigiila- 
tion in malar space: (0) coarsely 
substrigulate, ( I ) finely strigulate. 

27. Extension of strigulation from ma- 
lar space upward to lower face: (0) 
this pattern of strigulation not con- 
tinued upward to lower face, ( 1 ) 
strigulation continued upward to 
bases of antennae, (2) pattern of 
strigulation is bent as it extends up- 

28. Sculpture on elevated median qua- 
drangular area between antennae 
and clypeus: (0) mostly glabrous, 

(1) coriaceous, (2) mostly strigu- 

29. Sculpture on surface of clypeus: (0) 
glabrous, ( 1 ) strigulate near episto- 
mal sulcus, (2) embossed, (3) with 
a spherical elevation. 

30. Sculpture of vertex and upper face: 
(0) glabrous, ( I ) coriaceous to 
slightly strigulate. 

3 1 . Sculpture of gena, especially pos- 
teriorly: (0) glabrous, (1) strigulate, 

(2) coriaceous. 

32. Upper occiput: (0) glabrous, ( 1 ) 
finely, transversely strigulate, (2) 

33. Surface sculpture of pronotal plate: 
(0) mostly glabrous with sparse 
puctures, (1) strigulate with some 
punctures. (2) mostly punctate. 

34. Pubescence on pronotal plate: (0) 
absent to sparse, ( 1 ) present in two 
dorso-lateral patches, (2) short setae 
evenly scattered. 

35. Degree and strength of diagonal 
carinae on lateral surface of pron- 
otum: (0) mostly glabrous, carinate 
only toward margins, ( 1 ) largely di- 
agonally carinate. 

36. Suiface sculpture of mesoscutum: 
(0) glabrous, (1) coriaceous. 

37. Suiface sculpture of mesopleuron: 

(0) entirely glabrous, (1) mostly 
strigulate, (2) glabrous with a few 
central striae. 

38. Surface sculpture at bases of scu- 
tellar fovea: (0) generally glabrous, 
{ 1 ) entirely strigulate-reticulate or 
rugulose, (2) some strigulation pos- 

39. Sculpture on dorsal suiface of ax- 
illa: (0) glabrous except posteriorly, 

( 1 ) rugulose, (2) coriaceous. 

III. Body color and pubescence 

40. Color of head and mesosoma: (0) 
brown to reddish brown entirely or 
in part, ( 1 ) deep dark brown to 

41. Color of metasoma: (0) brown to 
reddish brown entirely or in part, 
( 1 ) deep dark brown to black. 

42. Infuscation of membranous areas of 
forewings often appearing as a lin- 
ear streak in Rs cell: (0) absent or 
very faint, ( 1 ) present. 

43. Presence of infuscation of fore- 
wings near veins (especially Rl and 
2r): (0) absent, ( 1 ) present, but in- 
distinct, infuscate area usually less 
than vein width. (2) infuscation 
dark and distinct, usually as wide as 

IV. Gall Features 

44. Position of gall growth: (0) Above 
ground, ( 1 ) below ground. 

45. Plant tissues affected by gall 
growth: (0) stem, (1) root, (2) leaf. 

46. Gall structure effects on plant sur- 
face: (0) internal and integral, ( 1 ) 
external and detachable. 

47. Suiface of gall indicating internal 
chambers: (0) individual, { 1 ) as sin- 
gle entire group, (2) multiple galls 
in close proximity. 


105(3). 2003. pp. 733-7?,'i 




John R. Grehan and John E. Rawlins 

(JRG) BiitYalo Mu.seum of Science. 1020 Humboldt Parkway, Buffalo. NY 1421 1-1293. 
U.S.A. (e-mail:; (JER) Section of Invertebrate Zoology, Car- 
negie Mu.seum of Natural History. 4400 Forbes Avenue, Pittsburgh, PA 15213, U.S.A. 

Abstract. — We present a description of chaetotaxy and selected moiphological features 
for an unidentified Phassus larva and examine the implications for hepialoid chaetotaxy 
and biogeography. The wood-boring genera Pluissiis. Endoclita. and Aenetiis represent a 
monophyletic lineage in reference to the presence of a microtrichiated field enclosing 
SDl. SD2 and D2. Other larval characters that may support this clade include a longi- 
tudinal pit posteroventral to LI on the meso and metathorax, and a medial triangular tooth 
on the labral margin. The wood-boring Zelotypia and Cibyra may represent more distant 
relatives within a monophyletic lineage of callus feeders and wood-borers within the 
Hepialidae sensu stricto. The spatial and nomenclatural problems in Lepidoptera chaeto- 
taxy are reviewed with respect to Phassus. The term "microtrichiated pit"" is distinguished 
from "microtrichiated field" refening to an extensive, concave or flat region that may 
enclose one or more setae. We argue that slight shifts in setal position and tonosensillar 
morphology for SD2 of the prothorax is more likely than convergent development of 
tonosensillar moiphology in D2. A monophyletic relationship between Phassus. Endoclita. 
and Aenetus is biogeographically congruent with a Pacific basin origin rather than a typ- 
ically 'Gondwanic" history. We suggest that much of the biogeography and evolution of 
the Hepialidae is closely associated with Pacific geology and tectonics and this would be 
consistent with what otherwise would be an "extraordinary and inexplicable"" absence of 
Exoporia from parts of West Africa and Madagascar. 

Key Words: Hepialidae. larva, chaetotaxy. biogeography. panbiogeography. Phassus 

The Hepialidae (Lepidoptera: Suborder alus Janse, Fraiis Walker, and Gazoryctra 
Exoporia) is almost global in distribution Hiibner lack derived features of other he- 
comprising 616 described species placed in pialids and comprise a basal group of un- 
68 genera (Nielsen et al. 2000). Phyloge- certain monophyly (Nielsen and Kristensen 
netic relationships within the Hepialidae are 1989, Kristensen 1998). The remaining 
poorly understood with most studies focus- genera constitute the great majority of spe- 
ing on the establishment and composition cies and are believed to represent a mono- 
of genera and subgenera (e.g.. Tindale phyletic assemblage, the Hepialidae sensu 
1932-1942, Viette 1946-1979, Nielsen and stricto (Kristensen 1998). 
Robinson 1983, Dugdale 1994). The genera Recent studies by Nielsen and Robinson 
Afrotheora Nielsen and Scoble, AnilhepI- (1983) and Nielsen and Kristensen (1989) 


attempted preliminary phylogenetic analy- 
sis of species relationships within selected 
genera, but intergeneric relationships re- 
main generally uncertain. Wagner and Ro- 
sovsky (1991) examined the relationships 
of ten genera (nine restricted to North 
America and Eurasia) where male courtship 
behavior was known, and an unpublished 
revision by Wagner (1985) hypothesized a 
monophyletic status for Phymatopiis from 
western North America and Eurasia. A 
morphological study by Brown et al. (2000) 
presented a cladistic phylogeny for the New 
Zealand Hepialidae, and Nielsen et al. 
(2000) catalogued the entire Hepialidae 
within an informal speculative phylogenetic 

Most hepialid larvae, including all the 
basal lineages, live in soil and feed on or 
within roots, or consume leaves and other 
herbaceous debris, including mosses and 
both monocotyledonous and dicotyledon- 
ous angiosperms. Host-plant relationships 
dissociated from soil microhabitats evolved 
in the genera Aenetus Hemch-Schiiffer, Ci- 
hyra Walker (= Aepytus Herrich-Schaffer), 
EndocUta Felder, Phassus Walker, Tricho- 
phassus Le Cerf, and Zelotypia Scott where 
larvae enter the host-plant above ground 
level and tunnel into stems and branches. 
Larvae of EndocUta. Trichophassus, and 
Zelotypia are known to feed on callus tis- 
sues forming around the tunnel entrance 
(Rojas de Hernandez and Chacon de Ulloa 
1982, Grehan 1987. Grehan 1989). The pri- 
mary food source for Cibyra remains un- 
confirmed (Rojas de Hernandez and Cha- 
con de Ulloa 1982, Hilje et al. 1992). Stem 
boring is documented for the monotypic ge- 
nus Leto Hiibner (Janse 1939, Duke and 
Taylor 1964) although whether larval activ- 
ity originates within stems or is an exten- 
sion of root-feeding is unknown. Fragments 
of the host-plant with tunnels (Peabody 
Museum of Natural History) do not show 
evidence of callus feeding. 

By applying Hennig's vicariance criteri- 
on (Craw et al. 1999), Grehan (1987) pre- 
dicted that the EndocUta lineage was most 

closely related to AenetiislZelotypia through 
vicariant differentiation of a widespread an- 
cestor. Morphological characters subse- 
quently identified in support of this lineage 
include the shared presence of sub-falcate 
forewings, reduced adult antennae, a weak 
truellum in the male genitalia, a small ven- 
tral spine crest on the seventh abdominal 
segment of pupae, and a prothoracic de- 
pression (or field) enclosing setae SDl, 
SD2, and D2 in larvae (Dugdale 1994). Ab- 
sence of this trisetose feature in primitive 
Fraiis and the hepialoid sister group Mne- 
sarchaeidae led Nielsen and Kristensen 
(1989) to suggest it represented an apo- 
morphy within Hepialidae. 

Kristensen (1999) suggested male meta- 
tibial androconial scales may support a 
monophyletic lineage within Hepialidae 
sensu stricto (Table 1 ), including the callus- 
feeding genera Aenetus, EndocUta, and Ze- 
lotypia. These genera are exclusively wood- 
borers. Of the remaining androconial gen- 
era, only the Mexican-South American 
Pluis.<ius is known as a wood-borer (Grehan 
1989). Root and stem boring is reported for 
the Phymatopiis-clade of Wagner (1985), 
while larvae of Sthenopus Packard and re- 
lated Zenophassii.s Tindale feed from roots 
into stems, although neither is reported 
feeding on callus (Grehan 1989). Larvae of 
Oncopera Walker feed on leaves of grasses 
and other herbaceous perennials (Grehan 
1989), but the feeding biology of monotyp- 
ic Piiennytrans Viette is unknown (Nielsen 
and Robinson 1983). 

The specialized trisetose prothoracic field 
represents a potential larval apomorphy for 
callus-feeding wood borers. Detailed larval 
descriptions of wood-boring larvae are lim- 
ited to Aenetus cofiici Viette (Boudinot 
1991) Aenetus virescens (Herrich-Schaffer) 
(Grehan 1981. Dugdale 1994) and EndocU- 
ta hosei Tindale (Yasuda and Abe 1986). 
Larvae of other wood-boring genera are un- 
described. or have received only general- 
ized treatment. Larval descriptions of 
American wood-borers are limited to a gen- 
eral account of Trichophassus i^igajiteus Le 

VOLUME 105. NUMBER 3 735 

Table 1. Feeding modes, geographical distributions, and important morphological features of ghost moth 
genera (Hepialidae). ? = character not recorded; * = inferred by probable relationship of Zenophassus with 














Cihyrii (Aepyms) 



South America 











North America 






Southwest Pacific 








South America 


South Africa 
































Cerf by Biiquelot (1956) and brief notes on 
Phassiis triangularis Henry Edwards 
(Schaus 1888, Dyar 1917). A color photo- 
graph of an unidentified Phassiis larva from 
Ecuador by Gara and Onore (1989) indi- 
cates a shaded area corresponding to a pro- 
thoracic sensory field. In this paper we de- 
scribe the larva of an undetermined species 
of Phassiis, confirm the presence of a pro- 
thoracic field, and discuss phylogenetic and 
biogeographic implications of this character 
for the evolutionary history of wood-boring 


Specimens examined. — One dried larva 
from Jalapa, Mexico (No. 15646. Collec- 
tion of Henry Edwards, American Museum 
of Natural History ( AMNH)), and four eth- 
anol-preserved specimens (National Muse- 
um of Natural History, Smithsonian Insti- 
tution (USNM)) found boring in living 
stems of Lantana camara Linnaeus by N. 
H. L. Kraus. as follows: one from Nogales, 
Veracruz, Mexico and one from Orizaba, 
Veracruz, Mexico, both during December, 
1954: one from Orizaba, Veracruz, Mexico 
(Kraus 5053), and one from Cordoba, Ve- 
racruz, Mexico (Kraus 5055). both in No- 

vember, 1954. The chaetotaxy of Piiassiis 
sp. was compared with specimens or de- 
scriptions of about 34 species of Hepialidae 
and one species of Mnesarchaeidae (Table 

The following description is based on 
study of the above larvae that did not differ 
significantly in moiphology. Measurements 
are given only for the single, most intact 
larva (Orizaba, Mexico. December, 1954). 
All are ultimate or penultimate instars based 
on head width and body size. 

Although the specimen from Jalapa 
(AMNH) and the two from Deceinber, 1954 
(USNM) were determined as Phassiis ar- 
geiitiferiis Walker by the collectors, only 
the generic determination can be accepted 
with confidence. No reared adult material is 
associated with these larvae, but they are 
most likely Phassiis as indicated by distri- 
bution and morphology. Three genera of 
Hepialidae are known as adults from Mex- 
ico (Phassiis Walker with more than 8 spe- 
cies in Mexico, monotypic Schaiisiana 
Viette. and Cibyra Walker (sensu Nielsen et 
al. 2000), the latter containing a few Mex- 
ican species formerly placed in Hanipson- 
iella Viette, Pseiidodakica Viette, and Gy- 
meUoxes Viette. all currently treated as sub- 
genera of Cibyra. We expect larvae of the 



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Fig. 1. PhoUigaiph ot preserved last larval inslar ol HInissiis. unidentified species. 

Cihyni alliance to lack a prothoracic micro- 
trichiated field enclosing macrosetae SDl. 
SD2, and D2 based on the larval descrip- 
tion of Cihyni serta (Schaus) (Rojas de 
Hernandez and Chacon de Ulloa 1982). 

Two larvae (from Nogales (USNM) and 
Jalapa (AMNH)) were macerated in warm 
lO'/f aqueous potassium hydroxide and soft 
tissues removed. The head capsule was re- 
moved by an incision along the posterit)r 
margin. The thoracic and abdominal cuticle 
was flattened under a glass slide for ex- 
amination and subsequently stored in 7()9f 

Terminology. — At the present time the 
larval chaetotaxy of Hepialidae is a confus- 
ing patchwork of prior systems of nomen- 
clature dating from Hinton (1946) and Ger- 
asimov (1952). The terminology used in 
tills paper follows in large part the general 
practice of recent authors (e.g., Nielsen and 
Kristensen 1989. Dugdale 1994. Zilli 1998) 
and makes no attempt to resolve homology 
issues with non-exoporian taxa. The chae- 
totaxy of the head capsule used here differs 
from that of Hinton ( 1946), following with 
few exceptions the nomenclature of Has- 
enfuss (1969) as a better-supported homol- 
ogy arrangement corresponding to ditrysian 
chaetotaxy (Leonard et al. 1992). Labeling 
of prothoracic setae SDl. SD2 and D2 fol- 
lows Wagner (1987) and Wagner et al. 
(1989). For consistency, nomenclature of 
other setae and pores follow recent descrip- 
tive work on larval hepialids (Nielsen and 

Kristensen 1989) with any deviations noted 
in the text. Chaetotaxy is illustrated by a 
semischematic setal map for thoracic ab- 
dominal segments (T1-A2 and A6-AI(), 
Fig. 4-5). Lengths of setae are given in 
general terms relative to a large, precisely 
measured seta on most segments. 


Last instar (Fig. 1 ). — Exoporian. hepi- 
aloid, hepialid (Nielsen and Kristensen 
1989). Length, 56 mm; maximum head 
width. 5.84 mm; head length from epicra- 
nial notch to apex of frontoclypeus, 4.88 
mm. Head weakly hypognathous. subspher- 
ical. maximum width slightly less than pro- 
thorax but greater than other segments. 
Body elongate, parallel-sided, narrowing 
from Ab8 to AblO; Ab8 slightly gibbose 
dorsally, longer than other segments except 
Ab7; setae short, inconspicuous, set in large 
flattened pinacula or plates paler than ad- 
jacent cuticle; cuticle of body between 
sclerotized plates and pinacula densely sha- 
greened with fine microtrichiae. Prolegs 
present on Ab3 to Ab6, subequal to each 
other, smaller than prolegs on AblO. 

Color: Head dark reddish brown, body 
paler grayish red brown except for pale yel- 
low to brownish-yellow pinacula, these 
prominent as pale transverse dorsal folds on 
Abl to Ab8; prothoracic dorsal shield red- 
dish brown, edged anteriorly and ventrally 
with brownish yellow; ventral areas con- 
colorous with non-sclerotized areas on rest 


Fig. 2. Chaetotaxy of head of last instar larva of Phassus sp., Veracruz, Mexico; dorsofrontal view of head 
capsule. Symbols used: AI-A3, anterior setae: Aa, anterior pore; AFI-AF2, adfrontal setae; CI-C2, clypeal 
setae; Fl, frontal seta; Fa, frontal pore; LI, lateral seta; La, lateral pore; Lrl-Lr5, labral setae; V1-V3, vertexal 
setae; Va. vertexal pore; GI, genal seta: P1-P2, posterior setae; Pb, posterior pore; 01-02, ocular or stemmatal 
setae; Sb, stemmatal pore; S02-S03. subocular or substemmalal setae. 

of body; prolegs paler, contrasting with 
ventral abdominal coloration. Setae brown, 
spiracles black. 

Head (Figs. 2-3): Epicranium subspher- 
ical, circular in dorsal view, smooth: post- 
occipital sclerites and sutures as in other 
Hepialidae; epicranial notch obscure; coro- 
nal suture ( = epicranial suture from epicra- 
nial notch to ecdysial lines) about twice 
length of epicranial suture from ecdysial 
lines to dorsal apex of front. Front fused to 
clypeus with frontoclypeal suture obsolete, 
anterior clypeal region with a large, scler- 
otized protuberance on each side. Six stem- 
mata on each side in two dorsoventral arcs, 
anterior arc of stemmata 3, 4, and 5 (dorsal 
to ventral), posterior arc of steinmata 2, 1, 
and 6 (dorsal to ventral); stemma 5 not dis- 
placed ventrally into paramaxillary region; 
distinct pore anteroventrad of stemma 4. 
Antennal fossa closed ventrally by pre-an- 
tennal bar that articulates with epicranium 
anterior to antenna and is posterior (not 

contiguous) to dorsal mandibular articula- 
tion. Antennal slit (sensu Dugdale 1994) a 
narrow strip of membranous cuticle contin- 
uous with that between antenna and base of 
mandible, extending posterodorsad and 
ending near two distinct pores just ventral 
to stemma 5. Antenna not studied. 

Setation of head as for other Hepialidae 
(Nielsen and Kristensen 1989). Dugdale 
(1994) following nomenclature of Hasen- 
fuss ( 1969) although homology of that sys- 
tem with ditrysian nomenclature (e.g., Stehr 
1987) doubtful. Seta VI macrosetose, setae 
V2 and V3 microsetose; pore present near 
seta PI (termed Pb by Nielsen and Kristen- 
sen 1989); pore La present; pore medial to 
seta SOI visible only in ventral view (per- 
haps homologous with pore SSa of ditry- 
sian system); pores Sa (= Oa of Hasenfuss 
1969), MGa, AFa, and Aa absent; two ge- 
nal microsetae; seta S03 minute, microse- 
tose (not visible or depicted in Fig. 3). 

Labium with five pairs of setae; anterior 



margin with medial, broadly triangular 
tooth. Maxillolabial complex with basisti- 
pes and dististipes sclerotized. Maxillary 
palp three-.segmented with distal segment 
subequal in size to large sensillum basicon- 
icum as in other Hepialidae; lateral pore on 
basal segment of palp. Medial maxillary 
lobe laterally sclerotized with seven sensilla 
on distal surface: three large flattened sen- 
silla dorsally, middle row of two apically 
rounded sensilla, lower row of two pointed 
sensilla with pitlike structure between them. 
Basistipes with two setae and a pore. Dis- 
tistipes with single lateral seta. Labial palp 
minute with long apical seta, arising from 
lateral subapical plates of premental lobe 
(maxillary features similar to those illus- 
trated for Fraus (Nielsen and Kristensen 
1989)). Mandible with four triangular teeth 
on distal extremity; oral surface of mandi- 
ble transversely rugulose without distinct 
molar area; mandible with two setae on ab- 
oral surface, basal seta Ml, distal M2. 

Prothora.x (Fig. 4): Entire dorsal surface 
of prothorax sclerotized, anterior margin re- 
llexed to join posterior margin of head cap- 
sule; prothoracic dorsal shield (sensu stric- 
tt)) thicker and indistinctly delimited from 
sclerotized marginal regions, yellowish 
brown; sclerotization of dorsal shield in- 
cludes L-group setae, naiTowly separated 
from sclerotized region around spiracular 

Seta D 1 strong, posterior to anterior mar- 
gin of dorsal shield at a distance subecjual 
tt) length of seta (0.76 mm); seta XDI di- 
rectly ventral to seta Dl, more than twice 
as long. Seta XD2 slightly anterior to XDI, 
subequal in length to Dl; seta D2 slightly 
ventral to level of XD2. slightly smaller 
than Dl, approximately midway between 
SDl and SD2. Three prothoracic pores 
(sensu Nielsen and Kristensen 1989): XDa 
directly ventral to Dl, pigmented dark 
brown; XDb posterodorsal to XDI, pale; 
XDc posterodorsal to XD2, strongly pig- 
mented; diaineter of all pores slightly less 
than socket of associated setae. 

Seta D2 ventral and slightly anterior to 

seta SD2, dorsal and slightly posterior to 
seta SDl; length of seta D2 0..'^9 mm; 
length of SDl 0.42 mm, equal to SD2. Both 
SD setae extremely slender, hiiform, not at- 
tenuate, arising from bottom of distinct con- 
ical pits with strongly microtrichiated walls 
(similar to tonosensilla of ditrysian larvae); 
cuticular articulation of D2 unmodified with 
setal alveolus (socket) sutix>unding visible, 
pale setal membrane bearing base of seta. 
Seta D2, both SD setae, and associated pits 
included within broad region (field) de- 
pressed below sun'ounding cuticle (Fig. 6), 
continuously microtrichiated (shagreened) 
more densely than adjacent cuticle or else- 
where on body; basal pits of setae SDl and 
SD2, and base of seta D2, darkly pigment- 
ed, contrasting with microtrichiated held; 
maximum dorsoventral dimension of mi- 
crotrichiated field 1.45 mm, width 0.55 
mm. Small unnamed pore on extreme dor- 
sal edge of microtrichiated field dorsal to a 
line between setae Dl and SD2, diameter 
smaller than alveolus of seta D2, but greater 
than diameter of hyaline setal membrane at 
bases of SDl and SD2. 

Seta LI near anterior edge of prothoracic 
shield directly anterior to middle of spira- 
cle, subequal in length to seta XDI. Seta 
L2 anterodorsal to seta LI, approximately 
half the length of seta LI. Seta L3 directly 
anterior to middle of spiracle, displaced 
from anterior peritreme by less than hori- 
zontal diameter of spiracle, shortest seta on 
prothoracic shield (half the length of seta 
L2). Setae SVl and SV2 below L-group se- 
tae on pinaculum narrowly separated from 
ventral edge of prothoracic shield; SVl 
subequal in length to XD2 or Dl, directly 
posterior to seta SV2 and twice its basal 
diameter; SV pinaculum yellowish tan, con- 
colorous with ventral lobe of prothoracic 

Seta VI subequal in size to seta L3, pos- 
terior to prothoraxic coxae on sclerotized 
mid-ventral plate. Seta MV3 macrosetose, 
subequal in size to seta L2, directly anterior 
to coxa; seta VI on large plate which cross- 
es ventral midline; seta MV2 macrosetose. 


Fig. 3. Chaetotaxy of head of last inslar larva of Pluissii.s sp.. Veracruz. Mexico; lateral view of head capsule. 
Symbols used as in Fig. 2. 

siibeqiial in size lo seta V 1 , anterior to mid- 
dle of prothoracic coxa on extreme anterior 
edge of cuticular fold such that seta is ap- 
pressed to head capsule midway between 
cranial setae Gl and G2. 

Spiracle vertically ovate, 0.89 mm high. 
0.35 mm long; outer peritreme heavily 
sclerotized, darkly pigmented; inner (filter) 
recessed into atrium approximately 0.10 

Prothonicic leg (Figs. 7-8): Prothoracic 
coxae proximate at base across ventral mid- 
line. Coxa with eight setae: anterodorsal 
pair (Cxi, Cx2) very small, subequal in 
size, near proximal edge of coxa with Cxi 
dorsalmost; anteroventral setal pair (Cx3, 
Cx4) unequal in size with anterior Cx3 as 
small as Cxi and posterior Cx4 longer and 
thicker; posteroventral setal pair (Cx5, Cx6) 
with Cx5 longest and most ventral; Cx6 
near middle of posteroventral expanse of 
coxa, slightly shorter than Cx5; posterodor- 

sal pair (Cx7, Cx8) subequal to Cxi; Cx7 
near proximal edge of coxa and dorsalmost; 
Cx8 near upper middle of posterodorsal 
swelling of coxa. Trochanteral seta Trl mi- 
crosetose. in extreme dorsal portion of tro- 
chanterofemoral membrane on anterior sur- 
face of leg; trochanteral pore Tra close by 
[second trochanteral pore Trb of Nielsen 
and Kristensen (1989) not evident]; tro- 
chanteral seta Tr2 microsetose, in trochan- 
teral membrane at ventral (adaxial) edge of 
leg; trochanteral seta Tr3 microsetose, in 
extreme dorsal portion of trochanterofe- 
moral membrane on posterior surface of 
leg. Femoral seta Fel the largest and lon- 
gest seta on legs, midlength on ventral edge 
of femur; femoral seta Fe2 near distal pos- 
teroventral edge of femur. Six tibial setae 
[nomenclature as Nielsen and Kristensen 
(1989)], all in distal half of tibia; Til, Ti3, 
and Ti5 on anterior surface of tibia, dorsal, 
subventral. and ventral respectively; Ti2, 



Ti4, and Ti6 on posterior surface of tibia, 
dorsal, subventral, and ventral respectively; 
tibial pore Tia conspicuous, in posterodor- 
sal surface of tibia near midlength. Four tar- 
sal setae; dorsal pair near distal end of tar- 
sus. Tal anterior, Ta2 posterior; ventral pair 
unmodified in shape, on ventral edge of tar- 
sus. Ta3 at distal extremity. Ta4 directly 
basal to Ta3 near midlength of tarsus, short- 
er and thinner than Ta4. Tarsal claw 
smoothly tapered with slight ventral im- 
pression near base; without teeth or other 

Mcsotlu'ia.x (Fig. 4): Two transverse 
tlorsal shields continuously fused across 
dorsal midline; anterior mesothoracic shield 
bearing seta Dl (0.76 mm in length, sube- 
cjual in size to Dl on prothorax). extending 
ventrad to level of seta XD2 on prothorax; 
Dl directly posterior to seta XDl of pro- 
thorax. Posterior mesothoracic shield with 
seta D2 nearly twice the size of seta Dl 
(length 1.37 mm. subequal to seta D2 on 
prothorax). directly ventral to seta SD2. 
posterior to Dl. and directly posterior to 
prothoracic seta SDl; posterior mesothorac- 
ic shield pale yellow with seta L3 in pos- 
teroventral corner; L3 very small, subequal 
to L3 on prothorax. Seta LI on anterior end 
of distinct lateral shield; longitudinal pit 
posteroventral to LI. brown, sclerotized. 
Seta L2 subequal in size to L3. on fold di- 
rectly below ventral extremity of anterior 
mesothoracic shield and directly anterior to 
SDl. Setae MDl. MSDl. and MSD2 lo- 
cated on surface of a single lateral fold 
which is occluded from view by posterior 
lobe of prothorax and bulging anterior 
shield of mesothorax (i.e., concealed in 
groove between prothoracic shield and me- 
sothoracic shield). MDl a microseta on out- 
er edge of fold, directly anterior to seta 
SD2. with apex in space between thoracic 
shield and adjacent mesothoracic shield: 
seta MSDl a microseta on anterior slope of 
fold, slightly ventrad of seta MDl, with 
apex in space between microsetal fold and 
prothoracic shield; seta MSD2 directly pos- 
terior to seta MDl on posterior declivity of 

fold, with apex in space between microsetal 
fold and anterior mesothoracic shield. Setae 
MSDl and MSD2 subequal in size, slightly 
smaller than seta MDl. Single SV seta in 
middle of large subventral, pale yellow 
shield; SV subequal or larger in size than 
setae Dl and SD2. but smaller than setae 
D2 and SVl. Two transverse plicae be- 
tween mesothoracic coxae and posterior 
margin of prothorax; anterior plica very 
small, not visible in external ventral view, 
and bearing seta MVl on its posterior face; 
posterior plica larger, visible in ventral 
view, bearing seta MV2 posterodorsal to 
seta MV3 on anterior declivity; MV2 di- 
rectly opposite dorsal extremity of anterior 
margin of mesocoxa with apex extending 
forward and contacting posterior ventral re- 
gion of prothorax. Setae MVl, MV2 and 
MDl subequal in length; setae MSDl, 
MSD2, and MV3 subequal in length, slight- 
ly shorter than setae MVl, MV2, and MDl. 
Mesothoracic coxae separate at base. Sub- 
dorsal peg organ located in membrane on 
extreme anterior edge of posterior mesotho- 
racic shield, opposite ventral posterior ex- 
tremity of anterior mesothoracic shield, 
dorsoventrally midway between level of se- 
tae D2 and SD2. Mesothoracic leg as for 
prothorax, in general all setae slightly 
smaller, especially anterior coxal setae. 

hietathorax (Fig. 4): As for mesothorax 
with the following differences; peg organ 
more exposed on leading edge of posterior 
metathoracic shield; anterior metathoracic 
shield slightly longer on midline than that 
shield on mesothorax; posterior metathorac- 
ic shield slightly shorter on midline than 
that shield on mesothorax; leg as on me- 

Ahdoiiifii (Figs. 4-5): Generalized ab- 
dominal segment consisting of four annuli: 
first (anteriormost) annulus small, nearly 
hidden from view between second annulus 
and posterior edge of preceding segment, 
diminishing laterally to simple plica; sec- 
ond annulus largest, bearing seta Dl; third 
annulus smaller than second, bearini: seta 


SV2 SV1 


Figs. 4-5. Chaetotaxy of thoracic and abdominal segments of Phassus sp. (semischematic views). 4. Left 
side of thorax and first two abdominal segments. 5. Left side of abdominal segments 6-10. Symbols used: Abl- 
2. Ab6-10. abdominal segments I. II. and VI through X; D1-D2, dorsal setae; L1-L3. lateral setae; LAa, lateral 
pore of AblO; MDI. dorsal proprioceptor seta; MSDI-MSD2. subdorsal proprioceptor setae; MVI-MV3. ventral 
proprioceptor setae; PPI. paraproct seta; SDI-SD2. subdorsal setae; SVI-SV3, subventral setae; TI-T3. thoracic 
segments I to III; VI, ventral seta; XD1-XD2, anterior dorsal setae of Tl; XDa-XDc, anterodorsal pores of Tl. 

D2; fourth annulus not sclerotized dorsally. 
subequal in size to third annulus. 

Generalized setation of uhdoiniiuil seg- 
ments: Setae Dl and D2 subequal in size, 
similar in position to same setae on T3, D2 
ventral to Dl. Seta SD2 in center of large 
pale pinaculum. subequal in size to seta Dl, 
about half size of seta SDl on anterodorsal 
corner of same pinaculuin. Peg organ con- 
spicuous, dark, located in membrane mid- 
way between ventral side of plate Dl and 

dorsal side of plate bearing setae SDl and 

Spiracle on first abdominal segment 
(Abl) largest, subequal to spiracle on Tl 
(height 0.92 mm, length 0.42 mm), that on 
Ab8 smaller (height 0.81 mm, length 0.42 
mm), that on A2 smaller still (height 0.78 
mm, length 0.42 mm) and those on A3-A7 
subequal and smallest (height 0.69 mm, 
length 0.42 mm). 

Abdominal segment 1: Seta MDI ante- 



unnamed pore 




Fig. 6. Detail of microtrichiated field of larval pro- 
thorax, Phassus sp. SymboLs used: D2. dor.sal .seta; L3, 
lateral seta; SD1-SD2. subdorsal setae. 

rior to peg organ, at bottom of groove be- 
tween first and second annulus. Cuticle en- 
folded around seta MDl forms open sub- 
spherical cavity, not collapsed by move- 
ment. Pinaculum of seta L2 directly 
posterior to spiracle; pinaculum of seta LI 
dorsoposterior to pinaculum L2. Seta L3 
slightly anterior to spiracle on extreme an- 
terior edge of large longitudinal plate ex- 
tending posterior to intersegmental groove. 
Setae SVl, SV3 located on dorsal half of 
same pale pinaculum. 

Abdominal segment 2: As for Abl with 
following differences: Seta MDl located 
near bottom of grove between annulus 1 of 
Ab2 and annulus 4 of segment Abl; large 
oval plate without setae on annulus 1 at lev- 
el of spiracle, subequal in size to spiracle; 
seta SD2 on small pinaculum approximate 
and dorsal to larger pinaculum bearing 
SDl; seta LI located on extreme dorsopos- 
terior edge of plate bearing L2, not on sep- 
arate plate; seta MV relatively large, more 

than twice length of seta MV on segment 

Abdominals segments 3—6: As for seg- 
ment Ab2 with following differences: seta 
SD2 on distinct pinaculum on segments 
Ab3— Ab4, that pinaculum reduced or ab- 
sent on Ab5-Ab7; seta LI on distinct small 
pinaculum posterodorsal to larger plate 
bearing seta L2; seta SVl largest, most pos- 
terior, and dorsalmost of any SV seta; seta 
SV2 anterovential to seta SVl on same 
plate, half the length of SVL seta SV3 an- 
teroventral to SV2 on same plate, about half 
length of seta SVl; seta VI directly be- 
tween base of prolegs on same sclerotized 
plate encircling pioleg and bearing all SV 
setae; seta MV3 directly anterior to proleg 
on extreme anterior edge of sclerite bearing 
setae VI and SV1-SV3. Crochet confor- 
mation as uniordinal biserial ellipse, outer 
series with crochets greatly reduced in 

Abdominal segment 7: As for segment 
Ab6 with following differences: setae SVl 
and SV2 located on same plate, half the di- 
ameter of homologous plate on Ab2; seta 
SV3 absent; seta VI directly ventral to seta 
SV2 on very small sclerotized plate; seta 
M V3 directly anterior of seta V 1 , subequal 
in size to seta MV3 on other abdominal 

Abdominal segment 8: As for segment 
Ab7 with following differences: spiracle 
subequal in size to that on segment Ab2; 
seta MDl exposed in flat region anterior to 
annulus 1; seta SDl on very small basal 
plate, SD2 anterodorsal to SDl, without 
sclerotized basal pinaculum; no prespira- 
cular plate on annulus 1; seta LI lacks pin- 
aculum; plate around seta L2 smaller than 
adjacent spiracle; seta L3 located on small 
pinaculum anterior to and slightly separated 
from larger lateral plate without setae; setae 
SVl and SV2 very close on shared basal 
pinaculum; seta VI on small distinct plate; 
seta MV3 much smaller than same seta on 
segments Ab2-Ab7, near outer edge of in- 
tersegmental groove, halfway between SVl 
and VI. 


Figs. 7-8. Chaetotaxy of prothoracic leg of last instar larva (if Pluissiis sp. 7, Anterior view of left prothoracic 
leg. 8, Posterior view of left prothoracic leg. Symbols used: Cxl-Cx8, coxal setae; FEI-FE2, femoral setae; 
MTRa. anterodorsal pore of trochanter; Tal-Ta4, tarsal setae; Til-Ti6, tibial setae; Tia, posterodorsal pore of 
tibia; Trl-Tr3, proprioceptor seta of trochanter. 

Abdominal segment 9: As for segment 
Ab8 with following differences: only a sin- 
gle obvious annulus; all setae with reduced 
basal pinacula; Dl setae much closer to 
each other across midline than are setae D2 
across midline; SDl directly ventral to Dl; 
seta SD2 missing; MDl anterior to SDl, 
extending into intersegmental groove on 
anterior slope of segment; setae LI, L2, and 
L3 in vertical row, equidistant, directly ven- 
tral to D2; setae SVl and SV2 as on seg- 

ment Ab8, directly ventral to L3 but not on 
shared basal pinaculum; seta VI strong, 
near ventral midline; seta MV3 slightly 
ventral to SV2, enfolded within interseg- 
mental groove. 

Abdominal segment 10: Three setae (Dl, 
D2, SDl) on dorsal anal shield; two setae 
(here interpreted as seta LI (anterior) and 
paraproct seta PPl (posterior)) and one pore 
(LAa) on posterior plate; two setae below 
anus and dorsal to proleg (interpreted as se- 


tae L2 (anterior) and L3 (posterior)); two 
lateral setae anterior to proleg (interpreted 
as setae SVl (dorsal) and SV2 (ventral)); 
two ventral setae anterior to proleg (inter- 
preted as seta VI (posterior near edge of 
planta) and nearly macrosetose MV3 (near 
intersegmental groove with segment Ab9). 
Crochet conformation triserial. uniordinal, 
arranged as two semicircular loops on each 
anal proleg. 

Problems with Homology and 
Setal Nomenclature 

The multiplicity of chaetotaxic systems 
for lepidopterous larvae, and in particular 
for hepialids, makes it difhcult to recognize 
and name setae that are homologous at the 
ordinal level. Resolution of this problem is 
not possible in this paper and awaits more 
extensive research on setation in Exoporia, 
other basal clades of Lepidoptera, and Di- 
trysia. As an initial set of concerns for fu- 
ture research in this area, we provide an an- 
notated list of the setal groups found to be 
problematic in this study. 

1. Thoracic and abdominal setae of the 
MV and V groups of Hinton (1946). Con- 
fusion arises from variation in stereographic 
position of setae and their relative size and 

2. Setae of the SV group, especially on 
the abdomen. This is a classic dilemma in 
many ditrysian superfamilies, especially for 
variation on the first two abdominal seg- 
ments, and segments where prolegs may be 

3. Setation of abdominal segment 10. 
This segment consists of problems of both 
number and position of setae on the anal 
shield (D and SD groups) and of presence 
and placement of setae on more ventral por- 
tions of the segment (L, SV, and V groups). 

4. MD and MSD microsetae on thoracic 
segments. The problem here may be due to 
inadvertent but alternate nomenclature ap- 
plied by Hardy ( 1973) and Yasuda and Abe 
(1986). We concur with Nielsen and Kris- 
tensen (1989). 

5. Head setae. Many problems with al- 

most every setal group on the head result 
from confusion about variable placement 
and size of setae. Although the chaetotaxy 
is not altered, a more thorough discussion 
of this problem is given below. 

6. D, SD, and MXD group setae and as- 
sociated pores on the prothorax. These are 
variable features unique to Exoporia. Pro- 
thoracic D and SD setae are discussed in 
detail below. 

Setation of the head. — Investigators us- 
ing setae in the comparative study of Lep- 
idoptera larvae have usually followed prior 
convention in chaetotaxy. Students of he- 
pialid larvae have revealed consistent dif- 
ferences in setal placement with ditrysian 
larvae, and these have resulted in a number 
of nomenclatural systems, each differing 
slightly from the others. This is especially 
true for setae of the head where a system 
dating from Heinrich (1916), Gerasimov 
(1935) and Hinton (1946) was modified by 
Hasenfuss (1969) for Hepialidae and by 
Stehr (1987) for Ditrysia, then variously 
modified again for Hepialidae by Wagner 
( 1987), Wagner et al. ( 1989), Leonard et al. 
(1992), and Nielsen and Kristensen ( 1989). 
To avoid confusion other workers have 
wisely followed the nomenclature of recent 
authors when comparing setae within He- 
pialidae (Dugdale 1994, Zilli 1998). No 
worker since Hasenfuss has proposed and 
adequately defended a chaetotaxic system 
that seeks to recognize homology across the 
entire order, as such an undertaking would 
require detailed study of all world lineages. 

For some groups of cranial setae there is 
no controversy among published descrip- 
tions in the last fifty years. All workers 
agree on the naming of setae on the frontal 
and adfrontal sclerites, and concur that pore 
AFa is absent in hepialids. They further 
agree that there are two mandibular setae, 
but do not distinguish between them (the 
basal seta is here named Ml, the distal seta 
M2). All workers agree on the number and 
placement of L-group setae, and of SS- 
group setae (sometimes labeled S01-S03 


(subocellar setae) as opposed to SSI-SS3 
(siibstemmatal setae)). 

For other groups of cranial setae there are 
major dift'erences in opinion as to nomen- 
clature. These differences may be clarified 
by noting how various authors have treated 
each majcir group of setae. Treatment of 
clypeal setae is divided into two groups: 1 ) 
those naming the lateral setae CI (Hinton 
1946. Hasenfuss 1969, Nielsen and Kristen- 
sen 1989, Dugdale 1994), and 2) those 
naming the medial setae CI (Wagner 1987) 
as in Ditrysia (Stehr 1987). 

Treatment of labral setae is ignored by 
all workers, but all concur in their drawings 
that hepialids have five pairs of externally 
visible labral setae. We here call these 
LRl-LR.S with a single pore, LRa, follow- 
ing the nomenclature of Rawlins (1992) as 
applied to ditrysian larvae. The homology 
of the lateral labral seta, LR5, with the di- 
trysian condition is uncertain as there are 
two lateral setae on each side in that line- 
age, LR5 and LR6. 

Numbering of stemmata in hepialids is 
not provided by any of these workers. The 
system followed here is consistent with that 
used for Ditrysia. assigning numbers for the 
left side of the head from 1 to 6 in a coun- 
ter-clockwise direction such that the most 
\entral stemnia near the posterior edge of 
the antennal fossa is 5. 

Treatment of S-group setae is divided 
into three groups: 1 ) those following the di- 
trysian system (Stehr 1987) and recogniz- 
ing three S-group setae with SI inside the 
stemmatal circle and S3 microsetose (Has- 
enfuss 1969), 2) those following the ditry- 
sian system but considering S3 to be absent, 
replaced by seta G2 or MG2 (Nielsen and 
Kj-istensen 1989, Dugdale 1994), and 3) 
those adopting a system with SI anterior to 
the stemmatal region and adjacent to the an- 
terior mandibular condyle. S2 within the 
stemmatal field, and S3 immediately pos- 
terior to the stemmata (Hinton 1946, Wag- 
ner 1987). 

Treatment of the genal setae may be di- 
vided into two groups: 1 ) those using the 

ditrysian system (Stehr 1987) with a single 
genal seta and pore, MGl and MGa (Has- 
enfuss 1969), and 2) those recognizing two 
genal setae and a pore, Gl, Ga, and G2 (or 
MGl, MGa, and MG2) (Hinton 1946, Wag- 
ner 1987, Nielsen and Kristensen 1989. 
Dugdale 1994). 

Treatment of the dorsal setae is also di- 
vided into two groups: 1 ) as in Ditrysia 
(Stehr 1987) with three dorsal setae and a 
pore (MD1-MD3 and MDa, or VI -V3 and 
Va) of which MDl or VI is macrosetose 
(Hasenfuss 1969, Nielsen and Kristensen 
1989, Dugdale 1994), and 2) those recog- 
nizing two dorsal setae and a pore, all mi- 
crosetose (MD2-MD3 and MDa. or V2-V3 
and Va) (Hinton 1946, Wagner 1987). 

All workers agree on naming PI setae, 
but treatment of P2 setae is divided into 
three groups: 1 ) the ditrysian condition with 
P2 posterodorsal to PI and associated with 
pore Pb (Stehr 1987), 2) the ditrysian con- 
dition with P2 posterodorsal to PI but not 
associated with a pore Pb (Hinton 1946, 
Wagner 1987), and 3) with P2 lateral or la- 
teroventral to PI and associated with a pore 
Pb (Hasenfuss 1969, Nielsen and Kristen- 
.sen 1989, Dugdale 1994). 

All past treatments agree on recognition 
of seta A2, and the absence of pores Aa and 
Pa. Treatment of other A-group setae is 
controversial and may be divided into two 
groups: 1 ) those naming the anteriormost 
seta near the anterior articulation of the 
mandible as Al with seta A3 near stemma 
2 (the ditrysian system (Stehr 1987) fol- 
lowed for Hepialidae (Hasenfuss 1969, 
Nielsen and Kristensen 1989, Dugdale 
1994)), and 2) those naming the seta asso- 
ciated with stemma 2 as Al, and that as- 
sociated with pore Pb as A3 (Hinton 1946, 
Wagner 1987). 

Careful study of the bewildering situation 
above reveals that the fundamental conflict 
for determining setal homologies is be- 
tween the size of setae (microsetose versus 
macrosetose) and their stereographic place- 
ment relative to each other and to pores. If 
the size of setae is ignored, then cranial se- 


tae of Ditrysia and Hepialidae are spatially 
and numerically consistent with one excep- 
tion: seta P2 in hepialids is displaced ven- 
trally and laterally from the expected posi- 
tion in Ditrysia. This anangement requires 
recognizing the ventral genal microseta in 
hepialids (MG2 of authors) as homologous 
with the posterior macroseta S3 of Ditrysia 
and treating the posterior dorsal macroseta 
of hepialids as homologous with the ante- 
rior dorsal microseta of Ditrysia (MDl of 
authors). Under this system there is no need 
to vii)late consistent spatial associations of 
A-group or S-group setae as proposed by 
Hinton (1946) and Wagner (1987). 

If the microsetose or macrosetose con- 
dition of setae is hypothesized to be so im- 
portant that setae of different sizes cannot 
be considered homologous, then a diversity 
of ad hoc hypotheses on setal homology are 
required to account for all setae. This re- 
quires switching the nomenclature of seta 
A I and SI, thereby changing their stereo- 
graphic placement on the head capsule rel- 
ative to the mandible and stemmata, in- 
creasing the number of genal microsetae 
from I to 2, decreasing the number of dor- 
sal microsetae from 3 to 2. hypothesizing 
the complete disappearance of macroseta 
S3, associating seta A3 with pore Pb, or 
removing seta P2 from association with that 
pore, and so on. 

Given that both ditrysian and exoporian 
larvae possess the same number of primary 
cranial setae in very similar spatial relation- 
ship to each other and to cranial landmarks 
such as the antennal fossa, mandibular con- 
dyles, adfrontal sclerites, and stemmata, a 
parsimonious hypothesis of homology for 
these setae involves accepting major chang- 
es in setal size and a lateroventral shift in 
the position of seta P2. A chaetotaxic sys- 
tem conesponding to such homology re- 
quires application of names for macrosetae 
to microsetae (the hepialid seta MG2 be- 
comes S3) or vice versa (the hepialid ma- 
croseta VI become MDl). In retrospect it 
is unfortunate that a special nomenclature 
fo]- microsetae arose following Hinton 

(1946) as this may have obscured major 
evolutionary shifts in setal size and func- 
tit)n. and in any event has greatly compli- 
cated hypotheses of homology with an al- 
ready abstruse chaetotaxy. This paper is not 
the place to present a testable, homologous 
system of nomenclature for setae on larval 
Lepidoptera or other holometabolous lar- 
vae, but the above discussion should un- 
derscore the need to do so in order to more 
clearly understand the evolution of setal 
size, placement, and function. 

Microtrichiated pits and Helds. — Previ- 
ous authors have used a variety of terms tt) 
describe regions of microtrichiated cuticle 
surrounding the base of setae in larval Lep- 
idoptera, including "pocket"" (Rawlins 
1984), "pigmented sensory pit"" (Wagner 
1987), "microtrichiated pit"" (Nielsen and 
Kristensen 1989), "pigmented pit with mi- 
crotrichiated walls"" (Rawlins 1992), "mi- 
crotrichial bed" (Leonard et al. 1992), and 
"felted pits"" (Dugdale 1994). These terms 
confuse two different ciiticLilar features, 
both distinguished by the presence of mi- 
crotrichiae: I ) a relatively small, deeply im- 
pressed pit surrounding the base of a single 
seta, and 2) a more extensive, concave or 
flat region that may enclose one or more 
setae. We limit the expression "microtri- 
chiated pit"" to the former condition, almost 
always in association with Hliform tonosen- 
silla. and use the expression "microtrichiat- 
ed field"" to describe the latter. Microtri- 
chiated pits and fields often occur indepen- 
dently, but pits can also be located within 
fields as in Phassiix. and may be develop- 
mentally related, differing only in degree of 

Prothoracic setae. — A distinctive, and 
possibly apomorphic feature of some hepi- 
alid larvae is the presence of three setae 
(D2, SDL and SD2) enclosed by a single 
continuously microtrichiated field, as op- 
posed to having the field divided into two 
separate regions, each enclosing a seta. Be- 
cause these setae are ventrally displaced 
from the position D2 occupies in Ditrysia. 
there has been confusion over their homol- 


ogy relative to other thoracic segments and 
prothoracic setae in other Lepidoptera. The 
problem is clarified but not rcsoheti in the 
following paragraphs. 

Criteria most often used to determine the 
homology of setae on a single larva may be 
broadly grouped into two categories: (';•/- 
terioii 1, stereographic position relative to 
other setae with respect to body axes (dor- 
sal, ventral, anterior, posterior), and Crite- 
rion 2. morphological details of the seta it- 
self (si/e, shape, color, surface microsculp- 
lure, and others) including region of artic- 
ulation with adjacent cuticle. Analysis of 
homology for setae results from compara- 
tive study of their position and morphology 
between developmental instars and between 
larvae of different taxa. Ontogenetic com- 
parisons are prt)blematic, especially those 
involving first instars. as there is no a priori 
reason to believe that apomorphic features 
could not have evolved in the first instar. 
Ni) first instar I'luissiis larvae were avail- 
able for study. 

To determine the homology of setae D2, 
SDI. and SD2 undei' the above criteria, we 
consider first the situation tor each coiuli- 
tion in Ditrysia. 

Seta D2 C'riicrion 1: D2 tlorsal and pos- 
terior to SDI and SD2 (or al 
most with D2 directly dorsal to 
those setae). 

Criterion 2: D2 variable in 
length, often shorter than SD2 
antl/or subet|ual in length to 
SDI; never positioned in a mi- 
crotrichiated pit on any segment, 
and always a typical macrosela. 
never a filiform tonosensillimi. 

Seta SDI Criterion I: SDI anterior and 
ventral to SD2, at most directly 
anterior or directly ventral to 
SD2 (ne\er posterior or dorsal to 

Criterion 2: Si3l shoiter than 
SD2 in many lineages, but in 
some subequal or greater in 
length than SD2; positioned in a 

microtrichiated pit in some line- 
ages; a hliform tonosensillum in 
some groups. 
Seta SD2 Criterion 1 : as abt)ve. 

Criterion 2: as above, not asso- 
ciated with microtrichiated pit 
and not a filiform tonosensillum. 

Contrast the above pattern to that observed 
in Pluissus using the terminology of Wag- 
ner (I W7): 

Criterion I. — D2 is ventral ami anterior to 
SD2, dorsal and posterior \o SDI. Con- 
clusion: Placciucnt of both D2 and SD2 
violalcs slcrcoi^rnphic coiulitious hol/i 
dorscil-vcntrol duel onlcrior-iuislcrior. 
Seta SDI, however, is in accordance with 
Criterion 1 in the Ditrysia. 

Criterion 2. — D2 slightly longer than SDI 
and SD2 which arc subequal in length; 
D2 a strong, typical tactile macroseta, but 
SDI and S132 are filiform tonosensilla 
positioned in microtrichiated pits; all 
three setae and jiits within a iiiicrolri- 
chiated field. Conclusion: D2 in accor- 
dance with Criterion 2 (large macroseta 
without a microtrichiated pit); SDI in ac- 
cordance (tonosensillum with a microtri- 
chiateil pit); SD2 not in ai^rccnicnt with 
Criterion 2 In-iiii; a lonosensilliini in a 
iiiierotrieliidted pit. 

Sv\itching names for setae D2 and SD2 
conforms to the nomenclature of Nielsen 
and Kristensen (1989) produces the follow- 
ing situation luider Criteria I and 2. 

Criterion I. — All setae are in accord with 
the condition in Ditrysia. 

Criterion 2. — D2 subequal in length to SDI 
and shorter than SD2; SD2 a strong, typ- 
ical tactile macroseta, but SDI and D2 
are filiform tonosensilla positioned in mi- 
crotrichiated pits; all three setae and pits 
within a microtrichiated field. Conclu- 
sion: SD2 and SDI in accordance with 
Criterion 2. hut 1)2 not in a)>reeineiu with 
Criterion 2 heiim a tonosensilhmi in ti 
niierotrichidled pit. 

The abo\e analvsis of setal condition in 

VOl.UMH :()5. Nl'MKHK ,^ 

/'lhi.s.\ii\ reveals a conflict wilh both Crile- 
lion I and Criterion 2. Resolution requires 
weighting one over the other. In this case, 
weighting Criterion 2 over Criterion I re- 
iiuires hypothesizing convergent develop- 
nieni of complex morphological leatmes 
(tonosensillum in a microtrichiated pit) for 
either seta SD2 (Wagner 1987) or seta D2 
(Nielsen and Kristensen 1989). Weighting 
Criterion 1 over Criterion 2 requires hy- 
pothesizing shifts in spatial placement lor 
setae D2 and SD2 under Wagner's termi- 
nology, hut not iMidei' liiat of Nielsen anil 
Krislensen (1989). It is Icnipting lo con- 
clude that the most parsimonious solution 
would be the latter system, but this ret|uires 
ilevelopinent of D2 as a tonosensillum in a 
microtrichiated pit, a situation not encoun- 
tered elsewhere in Lepidoptera. The alter- 
native system (Wagner 1987) requires rel- 
atively slight shifts in slereographic posi- 
tion for two setae and development of SD2 
as a tonosensillum, a condition we feel 
more likely than for D2 insofar as tonosen- 
silla are usually SI) group setae in olhci 

It is important to realize that a testable 
determination of which criterit)n to empha- 
size is not possible withoiU further morpho- 
logical and comparative study. For the tinie 
being we prefer to accept slight shifts in 
setal position and lonosensillar morphology 
for .SD2 as more likely than convergent de- 
vek)pmenl of tonosensillar morphology in 
D2. Favoring a chaetotaxic system empha- 
sizing the greatest likelihood of homology, 
we have used the terminology of Wagner 
( 1987) in agreement with the logic of Dug- 
dale (1994). Resolution of this problem 
may result from comprehensive study t)f to- 
nosensilla for all lar sal instars across Lep- 

ANt) Bl(Xii:(XiKAPIIV 

The inclusion of D2 with .SOI and ,Sn2 
within a conunon microtrichiated held in 
1'Ihi.\mi.\ supports a monophyletic relation- 
ship wilh the Asian/Australasian stem-bor 

ing Hcpialidae with the exception of Zclo- 
typia that is characterized by two separate 
micri)tiichiated areas for SDl and SD2. 
Dugdale's ( 1994) reference to all three setae 
being included in Zelolypia appears to be 
incorrect (Dugdale 1999, pers. et)mm.). A 
further larval character that may support a 
close aflinily between Aeiwlus and Pluissns 
is the elongate shape of the pit L.^a on the 
mesothorax and metathorax. This yi\\ is 
slighdy elongate in Zelolypia and ilmlocli- 
1(1. The pit is rt)und in larvae of the root/ 
stem boring Phyiiuilopiis califoniktis 
(Hoisduval) of North America, and the de- 
trital feeding Diinthleloiuiis iiniiucniildlii 
(.Salmon) (as Trioxycaiuis enysii of authors) 
in New Zealand (Grehan et al. 1983). The 
presence and shape of pits have been over- 
looked in many larval descriptions but may 
provide signilicani phylogenetic characters. 
The anterior ntargin of the labrum o\' Aene- 
tiis. /'luiMsiix. and luiclocliki is trilobate, a 
feature also recorded from Stliciiopis. Pliv- 
iiuitopiis. Zelotypia. and Ia'Io (Table I ). The 
larval description of Cihyra scrta (Schaus) 
by Rojas de Hernandez and Chacon de Ul- 
loa (1982) illustrates a prothoracic micro- 
Irichiated Meld common to SDl and SD2 
(hat excludes D2. Larval descriptions of the 
South American wood-borer Tr'uhophassiis 
are insuflicient lo conliini a triselose setal 
pi(, and the adul( male of I ricliopluissiis i-i- 
giinlciis lacks metatibial androconia (Bri- 
quolet 1956). Kristensen (1998) notes that 
metatibial androconia have evolveil several 
times in Lepidoptera. Androconia ol On- 
copcrii and Feiirniylrans may have origi- 
nated separately from those of the wood- 
borer lineages since the scales of Aenetiis, 
lindoclita and Pliiissus are pale reddish 
brown or orange brown in contrast to the 
gray brown androconia of Oncopcra and 
I'lwnnylraiis (Table 1 ). Al least one species 
of Afuctiis lacks metatibial androconia 
(Wagner and Rosovsky 1991). 

Kristensen's (1998) suggestion for a 
close relationship between the monotypic 
wood-boring genus Lclo of South Africa 
and Ihe Auslralasian Acitclus was based on 



biogeographic speculations and is not oth- 
erwise substantiated (Kristensen pars, 
comm., 1999). Metatibial androconia are 
absent from Leto veinis (Cramer) and larvae 
have no michrotrichiated fields at the base 
of SDl. SD2, or D2. Although larvae of L. 
veniis are wood-borers, callus feeding is not 
recorded and pupae reside in a unique tu- 
bular silk/frass extension of the tunnel be- 
yond the bark surface (Peabody Museum of 
Natural History specimens). We conclude 
the genera Phossiis. EiulocUta, and Aenetus 
represent a monophyletic lineage in refer- 
ence to the trisetose pit, with the possibility 
that Zelotypia and Cihyra are more distant- 
ly related, and possibly comprising a mono- 
phyletic lineage of callus feeders and wood- 
borers along with the Phymatopits-clade 
within the Hepialidae sensu stricto. 

Exoporian and hepialid lineages are pre- 
dicted by Nielsen et al. (2000: 832) to be 
'very much' older than the fragmentation of 
Gondwana. The oldest fossil record for he- 
pialoids are Paleocene Europe, mid-Mio- 
cene China (Kristensen and Skalski 1999) 
and Eocene New Zealand, the latter being 
fossil wing scales that may be referable to 
the extant genus Wiseana Viette (Evans 
1931. Harris 1984). A purported Upper 
Cretaceous amber mnesarchaeid wing is 
considered by Kristensen and Skalski 
(1999) to be unsubstantiated in the absence 
of "strong family autapomorphies" in the 
specimen. In the absence of a well-endowed 
fossil record, Holloway and Nielsen (1999) 

regard the age of Mesozoic events influenc- 
ing distribution patterns within Lepidoptera 
to be an open question. Grimaldi's (1999) 
inteipretation of fossil evidence proposed 
an Upper Jurassic origin for tongued Lepi- 
doptera (Glossata) and a Cretaceous origin 
for basal glossatan families (including He- 
pialidae). Grehan (1991) suggested the bio- 
geographic patterns of Lepidoptera and An- 
giospermae support a pre-Cretaceous origin 
for lepidopteran lineages to family level. 
Lack of evidence for discrete continental 
monophyletic exoporian faunas is contrast- 
ed by Nielsen et al. (2000:832) with their 
"temptation" to view the Exoporia as rel- 
icts of Gondwanic fragmentation and re- 
sulting isolation and speciation. 

Most Mesozoic rnodels of evolution are 
linked to the geological fragmentation mod- 
els of Pangaea or its Gondwanic and Laur- 
asian fragments (Craw 1982). Distribution 
patterns congruent with this history are ex- 
pected to exhibit distributional and phylo- 
genetic links across the Atlantic and Indian 
Oceans resulting from the breakup of an- 
cestral distributions on the supercontinents 
of Gondwana and Laurasia. This historical 
model is consistent with the distributions of 
primitive hepialid lineages (Fig. 9) com- 
prising a biogeographic track connecting 
Australia, Africa, North America, and Eur- 
asia (this connection does not assume a 
monophyletic status for these genera). In 
contrast, the Endoclita/Aenetus/Phassus 
clade is absent from Africa, although pres- 

Figs. 9-12. 9, Biogeography of "primitive" Hepialidae. A minimal .spanning link connects the generalized 
di.stributions of the African Afrotheora and Antihepialus with Fraus across the Indian Ocean basins (with baseline 
of track denoted by .sohd square) with an additional link to Gazoryctra between the African genera and the 
North American and Eurasian Gazoryctra. This pattern may be compatible with conventional "Pangaean" origin 
although the genera are currently not known to be monophyletic (distribution data from Holloway and Nielsen 
1999: Fig. 21-22). 10, Pacific biogeography of Entloclita, Aenetus, and Phassus. The nearest neighbor criterion 
links the distributions oX Aenetus (Australasia) and Entloclita (India-eastern Asia) with the American Phassus 
across the Pacific (baseline as a solid square). This spatial homology suggests the evolution of this lineage is 
more closely linked to the geological history of the Pacific basin than with the Atlantic or Indian oceans of 
Gondwana (distribution data from Grehan 1987, Nielsen and Robinson 1983). 1 1, Pacific interpretation for the 
distribution of Pliyniutof)us-c\ade. Although the North American ' Phymatopus' is in closest geographic proximity 
to the related European Phymatopus directly across the Atlantic, the western distribution of 'Phymatopus' may 
be the result of a former trans-Pacific connection through extinction of Asian representatives as indicated here 


by a north Pacific track and baseline (distribution data from Wagner 1985). 12. Pacific interpretation for the 
distribution of the Sthenopis-Zenophassus clade. As with the Phymatopus group, the Old World-New World 
disjunction between Sthenopis and Zenophassus may lie across the Pacific rather than Atlantic Ocean basins 
(dislribution data from Tindale 1941 — including Chinese records of 'Sthenopis' aucttorum nee Packard [1865] 
of uncertain status (Nielsen et al. 20001). 


Fig. 13-14. Pacific spatial patterns. 13, Track for perichaetine earthworms (compiled from Easton 1987). 
14. Track for the fabaceous genus Onnosia (from Croizat 1976, Fig. 1). 

ent in the Gondwanic fragments of India, 
Australasia, and South America (Fig. 10). 

A Gondwanic history for wood-boring 
Hepialidae may be supported by a close 
phylogenetic relationship being established 
with an African group such as Leio (Grehan 
1984). Alternatively, the African gap may 
be accounted for by extinction of Gond- 
wanic members or a biogeographic history 
for wood-boring and callus-feeding Hepi- 
alidae that bypasses Africa altogether. 
Gondwanic distributions bypassing Africa 
include tracks connecting Central and North 
America with the Mediterranean and Indian 
Ocean via the Tethyan geosyncline (Croizat 
1964). Absence of wood-boring Hepialidae 
from North Africa and Europe does not 
support this biogeographic history. Distri- 
bution of the EndocUtalAenetiislPhassus 
group is, however, consistent with Africa 
being 'bypassed" by a non-Gondwanic ori- 
gin involving the Pacific Basin. This bio- 
geographic connection may also be appli- 
cable to the Phymatopus-clade (Fig. 1 1 ) and 
to SthenopislZenophassus (Fig. 12). There 
are also similarities in the geographic rang- 

es of AenetitslEiuloclita with other Pacific 
groups such as the perichaetine earthworms 
(Fig. 13) in the Old World (Easton 1987) 
and the angiosperm genus Onnosia Jackson 
(Fig. 14) in both Old and New Worlds (Cro- 
izat 1976, fig. 1). 

The geological evolution of the Pacific is 
a controversial biogeographic and geologi- 
cal subject. Geohistorical reconstructions 
treating the Pacific as a permanent oceanic 
basin are contradicted by the extensive doc- 
umentation of allochthonous terranes 
around the Pacific Rim and Tethyan geo- 
syncline. These terranes are widely inter- 
preted as fragments of former Mesozoic 
and Tertiary island arcs or microcontinents 
of Pacific origin (Craw et al. 1999, Grehan 
2001 ). Geological efforts to resolve the his- 
torical relationships between the circum-Pa- 
cific terranes include proposals for disrup- 
tion of Mesozoic microcontinental frag- 
ments (e.g., Nur and Ben-Avraham 1977, 
1989), fragmentation of oceanic super- 
plume magmas (Kimura et al. 1994), and 
former island-arc bounded plates (Moores 
1998). Pacific distributions comprise a bio- 


geographic element distinct from Gond- 
wanian or Laurasian distributions spanning 
the Indian Ocean and Pacific basins (Cro- 
izat 1958, 1976) and current biogeographic 
studies continue to verify a distinct Pacific 
pattern of biogeography for groups ranging 
from cycads and conifers (Contreras-Me- 
dina et al. 1999) to angiosperms (Heads 
1999). dragonflies (De Marmels 2000), and 
dinosaurs (Rieppel 1999). A Pacific biogeo- 
graphic homology for the origin of an En- 
doclitalAenetuslPhassiis lineage, possibly 
along with other Hepialidae. provides a his- 
torical solution to the absence of Exoporia 
from West Africa (except for the marginal 
presence of Aiuihepiahis in western Congo/ 
Zaire) and Madagascar described by Niel- 
sen et al. (2000: 831 ) as "extraordinary and 
inexplicable." Absence of taxa from West 
Africa and Madagascar, far from extraor- 
dinary, is commonplace with many such 
groups being Pacific in origin whereas West 
Africa and Madagascar are regions central 
to the Atlantic and Indian Ocean biogeo- 
graphic patterns of Gondwana (Croizat 
I952. 1958. 1968a-b). The Pacific homol- 
ogy proposed here for wood-boring Hepi- 
alidae corroborates the caution expressed 
by Nielsen et al. (2000) against interpreting 
distribution of Exoporia as relicts of Gond- 
wanic fragmentation. The lack of exoporian 
monophyly within continents (Nielsen et al. 
2000) may be the result of ancestral differ- 
entiation predating geological dissolution of 
both Pacific and Gondwanic regions. 

We are grateful to D. R. Strong (Univer- 
sity of California, Davis) for providing lar- 
val specimens of Pliyinatopiis californiciis. 
E. L. Quinter (American Museum of Nat- 
ural History), D. R. Davis and P. Gentilli 
(National Museum of Natural History. 
Smithsonian Institution). P. T. Dang (Ca- 
nadian National Collection). T. L. McCabe 
(New York State Museum). M. Cochrane 
(South African Museum). R. Pupedis (Pea- 
body Museum of Natural History), and C. 
Young (University of Tasmania) for facili- 
tating loans, to J. S. Duadale (Landcare. 

New Zealand). N. P. Kristensen (University 
of Copenhagen). C. Young (University of 
Tasmania), and G. W. Gibbs (Victoria Uni- 
versity of Wellington. New Zealand) for 
comments on the manuscript, and to Jane 
Hyland (Carnegie Museum of Natural His- 
tory) for meticulous illustration of morpho- 
logical features. 

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105(3). 2003. pp. 756-768 





R.^CHEL Braud. D.'Wid R. Smith. John S. Strazan.AiC, .^nd Linda Butler 

(RB, JSS, LB) Division of Plant and Soil Sciences. West Virginia University, Morgan- 
town. WV 26506-6108. U.S.A. (e-mail;; 
lbutler@w\ (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research 
Service, U.S. Department of Agriculture, c/o National Museum of Natural History. Smith- 
sonian Institution. Washington. DC 20560-0168. U.S.A. (e-mail: dsmith@sel.barc.usda. 

Abstract. — Adult Symphyta richness and abundance were studied using data collected 
from 36 Malaise traps in the Monongahela National Forest. Pocahontas County. West 
Virginia, and in the George Washington National Forest. Augusta County, Virginia. A 
total of 8.884 adults representing 155 species in 49 genera and 8 families were collected. 
Diversity estimators suggest that approximately 81% of the actual species present on the 
two forests were sampled over five years (1995-1999). Fifty percent of all adults (4,481) 
were Acordiilecera dorsalis Say (Pergidae). The next most abundant species were Pris- 
tiphora biinksi Marlatt (Tenthredinidae) (12.8%), Pachynematiis comiger (Norton) (Ten- 
thredinidae) (4%), Pracharactus rudis (Norton) (Tenthredinidae) (3%), and Taxomis pal- 
lipes (Say) (Tenthredinidae) (2%). Larval symphytan richness and abundance were deter- 
mined by foliage collections from Quercus spp., Ccirya spp., and Acer spp. Symphytan 
larvae from foliage numbered 11,621 specimens representing eight genera. Sixty-three 
percent of all larvae (7,373) were Acordidecera spp. The next most abundant genus was 
Periclista (2,328) which accounted for 20% of the total larvae. Differences in species 
richness and abundance of both adults and larvae occurred between forests and between 

Key Words: Symphyta. richness, abundance, diversity. Acordidecera dorsalis 

Symphyta are an important and abundant Symphyta associated with forest ecosys- 

part of the insect fauna in hardwood forests, terns. Malaise traps commonly sample adult 

The adults are pollinators (Goulet 1996) sawflies, but have seldom been used to as- 

and the larvae of at least two species have sess sawfly community composition, 

been responsible for heavy defoliation of Though Symphyta comprised the bulk of 

oaks (Eidt and Nichols 1970. Matuzewski the Hymenoptera collected by Malaise traps 

and Ward 1977. Hutchinson 1998). Much in one New York survey (Matthews and 

attention has been given to the conifer-feed- Matthews 1970), the focus was not Sym- 

ing Symphyta: however, little is known phyta. so the researchers gave no details 

about the richness or abundance of the about the species captured or their abun- 

Symphyta that feed on hardwoods and other dance. Similarly, a study comparing the ef- 


Fig. 1. Location ot study plots in the George Washington National Forest and Monongahela National Forest. 

fectiveness of different type.s of Malaise 
traps also mentioned Symphyta (Darling 
and Packer 1988), but they gave no infor- 
mation regarding richness or abundance in 
their predi)minantly oak setting. Other sur- 
veys have been conducted in urban envi- 
ronments (Smith and Barrows 1987) or 
have focused on a single genus (Smith 
1991). The purposes of this study were to 
use Malaise traps for adults and foliage col- 
lections for larvae to determine which saw- 
fly species are present in oak-dominated 
forests, measure their relative adult abun- 
dances, and document their adult seasonal 

Materials and Methods 

This study was part of a long-term anal- 
ysis of nontarget effects of Bacillus tliiirin- 
giensis variety kurstaki and Gypchek® 
when used to suppress Lymontria dispor 
(L.) (Lymantriidae). Eighteen 200-ha study 
plots were established in gypsy moth sus- 
ceptible, oak-dominated forests. Plots one 
through nine were located in the Deerheld 
Ranger District of the Georsze Washington 

National Forest (GWNF), Augusta County, 
Virginia (Fig. 1 ). The GWNF plots range in 
elevation from 586 to 791 m and are locat- 
ed in a xeric forest of mixed oak and pine. 
Plots 10 through 18 were located in both 
the southern Greenbrier Ranger District and 
the Marlinton Ranger District of the Mo- 
nongahela National Forest (MNP), Poca- 
hontas County, West Virginia (Fig. 1). The 
MNF plots range in elevation from 860 to 
1,070 m and are more mesic than the 
GWNF plots (Butler and Strazanac 2000). 
These forests were selected for study in 
1994 because they contained a high per- 
centage of gypsy moth-preferred heists and 
were located ahead of the leading edge of 
gypsy inost movement. Each 200-ha plot 
contained a 30-ha subplot within which 
were two sites, one located on a ridge and 
the other in a valley or near a stream. 

Sampling adult Symphyta was done us- 
ing Townes-style Malaise traps (Townes 
1962). Each trap was designated by both 
plot number and lower or upper site. Two 
Malaise traps were operated per plot, one 
on each site for a total of 36 traps for the 


Study. Each trap was oriented on the plot 
so that its spine ran east-west. Each Malaise 
head contained a jar with 175 ml of 109c 
ethanol and was collected on the same day 
each week and replaced with a fresh jar of 
ethanol. Sampling was for 15 weeks from 
early May through mid-August for five 
years. In the laboratory, sawflies were sep- 
arated from other insects. Specimens were 
mounted and identified by DRS. 

For larvae, five foliage samples per plot 
were taken each of 15 weeks from the low- 
er and middle forest canopy using alumi- 
num pruning poles with large plastic catch 
bags. Foliage samples were taken well 
\\ ithin the boundaries of each plot, but just 
outside the bourdary of each subplot. One 
sample consisted of 21 branch-tip clippings 
from any species of maple (Acer spp.) ex- 
cept striped maple. The second sample con- 
sisted of 15 branch-tip clippings of any spe- 
cies of hickory {Caiyo spp.). The final three 
samples each consisted of 21 clippings of 
oak and contained either members of the 
red oak group (including scarlet oak (Qiier- 
ciis coccinea Muenchh.), black oak {Q. ve- 
lutina Lam.), and red oak (Q. rubra L.)]: or 
the white oak group [chestnut oak (Q. pri- 
iiiis L.); or white oak (Q. albo L.)]. The 
foliage samples were taken to the labora- 
tory and stored in a walk-in cooler until the 
arthropods could be removed from the fo- 
liage by hand during the following two 
days. All symphytan larvae were removed 
from the foliage and preserved in 709^ al- 
cohol for identification. Specimens were 
identified by DRS and RB. 

Voucher specimens of adults and lar\ ae 
are deposited in the National Museum of 
Natural History, Smithsonian histitution. 
Washington, DC, and the West Virginia 
L'ni\ersity Arthropod Collection, Morgan- 
town, WV. 


Malaise samples provided a wealth of in- 
formation about adult symphytan richness, 
abundance, and seasonal occurrence in a 
hardwood forest. Data were analvzed usint; 

the Estimates 5 program (Colwell 1997). 
Diversity estimators produced a fairly nar- 
row range of species estimates for both for- 
ests combined and each forest individually. 
The bootstrap estimations were at the low 
end of the range for both forests combined 
and each forest separately. For example, 
bootstrap estimated that both forests com- 
bined may have 174 species. The high end 
of the range was determined by second or- 
der jackknife, which estimated 214 species 
for both forests (Table 1). The Chao 2 and 
ICE estimators produced moderate esti- 
mates (191 and 192, respectively) and may 
be more realistic for this data set. These two 
estimators suggest that approximately 81% 
of the actual species present on the two for- 
ests were sampled over five years. Species 
accumulation curves and weekly richness 
were plotted for the combined forests (Fig. 
2), the GWNF (Fig. 3), and the MNF (Fig. 

The number of adult specimens captured 
over five years was 8,884 with 155 species 
identified: 104 from the GWNF and 127 
from the MNF (Table 2). These adults rep- 
resent eight families, with most species 
Tenthredinidae (121), followed by Pamphi- 
liidae (14). Argidae (7), Pergidae (4), Ce- 
phidae (3), Xiphydriidae (3), and Xyelidae 
(2). Pergids were the most abundant with 
4,529 individuals, 4,481 being Acorditle- 
cera dorsalis Say. Tenthredinid total abun- 
dance was 4,240. followed by Xyelidae 
(33), Argidae (31), Pamphiliidae (28). 
Xiphydriidae (15), Cephidae (7), and Di- 
prionidae ( 1 ). Pergids made up 59% of the 
abundance in the GWNF, but only 4% of 
the species richness. In the MNF, pergids 
made up 38.6% of abundance and account- 
ed for 3% of the richness. However, the ten- 
thredinids collected in the GWNF com- 
prised 76% of the richness and 39.9% of 
the abundance. MNF tenthredinids account- 
ed for 81% of richness and 59.7% of abun- 
dance. The more abundant species follow- 
ing A. dorsalis were Prisriphora baiiksi 
Marlatt, Pachynematus corniger (Norton), 
ParacharactKs rudis (Norton). Ta.xoniis 



1996 1997 1998 1999 

Richness Species Accumulation 


o 125 

I.I Iiillliililllllli.lii.llll.liilll.i.lllli 





Richness Species Accumulation 

Figs. 2-3. Species accumulation curves. 2 (Top), Combined George Washington and Monongahcla national 
forests for \5 weeks each year from 1995-1999. 3 (Bottom), George Washington National Forest for 13 weeks 
each year from 1993-1999. 

pill li pes (Say), Aneuginenus jiavipes (Nor- 
ton), Taxoniis pallidicornis (Norton). Ne- 
iiuitiis radialis Smith, Taxoims nifociiictiis 
(Norton), and Ametastegia pulchella (Roh- 
wer). All of these species were taken from 
both forests, though not in the same order 
of abundance (Table 3). Although species 
richness was high, as measured by the 155 
species of Symphyta caught in the forests. 

87 were represented by five or fewer spec- 
imens (Table 4). 

Many species were collected from only 
one of the two forests. Twenty-eight species 
were found exclusively in the GWNF. Fam- 
ilies which contained species unique to 
GWNF were Argidae (3), Cephidae ( I ), Di- 
prionidae (1), Pamphiliidae (5), and Ten- 
thredinidae (18). Fifty-one species were 


Table \. Species richness estimates using several 
diversity estimators for the George Washington Na- 
tional Forest (GWNF), Monongahela National Forest 
(MNF), and both combined. 


















2nd-order jackkn; 





found only in MNF, and these were in the 
famihes Cephidae (1), Pamphihidae (6). 
Tenthredinidae (42), and Xiphydriidae (2). 
Sixty-one species were collected from both 

Trends in abundance for certain species 
were evident over the five years, Acordii- 
h'cera dorsalis has a peak flight time at the 
end of May. Sampling after that yielded 
very few specimens. The flight time of 
Pristipliora banksi peaks during the second 
or third week in June, 

Over the five years of sampling, adult 
Symphyta were collected from Malaise 
traps from the earliest date (6 May) through 
the latest (18 August) (Table 4), The ten 
more abundant species usually flew from 
early May until late August. While abun- 
dance peaked for these species at different 
times of the season, they were caught with 
some consistency during the entire three- 
month sampling period. The first five weeks 
of each sampling season were critical and 

accounted for 60% of 1995 adults, 82% of 
1996 adults, 32% of 1997 adults, 47% of 
1998 adults, and 66% of 1999 adults (Table 

Foliage samples produced 11,621 larval 
individuals from 1995 through 1999. Eight 
different genera were represented. Acardii- 
lecera comprised 63.5% of all symphytan 
larvae with 7,373 specimens, while Peri- 
clista (several species) comprised 20.0% 
with 2.328 specimens. Other larvae were 
Nematinae [probably mostly P. chlorea 
(Norton) and Craterocerciis spp.] (1,220), 
Caliroa spp. ( 108), Pamphihidae (3 1 ), Arge 
sp. ( 14). Dimorphopteryx sp. ( 14), and Me- 
gaxyela sp. (3). 

The general trend for Acorduleceni lar- 
vae was a shaip increase over the first few 
sampling weeks, peaking in late May, then 
shaiply declining. Periclista larvae also de- 
clined sharply after their peak in early June. 


While a high diversity of Symphyta oc- 
curred in our samples, there are a few fac- 
tors that may have influenced sample con- 
tent. First, sampling by Malaise traps cap- 
tures only those species that fly at the level 
of the collecting net. Species that live pri- 
marily in forest canopy probably were not 
in the samples or occurred in small num- 
bers. Periclista, for example, comprised 
20% of the total larvae captured but only 
0.01% of the adults. Second, Malaise traps 

Table 2. Adult Symphyta species richness and abundance by family as sampled from 1995 through 1999 in 
the George Washington National Forest (GWNF) and Monongahela National Forest (MNF). 

Speciex Richness 






































































Table 3. The ten more abundant species of adult Symphyta sampled from 1995 through 1999 in the George 
Washington National Forest (GWNF) and Monongahela National Forest (MNF). 




1'^, Total 1 


l'^ lolah 





dor.salis Say 


dor.salis Say 






hanksi Marlatt 


corniger (Norton) 






flcnipes (Norton) 


ritdis (Norton) 






pallidicornis ( Norton ) 


pal Ii pes (Say) 


cincui NewiTian 


hanksi Marlatt 






coini)>ci- (Norton) 


radialis Smith 






pukhclUi (Rohwer) 


riifociiicliis ( Norton ) 






patlipes (Say) 


multicolor ( Norton ) 



fi'imnsa (Klug) 





qiiattiiordecimpiinctaliim ( Norton ) 



ostryae ( Marlall ) 


chlorca (Norton) 


were in use for only 15 weeks starting in excluded entirely. The traps were placed in 

early May. While traps were present during the same location each year, and some spe- 

most species" peak fight times, some spe- cies may have avoided the traps. In most 

cies which were more active either before cases, the species that appeared in low 

or after traps were present may have been numbers such as Sphacophilus cclliikiris 


Fig. 4. 

1995 1996 1997 1998 1999 

^H Richness Species Accumulation 

Species accumulation curve for the Monongahela National Forest for 15 weeks each year from 1995- 


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(Say), Sterictiphora serolino Smith, and Ja- 
nus abbreviatus (Say), have hosts such as 
morning glory, Ipomoea purpurea (L.) 
Roth, black cherry, Prunus serotina Ehrh., 
and Sali.x and white poplar. Popuhis alba 
L., respectively, which are uncommon in 
both forests. Alternatively, species such as 
Acordulecera dorsalis. whose hosts domi- 
nate the forest (Quercus spp. and Carya 
spp.), were found in greater abundance. 
However, some Symphyta did not fit this 
pattern. The host plant for Pristiphora 
banksi, Vacciniuin, was not found in great 
abundance in the MNF and yet it was the 
second most abundant species of Symphyta. 

A high diversity of symphytan larvae 
was collected even though only five host 
plant groups were sampled. These larvae as 
a group show considerable variability in 
their host plants and can feed on foliage of 
larger trees or small groundcover flora such 
as grasses, blueberries, ferns, and other 
plants. Another factor which limited our 
larval diversity is that only foliage from the 
middle to lower canopy was sampled. Sym- 
phytan larvae on other food plants in the 
forest or feeding on upper canopy foliage 
were not sampled using this method. The 
adult abundance and richness were affected 
by seasonal sampling time; the same is true 
of the larvae. Assessing population fluctu- 
ations and abundances is limited since our 
sampling period was only 15 weeks from 
early May through mid-August. Some sym- 
phytan larvae are present before, or after 
this period, or both. Seasonal population 
changes over the five-year period could be 
responsible for overestimated and underes- 
timated abundances when comparing sam- 
ples to predict species diversity. Larval 
feeding habits may have been a factor. For 
instance, Acordulecera larvae were espe- 
cially problematic because they were in the 
only genus sampled that existed in large 
gregarious clusters. A single sample can 
contain hundreds of larvae. 

As with other species rich taxa, Symphy- 
ta diversity plays an important role in the 
forests. They are likely also important in 

Table 3. Weekly number of adult Symphyta col- 
lected by Malaise trap.s from 1995 through 1999 in 
both the George Washington National Forest and Mo- 
nongahela National Forest. 


Week ]'-!•->> IWfi IW7 IWS mw 





























































1 1 






























forest food webs as herbivores preyed upon 
by numerous species of invertebrates and 
vertebrates. The cruciform larvae of Sym- 
phyta are similar to lepidopteran larvae 
which are recognized as having both high 
fat ( Redford and Dorea 1 984 ) and very low 
chitin content relative to other insects, thus 
making them especially valuable in the di- 
ets of songbirds (Sample et al. 1993). Some 
ornithologists collectively refer to larval 
sawflies and lepidopterans as caterpillars 
when recording dietary habits of songbirds 
(Rodenhouse and Holmes 1992). As abun- 
dance of spring defoliating lepidopterans 
may be reduced during Btk applications for 
gypsy moth suppression (Marshall et al. 
2002), sawfly larvae may survive to be 
preyed upon by songbirds. Understanding 
the diversity, abundance, and seasonality of 
adult and larval Symphyta in a hardwood 
forest ecosystem may help make predic- 
tions about how certain chemical or biolog- 
ical controls effect Symphyta and the forest 
food web. 


We thank Cynthia J. Frizler, Crystal B. 
Mayle, graduate students, and our summer 


Student workers for assisting in collecting 
and sorting the specimens. Special thanks 
to Cathy J. Apgar. Systematic Entomology 
Laboratory (SEL). USDA, who mounted, 
labeled, and did preliminary sorting of 
many thousands of specimens. We also 
thank the following reviewers for their 
comments: H. Goulet. Agriculture Canada. 
Ottawa and A. Konstantinov and J. Brown 
(SEL). This research was funded by the 
USDA Forest Service as cooperative agree- 
ment 42-793, and we are grateful for that 
support. Published with the approval of the 
Director of the West Virginia Agriculture 
and Forestry Experimental Station as sci- 
entific article #2839. 

Literature Cited 

Butler, L. and J. Strazanac. 2000. Macrolepidopteran 
larvae sampled by tree bands in temperate mesic 
and xeric forests in eastern United States. Pro- 
ceedings of the Entomological Society of Wash- 
ington 102: 188-197. 

Colwell, R. K. 1997. EstimateS: Statistical estimation 
of species richness and shared species from sam- 
ples. Version 5. 

Darling, D. C. and L. Packer. 1988. Effectiveness of 
Malaise traps in collecting Hymenoptera: The in- 
fluence of trap design, mesh size, and location. 
The Canadian Entomologist 120: 787-796. 

i;idt. D. C. and J. O. Nichols. 1970. An outbreak of a 
sawfly, Pamphilius phyllisae (Hymenoptera: Pam- 
philiidae), on northern red oak, with notes on lar- 
val morphology. The Canadian Entomologist 102: 

Goulet, H. 1996. Revision of the Nearctic species of 
the arcuata group of the genus Tenthredo with 
notes on the higher classification of the Tenthre- 
dinini (Hymenoptera: Symphyta: Tenthredinidae). 

Contributions of the American Entomological In- 
stitute 29(2): 1-135. 

Hutchinson. S. F 1998. 1998, Oak slug sawfly survey 
{memo I. West Virginia Department of Agricul- 

Marshall, M. R., R. J. Cooper, J. A. DeCecco, J. Straz- 
nac, and L. Butler. 2002. Effects of experimentally 
reduced prey abundance on the breeding ecology 
of the red-eyed vireo. Ecological Applications 12: 

Matthews. R. W. and J. R. Matthews. 1970. Malaise 
trap studies of flying insects in a New York mesic 
forest I. Ordinal composition and seasonal abun- 
dance. Journal of the New York Entomological 
Society 78: 52-59. 

Matuszewski. M. and J. D. Ward. 1977. Oak sawfly 
infestaion. Kentucky — 1997. Division of Forestry, 
Frankfort, KY. 

Redford, K. H. and J. G. Dorea. 1984. The nutritional 
value of invertebrates with emphasis on ants and 
termites as food for mammals. Journal of Zoolo- 
gy. London 203: 385-395. 

Rodenhouse, N. and R. T Holmes. 1992. Results of 
experimental and natural food reductions for 
breeding black-throated blue warblers. Ecology 
73: 357-372. 

Sample, B. E., R. J. Cooper, and R. C. Whitmore. 
1993. Dietary shifts among songbirds from a di- 
flubenzuron-treated forest. The Condor 95: 616- 

Sniith. D. R. 1991. Flight records for twenty-eight spe- 
cies of Macrophya Dahlbom (Hymenoptera: Ten- 
thredinidae) in Virginia, and an unusual specimen 
of M. epinnta (Say). Proceedings of the Entomo- 
logical Society of Washington 93: 172-775. 

Smith, D. R. and E. M. Barrows. 1987. Sawflies (Hy- 
menoptera: Symphyta) in urban environments in 
the Washington, D. C. area. Proceedings of the 
Entomological Society of Washington 89: 147- 

Townes, H. 1962. Design for a Malaise trap. Proceed- 
ings of the Entomological Society of Washington 
64: 253-262. 


105(3). 2003. pp. 769-775 


E. Richard Hoebeke. Emily J. Sensenb.^ch. John P. S.'^nderson, and 
Stephen P. Wraight 

(ERH. EJS, JPS) Department of Entomology. Comstock Hall, Cornell University, Ith- 
aca, NY 14853, U.S.A. (e-mail:; ejs47(<?; jps3@cornell.edii), 
(SPW) USDA, ARS, Plant, Soil, and Nutrition Laboratory, Tower Road, Ithaca, NY 
14853. U.S.A. (e-mail: 

Abstract. — The Palearctic muscid Coenosia attcmiata Stein is reported for the hrst time 
in North America based on collections from the U.S.A. and Canada. Adult flies were 
collected from commercial greenhouses in Erie, Monroe, Onondaga. Suffolk. Tompkins, 
and Wayne counties of New York State and Ontario Province. Canada, and from a Malaise 
trap in a suburban backyard of Los Angeles County, California. A diagnosis, redescription, 
and photographs of the adult are provided to help distinguish it from other North American 
Muscidae. Its biology and habits are summarized from the European literature, and its 
Old World distribution is reviewed. 

Kcx Words 

Diptera, Muscidae, Coenosia atlenuata. North America, immigrant species, 
new record. New York. California, Ontario 

In late August of 2002, several speci- 
mens of a small muscid fly, collected by 
EJS in a greenhouse in East Syracuse. New 
York, were submitted to ERH for identifi- 
cation. They proved to be Coenosia atten- 
iicita Stein, an obligatory predaceous fly in- 
digenous to the Paleotropical Region and 
not known to occur in North America. 

In this paper, we give the first North 
American records for C. attenuata. review 
the Old World literature treating this spe- 
cies, provide notes on its biology and hab- 
its, and redescribe and provide photographs 
of the adult to enable recognition of this 
species in the North American fauna. 

Specimens of C attenuata were first no- 
ticed in early October 1999 by Elise Schil- 
lo-Lobdell, an IPM scout, at a commercial 
greenhouse specializing in the large-scale 
production of poinsettias. It was at this 

same greenhouse where specimens were 
later collected by EJS on 28 August 2002 
and identified by ERH. 

Since this initial collection, numerous 
other specimens have been collected on yel- 
low sticky card-traps set for monitoring 
pest populations (including shore flies and 
fungus gnats) in other greenhouses across 
New York as well as at a site in Ontario, 
Canada. Quite by accident and about the 
same time, the senior author also learned 
from Or Adrian Pont, a muscid fly spe- 
cialist from Great Britain, that he had re- 
cently identified specimens of C attenuata 
from California; these specimens were col- 
lected from a Malaise trap in a suburban 
backyard in Los Angeles County in mid- 
September of 2002 (Brian V. Brown, per- 
sonal communication ). 


Coenosia atteniiata Stein 

(Figs. 1-4) 

(Synonymy after Pont 1986) 

Coenosia atteniiata Stein in Becker 1903: 

Coenosia (Caricea) Jiavicornis Schnabl in 

Schnabl and Dziedzicki 1911: 80. 
Coenosia confalonierii Seguy 1930: 86. 
Coenosia (Caricea) afftnis Santos Abreii 

1976: 13. 
Coenosia [Caricea) fiavipes Santos Abreu 

1976: 13. 

Diagnosis. — Coenosia atteniiata is a 
member of the nominate subgenus Coeno- 
sia Meigen {sensii Huckett 1934) or the ti- 
grina-group (sensii Hennig 1961), charac- 
terized by the hind tibia with two closely 
adjacent bristles at the middle, situated on 
the anterodorsal and anterior surfaces re- 
spectively (illustrated by Hennig 1961: 521, 
tig. 210B. and Fig. 1, herein). The Old 
World species C. tigrina F, C. atra Meigen. 
C luimilis Meigen, and C. strigipes Stein 
also belong to this tigrina-group. occasion- 
ally placed in its own genus Caricea Ro- 
bineau-Desvoidy (Skidmore 1985): all are 
members of the greenhouse predator com- 
munity (Kiihne 2000), and are occasionally 
referred to by Old World workers as either 
■"hunter flies" or "killer flies." Coenosia 
tigrina and C. Ininiilis are recorded from 
North America (see remarks). From these 
two introduced species, C. atteniiata can be 
easily differentiated by its smaller size 
(2.5-3.0 mm, male; 3.0-4.0 mm, female) 
(vs. 4.75-5.75 mm, male and 5.75-7.0 mm, 
female for C tigrina: and 3.0-3.5 mm, 
male and 3.0-5.5 mm, female for C. hti- 
niilis: measurements from d'Assis Fonseca 
1968), legs of the male entirely pale yellow 
(vs. femora mostly black but broadly red- 
dish yellow at apex in male of C. tigrina. 
and femora entirely black in male of C lui- 
milis). and the distinctive male genitalia 
(Fig. 2). 

Redescription [terminology for structures 
follows McAlpine (1981)]. Male and fe- 
male habitus. Fins. 3 — t. Redescribed from 

29 specimens (East Syracuse, Onondaga 
Co., NY). Length: 2.5-3.0 mm, male (n = 
20); 3.0-4.0 mm, female (n = 9). 

Male. — Head: Parafacial, fronto-orbital 
plates, frontal vitta, and face silvery-white 
pruinose, with yellowish tinge; black 
ground color not visible under pruinosity 
except slightly on face. Pedicel silvery- 
white pruinose (with yellow tinge), black 
ground color visible at exteme base. Fla- 
gellomere 1 also silvery-white pruinose 
(with yellow tinge). Aristomere 1 very 
short; aristomere 2 not more than 2x as 
long as wide; aristomere 3 very long, even- 
ly tapered to tine tip, with dense short hairs 
on basal half, longest of which equal basal 
diameter of aristomere 3. Orbital plate with 
3 strong setae: lower and middle setae la- 
teroclinate; upper seta reclinate. Ocellar and 
postocellar setae well developed. Inner ver- 
tical setae strong, parallel. Parafacial and 
facial ridge bare. Lower facial margin pro- 
truding slightly just below vibrissa when 
\ iewed in profile. Maxillary palpus yellow, 
slightly clavate apically, moderately haired. 

Thorax: Black in ground color, except 
dorsum of scutum and scutellum light gray 
pruinose, with slight metallic blue tinge 
scarcely visible. In some specimens, 3 in- 
distinct brownish lines or stripes visible on 
scutum. Acrostichal setae small to medium, 
arranged in roughly 2 irregular rows: ap- 
proximately 4-6 presutural setae, and 6—8 
postsutural setae. One strong presutural 
dorsocentral seta. Three strong postsutural 
dorsocentral setae. Two medium intra-alar 
setae. Two (one small, one strong) postalar 
setae. One strong supra-alar seta. Scutellum 
with one pair of strong crossed apical setae, 
one pair of strong subapical setae, and one 
pair of small basal setae. Dorsum of scu- 
tellum with several small discal setae. Ane- 
pisternum with at least 4-5 setae, 2 of these 
strong and extended to margin of lower ca- 
lypter. Katepisternum with three strong se- 
tae. Wings with veins rj+j and m parallel to 
margin. Halter yellow. Legs entirely yel- 
low; tarsi appearing darker due to dense 
coverinc of numerous small black setulae. 


FigN. 1-2. Coenosia attcinuita. 1, Hind tibia. 2, 
Male genitalia (epandriuni, cerci. and .surstylu.s). Both 
figures redrawn from Hennig (1961: Te.xtfig. 210B. p. 
521 and Taf. XXVIII, fig. 533). 

Hind tibiae with two closely adjacent bris- 
tles at middle, situated on anterodorsal and 
anterior surfaces respectively (Fig. 1 ). 

Abdomen: Short, ovoid. T,^, to T^ dark 
gray pruinose, without dark maculations. 
All tergites with numerous small to medium 
discal and marginal setae. Epandrium of 
terminalia gray pruinose, cercus and sursty- 
lus yellow to orangish. 

Male terminalia (Fig. 2); As illustrated 
by Hennig (1961: Taf. XXII, fig. 453; Taf. 
XXVIII, fig. 533) and Cui (1999: 929. fig. 

Female. — Similar to male in most re- 
spects, except larger in size, and in color- 
ation of head, antenna, and legs. Frontal vit- 
ta (not silvery-white pruinose as in male) 
generally dark with metallic blue-green 
pruinosity. Antenna darkened. Legs black 
(with gray pruinosity), except trochanters, 
extreme bases and apices of femora, tibiae, 
and tarsi yellowish. 

Remarks. — Two other species of the ti- 
,i^;7«(v-group are known from North Amer- 
ica, both presumably introduced in the early 
to mid 1800s. Coenosia tigrina was first re- 
ported in North America by Walker ( 1849). 

who originally described it as a new species 
(C. sexmaculata). from Hudson Bay. On- 
tario (later synonymized with C. tigrina by 
Stein 1901). Its known geographic distri- 
bution is decidedly disjunct, occumng from 
British Columbia and Alberta to California 
in the west and from northwestern Ontario 
and Quebec to Michigan and Maine in the 
east (Huckett 1965). Coenosia humilis is 
widely distributed in North America, rang- 
ing from Washington to California and Col- 
orado, and from Wisconsin and Illinois to 
Quebec and New Jersey (Huckett 1934. 
1965). Huckett (1934) reported that the lat- 
ter species occurs frequently in greenhous- 

Biology, habits, and immature stages. — 
In Europe, Coenosia atteimata, along with 
C. atra, C. humilis, C strigipes. C. tigrina. 
and C. testacea Robineau-Desvoidy, are 
members of a greenhouse predator com- 
munity (Kuhne et al. 1994, Kuhne 2000). 
These predaceous flies not only colonize 
greenhouses from the outside environment, 
but may also complete their development in 
greenhouse soil and become established 
there for long periods of time (Kiihne 

Both the larval and adult stages of C at- 
tenitata are predaceous. Larvae have been 
mass reared on dipterous larvae of two 
taxa: Bradysia dijformis Frey (= paiipera 
Tuomikoski) (Sciaridae) (Kiihne 2000) and 
Scatopse transversalis Loew (Scatopsidae) 
(Kiihne 2000). The prey spectrum of adult 
C. attenuata in greenhouses in Baden- 
Wiirttemberg. Germany, includes the green- 
house pests Trialeiirodes vaporarionim 
Westwood and Bemisia tabaci (Gennadius) 
(Aleyrodidae), Empoasca sp. (Cicadelli- 
dae), and Sciaridae (Schrameyer 1991). 
Kiihne et al. ( 1997) also recorded Psychodi- 
dae. Chironomidae, Ephydridae, and Dro- 
sophilidae as prey groups. In captivity, 
adult C. attenuata also consumed the leaf- 
miner Liriomyza huidobrensis Blanchard 
(Agromyzidae) (Schrameyer 1991), and the 
scavenger midge S. tranversalis (Kuhne 


Figs. 3-4. CoenosUi atleniuihi. lateral aspect. .^. Male. 4, Female. Scale line. 1 nun. 


Acct)iding to Schrameyer (1991). adult 
C. cittcmiahi employ a sit-and-wait strategy 
and pursue only prey that are in flight. Am- 
bulatory prey are disregarded, even in ex- 
tremely close proximity. The prey are pur- 
sued, caught, punctured with the proboscis, 
and the liquid body contents ingested. Ma- 
nipulation of the food is acct)mplished by 
specialized mouthparts. A projection on the 
mid-labellum forms a daggerlike tooth to 
puncture prey, and 4-5 "teeth" and a raspy 
"tongue" structure tear the cuticle of the 
prey and mechanically grind it for ingestion 
(Kiihne 2(){)()). The daily capture of a single 
C. attenuata adult appears to be indepen- 
dent of its age. When food is in short sup- 
ply, both adults and larvae may become 
cannibalistic (Kiihne 2(X)()). 

In the laboratory, the adult reproductive 
behavior and duration of the developmental 
stages have been studied by Kiihne et al. 
(1997) and Kuhne (2000). The pre-ovipo- 
sition period is approximately 4 days 
(Kiihne et al. 1997), and most eggs are laid 
within a three week period at 25° C (Kiihne 
2000). The eggs of C. attemiata are of the 
Pluionici-type, sensii Ferrar (1987), and re- 
quire a developmental period of 6.9 days at 
21° C and 5.7 days at 25° C (Kuhne et al. 
1997). The number of larval instars has not 
been directly determined for C. attenuata. 
On the basis of morphological data, how- 
ever. Kiihne (2000) suggests that the larva 
emerges from the egg as a third instar. 
LeRoux and Perron (1960) found this phe- 
nomenon also to be true for C. tigriiun 
based on morphological evidence and the 
fact that larvae do not molt during larval 
life, they concluded that for this species the 
larvae emerge from the egg in the third in- 
star. Complete larval development for C. at- 
tenuata requires 15.4 days at 21° C and 
10.4 days at 25° C, while the pupal stage 
averaged a duration of 15.9 days at 21° C 
and 10.5 days at 25° C (Kuhne et al. 1997). 
The maximum adult lifespan is as long as 
7 weeks in captivity, but only ca. 50% of 
captive populations survive longer than 3 
weeks (Kiihne et al. 1997). 

Distribution. — Coenosia attenuata is re- 
ported from southern Europe (Germany, 
France, Spain, Italy, Greece, Malta, Cy- 
prus); Asia (Syria, Iraq, Israel. Tajikistan. 
Afghanistan); North Africa (Morocco. Al- 
geria. Libya. Egypt); Madeira. Canary Is- 
lands; and the Oriental (widespread), Aus- 
tralasian (Papua New Guinea, Australia), 
and Afrotropical (Cape Verde Islands. So- 
cotra. South Africa) regions (Pont 1986 and 
A. C. Pont, pers. comm.). Most recently. C. 
attenuata has been reported from South 
America (Ecuador, Peru) (Marli'ne/San- 
che/. et al. 2002). 

New York, including at least six counties 
to date (Erie, Monroe. Onondaga. Suffolk, 
Tompkins, and Wayne), and California (Los 
Angeles Co.) in the United States, and On- 
tario. Canada are new records for North 

Material examined. — All specimens de- 
posited in the Cornell University Insect 
Collection. Ithaca, NY, except as noted. 
UNITED STATES: California: Los Angeles 
Co., Monrovia, 34.15°N. 1 17.99° W, 13-14 
Sept. 2002. B. V. Brown, ex Malaise trap 
in suburban backyard (5 i. 2 9); same 
data, except 14-15 Sept. 2002 (1 (5, 4 9) 
(California specimens in the collection of 
the Natural History Museum of Los An- 
geles County). New York: Erie Co., Eden, 
24 Oct. 2002. E. J. Sensenbach. ex sticky 
trap (1 6). Monroe Co.. Gates. 15 Oct. 
2002, EJS (1 (J, 2 9). Onondaga Co.. East 
Syracuse, 28 August 2002, EJS and J PS (8 
6,1 9; of these, 3 S and 3 9 in the col- 
lection of The Natural History Museum. 
London, UK); same data, except 12 Sept. 
2002, ERH (20 6,9 9); same data, except 
12 Sept. 2002, EJS, ex sticky cards (2 6,1 
9). Suffolk Co.. Mattituck. 29 Oct. 2002, 
M. Daughtrey, ex sticky cards (13 6,2 9); 
Riverhead, 29 Oct. 2002, M. Daughtrey, ex 
sticky cards (2 6). Tompkins Co., Lansing, 
30 Oct. 2002, W. Nelson, ex sticky cards (2 
6). Wayne Co.. Macedon. 15 Oct. 2002, 
EJS ( I cJ. 1 9 ); Newark. 15 Oct. 2002, EJS 
(\ 6). CANADA: Ontario: Jordan Station, 
17 Oct. 2002, G. Murphy, ex sticky cards 


(9 6.3 9); same data, except 25 Oct. 2002, 
T. Thiesen (1 cJ. 6 9). All specimens taken 
in New York and Ontario were from com- 
mercial production greenhouses. 

Mode of introduction. — Because delib- 
erate releases have only been reported in 
Germany (Kiihne 2000), we doubt very 
strongly that C. atteiniata was purposefully 
introduced into North America. According 
to Stefan Kiihne (pers. comm. to EJS). in 
the last two years rearings of this predator 
Hy have been initiated in Spain, Italy, and 
Portugal. Moreover, papers by Kiihne, and 
especially K. Schrameyer, suggest these 
Hies are being moved from place to place 
unintentionally with plant material, al- 
though there is no strong evidence to sup- 
port this hypothesis. We would advocate 
that potting media are the most likely 
source for the accidental introduction of C. 
attenuata into North America. 


We thank Kent Loeffier (Dept. of Plant 
Pathology, Cornell University) for provid- 
ing the photographs of the male and female 
in Figs. 1-2. We thank Graeme Muiphy, 
Walter Nelson, and Margery Daughtrey for 
collecting additional samples for us, and 
Elise Schillo-Lobdell for originally alerting 
us to the presence of this predaceous fly in 
New York greenhouses. We also are grate- 
ful to Adrian C. Pont (Oxford University 
Museum of Natural History, Oxford, UK) 
for confirming the identity of C attenuata, 
to him and Brian V. Brown (Natural History 
Museum of Los Angeles County, Los An- 
geles, CA) for each providing a critical re- 
view of a draft of this paper, and to Stefan 
Kiihne (Institut fiir integrierten Pflanzen- 
schutz. Kleinmachnow, Germany) for shar- 
ing information on the rearing of C. atteii- 
itata in Europe (with EJS via e-mail). 

This research was funded in part through 
a Specific Cooperative Agreement between 
the USDA/ARS Plant Protection Research 
Unit and the Cornell University Department 
of Entomolosv, Ithaca, NY (SCA #58- 

1907-9-033) funded by the USDA/ARS. as 
part of the Floriculture and Nursery Re- 
search Initiative. 

Literature Cited 

Becker. T 1903. Aegyplische Dipleren (Fortsetzung 
und Schluss). Mitteilungen aus dem Zoologischen 
Museum in Berlin 2: 67-195. 

Cui. Y. 1999 [1996). Family Muscidae (Coenosia Mei- 
gen). pp. 925-934. In Xue. W.-C. and C. Chao, 
eds. Flies of China. Volume 1 . Liaoning Science 
and Technology Press. Shenyang. 

d'Assis Fonseca, E. C. M. 1968. Diptera Cyclorrhapha 
Calyplrata. Section (b) Muscidae. Handbooks for 
the Identification of British Insects. Vol. X. Part 
4(b). London. Royal Entomological Society. 119 

Ferrar. P. 19S7. A guide to the breeding habits and 
immature stages of Diptera Cyclorrhapha. E. J. 
Brill. Leiden. 2 volumes: 907 pp. 

Hennig, W. 1961. 63b. Muscidae. Gattung Coenosia 
Meigen, pp. 518-619. In Lindner. E.. ed. Die Flie- 
gen der Palaearktischen Region 7(2). 

Huckett. H. C. 1934. A revision of the North American 
species belonging to the genus Coenosia Meigen 
and related genera (Diptera: Muscidae). Part I. 
The subgenera Nemlexiopsis, Coenosia, Hoplo- 
gaster and related genera Allognota, Bithoraco- 
chaeta and Schoenomyza; Part II. The subgenus 
Limosia (Coenosia of authors). Transactions of the 
.Jimerican Entomological Society 60: 57-119; 

1965. Family Muscidae. pp. 869-915. In 

Stone. A.. C. W. Sabrosky. W. W. Wirth, R. H. 
Foote, and J. R. Coulson. eds. A Catalog of the 
Diptera of America north of Mexico. U.S. De- 
partment of Agriculture. Agricultural Research 
Service. Agriculture Handbook No. 276. 

Kiihne. S. 2000. Riiuberische Fliegen der Gattung Coe- 
nosia Meigen. 1826 (Diptera: Muscidae) und die 
Moglichkeit ihres Einsatzes bei der biologischen 
Schiidlingsbekampfung. Studia Dipterologica. 
Supplement 9. 78 pp. 

Kiihne, S.. K. Schrameyer. R. Mullen and E Menzel. 
1994. Riiuberische Fliegen — ein bisher wenig 
beachteter Niitzlingskomplex in Gewach.shausern. 
Mitteilungen aus der Biologischen Bundesanstalt 
fiir Land und Forstwirtschaft Berlin Dahlem, Heft 
302. 75 pp. 

Kuhne. S.. K. Schiller, and U. Dahl. 1997. Beitrag zur 
Lebensweise. Morphologic und Entwicklungs- 
dauer der rauberischen Fliege Coenosia attenuata 
Stein (Diptera: Muscidae). Gesunde Pflanzen. 
Bonn 49(3): 100-106. 

LeRoux. E. J. and J. P Perron. 1960. Descriptions of 
immature stages of Coenosia lii^rimi (F) (Diptera: 
Anthiimviidae). with notes on hibernation of lar- 


vae and predalion by adults, Canadian Entomol- 
ogist 92: 284-296. 

Marti'nez-Sanche/,. A., M. A. Marcos-Garci'a. and A. 
C. Pont. 2002. Coenosia atteiniala Stein. 1903 
(Diptera, Muscidae) nueva especie para la fauna 
Neotropical. Bollettino di Zoologia Agraria e di 
Bachicoltura 34(2): 269-272. 

McAlpine, J. F. 1981. Morphology and terminology — 
adult,s. pp. 9-63. In McAlpine. J. F. ei al.. eds. 
Manual of Nearctic Diptera. Vol. 1. Agriculture 
Canada Monograph 27: 1-674. Ottawa. Canada. 

Pont, A. C. 1986. Muscidae, pp. 57-215. In Soos. A. 
and Papp. L., eds. Catalogue of Polaearctic Dip- 
tera. Volume 1 1 (Scathophagidae — Hypoderma- 
tidae). Budapest. Hungary. 

Santos Abreu, E. 1976. Monografia de los Anthomyi- 
dos de las Islas Canarias (Dipteros). Servicio del 
Aula de Cullura "Elias Santos Abreu." La Palma. 
175 pp. 

Schnabl. J. and H. Dziedzicki. 1911. Die Anthomyi- 
den. Nova acta Academiae Caesareae Leopoldino- 

Carolinae Germanicum Naturae Curiosorum 
95(2): 53-358. 

Schrameyer, K. 1991. Die rauberischen Fliegenarten 
[Coenosia allenuata. C. humilis) ein bedeutender 
Faktor der biologischen Schadlingsbekiimpfung. 
Gesunde Pflanzen 43(12): 398-400. 

Seguy, E. 1930. Risultati zoologici della Missione in- 
viata dalla R. Societa geografica italiana per 
Tesplorazione dell'oasi di Giarabub (1926-1927). 
Insectos Dipteres. Annali Museo Civico di Storia 
Naturale ""Giacomo Doria" 55: 75-93. 

Skidmore, R 1985. The biology of the Mu.scidae of the 
world. Series Entomologica 29: 1-550. 

Stein. P. 1901. Die Walker"schen aus.sereuropaischen 
Anihomyiden in der Sammlung des British Mu- 
seum vu London (Dipt.). Zeitschrift fiir Systema- 
lische Hymenoplerologie und Dipterologie I: 
1 85-22 1 . 

Walker, F. 1849. List of the specimens of dipterous 
insects in the collection of the British Museum. 
Vol. 4. pp. 689-1 172. London. 


105(3). 2003. pp. lib-Ill 


A New Synonym of CuiuUitclUi hystrix (Traver) 
(Ephemeroptera: Ephemerellidae) 

Caiidatella Edmunds (Ephemeroptera: 
Ephemerellidae) was described as a Nearc- 
lic subgenus of Ephemerella Walsh (Ed- 
munds 1959) for species included in 
McDunnough's (1935) heterocaudata spe- 
cies group. Allen (1980) later raised Caii- 
clatellu to generic status. Allen and Ed- 
munds (1961) constructed a Caiidatella 
species key. in which specimens with well- 
detined ventral color patterns would "key 
with ease"; however, specimens with less 
distinct \entral markings would "key 
llnough a more difhcult series of couplets." 

Allen and Edmunds ( 1961) differentiated 
C. cascadia (Allen and Edmunds) and C. 
hystrix (Traver) (two species with overlap- 
ping geographic distribution) by ventral ab- 
dominal coloration, the respective presence 
or absence of occipital tubercles, and the 
relative development of abdominal tuber- 
cles. Examination of series of C. hystri.x lar- 
vae, C. cascadia type material, and addi- 
tional material from the C. cascadia type 
locale revealed that variability of occipital 
and abdominal tubercles development and 
ventral color pattern in C. hystri.x encom- 
passes the concept of C. cascadia. Some 
individuals from populations of C. hystri.x 
have occipital tubercles such as described 
for C. cascadia. Additionally, we have 
found that the ventral color patterns asso- 
ciated with C. cascadia and C. hystri.x are 
variable within single populations and have 
overlapping development. Therefore, we 
propose the following new subjective syn- 
onymy: C. hystri.x (Traver) 1934 [= C. cas- 
cadia (Allen and Edmunds) 1961, new syn- 
onym]. Previously, Day (1954) recognized 
Ephemerella spinosa Mayo (nee E. spitwsa 
Morgan, nee E. spinosa Ikonomov) as a 
synonym of C. hystri.x. 

All material examined is housed in the 

Purdue University Entomological Research 
Collection. West Lafayette. Indiana. George 
E Edmunds. Jr (Salt Lake City. Utah) and 
Bob Wissemann (Corvallis. Oregon) donat- 
ed material for study. This study was fund- 
ed in part by USEPA fellowship 91601701- 

to LMJ and NSF grant DEB-9901577 to 

Material examined. — IDAHO: Lemhi 
Co., Opal Cr, 23-VII-1964, three larvae, 
one set larval exuviae. MONTANA: Judith 
Basin Co., Running Wolf Cr, 47°00'33"N, 

1 10°22'18"W, 12-VI-200(), WP McCafferty 
et al.. 25 larvae; Ravalli Co., Bitterroot R.. 
N. of Sula on US93. AR GauHn. 29-VI- 
1963. one larva; E. Fk. Bittenoot R.. 10 mi. 
above jet. W. Fk. Bittenoot R., 24-VI-1965, 
JR Grierson. one larva. NEVADA: Washoe 
Co., Incline Cr. 2 mi. W. Mt. Rose Summit. 
GF Edmunds. Jn & RK Allen, 21 -IX- 1957, 
six immature larvae. OREGON: Clackamas 
Co.. Branch of Still Cr. on rd. to Timberline 
Lodge. Mt. Hood, 30-VIII-1958, GF Ed- 
munds, Jr. & RK Allen, 18 larvae (C. cas- 
cadia holotype, paratopotype, and topo- 
types); Douglas Co., Jackson Cr headwater 
trib., Umpqua NE 43°03'N, 122°30'W, 14- 
VI- 1990. RW Wissemann. four larvae; 
Klamath Co.. Sun Cr., Crater Lake NR 14/ 
22-IX-1993. 10 larvae; Linn Co., Sweet- 
water Cr. Willamette NF. Rigdon Ranger 
District. IX- 1997, three larvae. WYO- 
MING: Park Co.. Antelope Cr. Yellow- 
stone NP 17-VI-1949. GF Edmunds. Jr. 
one larva; Gibbon R. at Virginia Cascade, 
I2-VM963, ER Vincent, 15 larvae; Iron 
Cr. YNR 21-VM949. GF Edmunds, Jr, 
one larva; Teton Co.. Firehole R., Old 
Faithful, YNR 6-V1-1942, GF Edmunds. Jr, 
one larva. 

Literature Cited 

.Mien. R. K. 1980. Geographic distribulion and reclas- 
sitication of the subfamily Ephemerellinae 



( Ephemeroptera: Ephemerellidae). pp. 71-91. In 
Flannagan, J. F. and K. E. Marshall, eds. Advances 
in Ephemeroptera Biology. Plenum, New York. 

Allen, R. K. and G. E Edmunds. Jr. 1961. A revision 
of the genus Epheinerella (Ephemeroptera: 
Ephemerellidae). II. The subgenus Caiuiatel/a. 
Annals of the Entomological Society of America 
54: 603-612. 

Day, W. C. 1954: New species and notes on California 
mayflies. II. (Ephemeroptera). The Pan-Pacific 
Entomologist 30: 15-29. 

Edmunds. G. F, Jr. 1959. Subgeneric groups within the 

mayfly genus Epheinerella (Ephemeroptera: 
Ephemerellidae). Annals of the Entomological So- 
ciety of America 52: 543-547. 
McDunnough, J. 1935. Notes on western species of 
Ephemeroptera. The Canadian Entomologist 67: 

Luke M. Jacobus and W. P. McCafferty, 
Department of Entowology. Purdue Uni- 
versity, West Lafayette. /N 47907. U.S.A. 

105(3). 2003. pp. 778-780 


Food Plant, Life History Notes, and Distribution of Nenuitits atriceps (Marlatt) 
(Hymenoptera: Tenthredinidae) 

Sawfly larvae were discovered feeding 
on Trifolium wonnskjolkii Lenm. (Legu- 
minosae) by GP at Seal Rock, Lincoln Co., 
Oregon, in the summer and fall of 2002. 
The reared adults were Nematus atriceps 
(Marlatt) (Nematinae) which occurs in the 
western United States and Canada. This is 
the first food plant record for this sawfly 
and the first record for any of the species 
placed in the Nematus atriceps group by 
Smith ( 1979). The food plant is unusual for 
Nematus in that most species feed on Sali- 
caceae, Betulaceae, or other woody trees or 
shrubs. One Palearctic species, Nematus 
myosotidis (E), is known to feed on Trifo- 
lium sp. (Taeger et al. 1998). Nematus myo- 
sotidis is very similar to N. atriceps, and 
could be placed in the same species group. 

At Seal Rock, the clover plants were 
growing in a seepage area about 12 feet 
from the high tide level. Nematus larvae 
(Fig. 1) were first seen and collected while 
feeding on the flowers on July 12, 2002. 
Some lepidopteran larvae were found in 
other flower heads at the same site. On re- 
turning to the site on August 14, 2002, no 
insects were found on the same clover 
plants. However, on September 6, a number 
of small Nematus larvae were found eating 
clover leaves but leaving the midrib and 
larger veins. Three sawfly larvae were col- 
lected on September 23, 2002, from the 
leaves, and, on October 7, 2002, three ad- 
ditional small larvae were found on the 
leaves. After September, flowering drops 
off considerably, so the leaves are the major 
source of nourishment. Young larvae were 
reared in the laboratory on flowers and 
leaves of the same clover species. They 
would not eat white clover, Trifolium re- 
pens L. In rearing, they readily spun a 
tough, papery cocoon on the flowers (Fig. 

3), or, if no flowers were present, under the 
filter pad in the petri dish. If the mature 
larvae are continually moved, they often 
did not form a cocoon and pupated naked 
(Fig. 2). Adults (Fig. 4) emerged in about 
7 to 10 days. In the field, pupae were found 
only on flowers; however, they could have 
been in the duff even though it was quite 
wet under the plants. 

The food plant is a native species com- 
monly called marsh or spring bank clover. 
It is distributed from British Columbia to 
Mexico along the coast and also has inland 
populations in Idaho, New Mexico, and 
Colorado. It is a weak-stemmed perennial 
with large red. pink, or purple flowers. 
There is a mountain meadow race found 
along streams, as well as a coastal race 
found on beaches or edges of salt marshes. 
The coastal race is a rhizomed, matted form 
(Hickman 1996). Populations of the coastal 
race were examined by GP all along 
Oregon and into northern California (Lake 
Earl). The only locality where Nematus was 
found was at Seal Rock. The sawfly may 
have been missed at the other sites, al- 
though other insects were found in the 
flowers, such as larvae of Hypero punctata 
(¥.). an Old World weevil. 

Though Nematus atriceps feeds both on 
the leaves and inflorescence of Trifolium 
wormskjoldii, few other sawflies have been 
recorded feeding on inflorescences and N. 
atriceps provides one of the few examples. 
Other North American species include Ten- 
thredo sp. on the flowers of Ranunculus 
californicus Benth. (Ranunculaceae) in Cal- 
ifornia (Linsley and MacSwain 1959), and 
Rhadinoceraea zigadenusae Smith (Smith 
and McDearman 1990) and R. sodsensis 
Smith and Barrows (1995) on species of Z/- 
gadenus (Liliaceae) in southeastern U.S. 
and West Virginia. 


Figs. 1-4. Nematiis atriceps. 1. Mature larva feeding on a leaflet of Trifolium: length. 13 mm. 2, Pupa, 
without cocoon; length 8 mm. 3. Cocoon attached to florets of the clover flower; the larva developed on the 
petals; length. 1 1 mm. 4. Recently emerged adult, length. 12 mm. 

The distribution of Neinatiis atriceps has 
not been recorded. It is found throughout 
western United States and in western Can- 
ada from the Rockies westward with sev- 
eral records east to Wisconsin and Mani- 
toba. It generally overlaps the distribution 
of the food plant, but it is also possible N. 
atriceps feeds on other species of clover 
Material examined by DRS is as follows: 
ALASKA: Nazan Bay. Atka. Aleutian Is.. 
VII-27-1907, VIII-1-1907. ALBERTA: 
Beaverlodge. VI-7-1931; Wabamum. VII- 
31-1929. BRITISH COLUMBIA: "Vane." 
CALIFORNIA: Smith River, VII- 17: Rat- 
tlesnake Mdw.. Siskiyou Co., 5800'. VIII- 
9-1970; Twin Lake. Siskiyou Co.. VII-27- 
1971; Young's Valley, Siskiyou Co., 4600', 
VII-4-1971; Swift Cr. Trinity Co.. 5700'. 
VII-31-1973: Black Bsn. Trinity Co., 7100'. 
VII-29-1 972: Humboldt Co.: Mumford 
Bsn, Trinity Co, 6400', VII-3I-1972: 7 mi 
S Yellowjacket Camp, El Dorado Co.. VI- 

29-1980; Cracker Mdws.. Siskiyou Co., 
5000', VIII-16-1971; Cazadero. IV-12- 
1918: Oakland, Alameda Co., III-22-1952; 
Strawberry, Tuolumne Co.. VI-2 1-1951; 
Tahquitz Mdw. San Jacinto Mts.. VI-3- 
1940: Buck's Lake. Plumas Co., VII- 1- 
1949: Muir Woods, V-19-1915; Cona Cr. 
Napa Co., IV-23-1949: 3 mi E. Mt. Lassen, 
VII-19-1953: Trinity Co., V-25-1934: Sa- 
gehen nr Hobart Mills, VII-9-1954, VII-25- 
1954; Woodacre. III-28-1955; Pine Crest, 
Tuolumne Co.. VII- 1-1951; Kings Crk. 
Mdw.. Shasta Co., VII-2-1947. COLORA- 
DO: "Colo."': 15 mi N Jet R. 14 & 40, VII- 
2-1962; Pitkin Co.. Weller. 9360'. 8 rd. mi 
SE Aspen. VII-26-1977: Big Spring Ranch. 
Florissant. VII- 18- 1962. IDAHO; Lenore. 
V-7-1938; Weippe. 3006'. VII-5-1935; 
Vollmer. V-1930. MANITOBA: Aweme, 
VI- 10-1928. MONTANA: Fairy Lakes, 
VIII-IO-1968. NEVADA: "Nev " (holo- 
type of atriceps): "Ormsby Co.," VII. 


NEW MEXICO: Magdalena Mts.. VIII- 
1894. NORTH DAKOTA: Bottineau Co.. 
Malaise trap, VII- 17- 1 972, VII-28-1974. 
OREGON: Seal Rock, Lincoln Co.. 2002 
(reared specimens); Cannon Beach. VIII-9- 
1940: Wheeler Co., road to Grant Spg., 500 
yds SW Guard Station, VII-20-I962: Wal- 
lowa Co.. 8 mi N Flora, VI-7-1963; Iron 
Mtn., 8 mi E Upper Soda, Linn Co., 5000', 
VIII-1 1-1962: Kelsey Valley, Douglas Co., 
VI-26-1962: Horse Lake, High Cascade 
Mountains, Lane Co, VII-25-3 1-1909: 
Jackson Co., Squaw Lake, 7 mi E Copper. 
V- 19- 1962; Jackson Co., Dead Indian Soda 
Springs, 12 mi SE Lakecreek, 2500', V-21- 
1964; Jackson Co., Pinehurst. 3375'. V-19- 
1960: Jackson Co., Buckhorn Mineral 
Spgs., 1 1 mi ESE Ashland, 2800', Emigrant 
Cn, V- 19- 1960; Benton Co., Rock Creek, 4 
mi S Philomath, IV-28-1963; Wasco Co.. 
Bear Springs. 3180', 25 mi W. Maupin, V- 
21-1959. WASHINGTON: Mt. Adams, VI- 
26-1931; Westport, VIM 7- 1949; Seattle, 
IX- 17- 1891. WISCONSIN: Polk Co., VII. 
UTAH: Bear R., n. si. Uinta Mts., 8000'. 

Acknowledgments. — Some specimens of 
the material examined are from the Cana- 
dian National Collection. Ottawa. Univer- 
sity of California at Berkeley. Cornell Uni- 
versity, and the Natural History Museum of 
Los Angeles County. We extend thanks to 
the curators for allowing examination of 
specimens. Thanks to N. Schiff, USDA, 
Forest Service, Stoneville, MS, and S. Cod- 

ella. Kean University, Union, NJ, for re- 
viewing the manuscript. 

Literature Cited 

Hickman. J. C, ed. 1996. The Jepson Manual. Llni- 
versity of California Press. Berkeley. 1 ,400 pp. 

Linsley. E. G. and J. W. MacSwain. 1959. Ethology of 
some Ranunculus insects with emphasis on com- 
petition for pollen. University of California Pub- 
lications in Entomology 16: 1-46. 

Smith. D. R. 1979. Symphyta. pp. 3-137. //; Krom- 
bein. K. V., R D. Hurd. Jr., D. R. Smith, and B. 
D. Burks, eds. Catalog of Hymenoptera in Amer- 
ica North of Mexico, Vol. 1. Smithsonian Insti- 
tution Press, Washington, DC. 

Smith. D. R. and E. M. Barrows. 1995. Rhadinoceraea 
n. sp. (Hymenoptera: Tenthredinidae) from West 
Virginia, a second species on Zigadenus (Lili- 
aceae). Entomological News 106: 237-240. 

Smith. D. R. and W. McDearman. 1990. A new Rhad- 
iniiceraea (Hymenoptera: Tenthredinidae) feeding 
on Zigadenus (Liliaceae) from southeastern Unit- 
ed States. Entomological News 101: 13-19. 

Taeger, A., E. Altenhofer, S. M. Blank. E. Jansen, M. 
Kraus, H. Pschron-Walcher. and C. Ritzau. 1998. 
Konimentare zur Biologic. Verbreitung und Ge- 
fahrdung der Pflanzenwespen Deutschlands (Hy- 
menoptera, Symphyta). pp. 49-135. In Taeger. A. 
and S. M. Blank, eds. Pflanzenwespen Deutsch- 
lands (Hymenoptera. Symphyta). Verlag Goecke 
& Evers. Keltern, 364 pp. 

George Poinar, Jr., Department of Zool- 
ogy. Oregon State University. Con'allis. 
OR 97331. U.S.A. {e-mail: poinarg@bcc. and David R. Smith, Systematic 
Entomology Laboratory. PSI. Agricultural 
Research Senice. U.S. Department of Ag- 
riculture. % National Museum of Natural 
History, Smithsonian Institution. Washing- 
ton. DC 20560-0168. U.S.A. (e-mail: 


105(3). 2003. pp. 781-782 


Notes on the Biology of Syna.xis forniosa (Hiilst) (Lepidoptera: Geonietridae) in 
South Central Washington State 

Scoble (1999) recorded ten species of 
Syna.xis Hulst from North America while 
McGuffin (1987) discussed four from Can- 
ada. McGuffin (1987) indicated that several 
of the species fly late in the season and 
probably overwinter in the egg stage; the 
larvae appear in the spring, feeding on co- 
nifers and hardwoods. For three of the Ca- 
nadian species, he referenced significant 
host data along with flight periods. One 
species, Syna.xis formoso (Hulst). had un- 
known immature stages and no host infor- 
mation. McGuffin indicated the flight peri- 
od as October and recorded the species only 
from British Columbia. Hulst (1896) de- 
scribed the species from Colorado and 
southern California. 

Since 1995, we have been conducting a 
survey of the biological diversity of select 
groups of arthropods on the Hanford Site 
located in south central Washington State. 
Consult Zack (1998) and Zack et al. (1998) 
for a description of the Hanford Site and 
the results of other aspects of the overall 
project. One of the groups of primary in- 
terest has been the moth fauna. Herein we 
report on the life history, host plant, and 
light trap captures of S. formoso. 

Larvae of S. formoso were collected at 
the Chamna Natural Area (46°15.15'N 
119°16.85'W) located south of Richland 
(Benton Co.) on 31 March 2001. This site is 
not within the Hanford Site but lies along 
the Yakima River just outside the boundaries 
of the Site. The habitat contains significant 
stands of big sagebrush (Artemesia triden- 
tata Nuttall) bordering the riparian habitat 
along the river. Larvae were taken by beat- 
ing big sagebrush plants and observing ma- 
terials falling to sheets placed under the 
plants. Two geometrid larvae less than 1 cm 
in length (probably second instars) were col- 

lected; they appeared to be conspecific. Both 
larvae were placed on a potted sagebrush ta- 
bling and protected by placing a transparent 
plastic cover, with screened top, over the 
plant. The plant was kept outdoors, but out 
of direct sunlight to protect from overheat- 
ing. One of the larvae died before attaining 
1 .3 cm, but the second survived, pupated, 
and emerged as an adult on 6 October 200 1 . 
The larva is a twig mimic as was noted by 
McGuffin ( 1 987 ) for other species of Syn- 
a.xis. The larva had pupated by mid-May and 
remained dormant throughout the summer. 
The pupal period was thus 130-140 days, 
much longer than the time period suggested 
by McGuffin (1987) for Syna.xis jubararia 
Hulst (58-64 days). This longer pupal peri- 
od may reflect the relatively long hot and 
dry summer period found in the south cen- 
tral Columbia Basin of eastern Washington 
where these observations were made. The 
reared adult was slightly smaller than wild 
caught individuals. 

Adults were collected at several locations 
on the Hanford Site at 15-watt "black 
light"" or 150-watt mercury vapor light. 
Collecting locations and dates are as fol- 
lows: Benton Co. (all Hanford Site), SE 
base of Gable Mtn., 46°35.286'N 
119^27. 868'W, 15 Oct 1997 (5); Rattle- 
snake Spring, 46°30.447'N 1 19°41.887'W, 
18 Oct 1996 (3) and 20 Oct 2001 (1); sand 
dunes, 46°31.369'N 1 19°21.192'W, 9 Oct 
1996 (5) and 23 Oct 1996 (1). Grant Co. 
(all Hanford Reach National Monument — 
Saddle Mountain National Wildlife Refuge), 
46°42.155'N 1 19°37.230'W, 18 Oct 2002 
(3); 46°42.064'N 119°38.27rw, 18 Oct 
2002 (3); 46°42.117'N 119°36.282'W. 18 
Oct 2002 (1); 46°42.064'N 1 19°38.271'W. 5 
Oct 2002 ( 14) and 25 Oct 2002 ( 1 ). 

This work was funded by The Nature 


Conservancy and the U.S. Department of 
Energy. The U.S. Fish and Wildlife Service 
allowed us access to sites on the Hanford 
Reach National Monument and was sup- 
portive of our studies. 

Literature Cited 

Hulst. G. D. 1896. A classification of the Geometrina 
of North America with descriptions of new genera 
and species. Transactions of the American Ento- 
mological Society 23: 245-348. 

McGuffin. W. C. 1987. Guide to the Geometridae of 
Canada (Lepidoptera). II. Subfamily Ennominae. 
4. Memoirs of the Entomological Society of Can- 
ada 138: 1-182. 

Scoble. M. J. 1999. Geometrid Moths of the World: A 
Catalogue (Lepidoptera. Geometridae). Volume 2. 
CSIRO Publishing. CoIIingwood. Australia. 

Zack. R. S. 1998. Shore flies (Diptera: Ephydridae) of 
the Hanford Site. Washington. Northwest Science 
72: 127-141. 

Zack. R. S., N. D. Penny, J. B. Johnson, and D. L. 
Strenge. 1998. Raphidioptera and Neuroptera 
from the Hanford Site of southcentral Washington 
State. Pan-Pacific 74: 203-209. 

Dennis L. Strenge. BatteUe, Pacific 
Northwest Division, Richland, WA 99352, 
U.S.A. (e-mail: and 
Richard S. Zack. Department of Entomol- 
ogy, James Entomological Collection, 
Washington State University, Pullman, WA 
99164-6382. U.S.A. (e-mail: zack@wsu. 

105(3). 2003. pp. 783-785 


Contiimation of the Presence o\' Amhlyoninici ovale Koch 1844 and First Records of 
Ainhlyoiuina scalpuiniium Neumann 1906 (Acari: Ixodida: Ixodidae) in the Amazonian 

Region of Ecuador 

The senior author visited the Centro de 
Rescate Amazoonico (OTOS'S. 77°32'W). 
Napo Province, in the Amazonian region of 
Ecuador in February. 2002. The following 
species of vertebrates were inspected re- 
peatedly for ticks during that month: do- 
mestic dog (Carnivora: Canidae); Nasiia 
nasiui (L. 1766) (Carnivora: Procyonidae); 
Tapirus terrestris (L. 1758) (Perissodactyla: 
Tapiridae); monkeys (Primates: Cebidae), 
Saimiri sciiireus (L. 1758), Cebus capuci- 
nus (L. 1758) and Lagothrix lagothcha 
(Humboldt 1812). Tick specimens obtained 
were preserved in 70% ethanol and depos- 
ited in the Tick Collection of Estacion Ex- 
perimental Agropecuaria Rafaela, Instituto 
Nacional de Tecnologi'a Agropecuaria, Ra- 
faela, Santa Fe, Argentina (INTA). 

No ticks were found on monkeys. A 
nymph of Amblyomma spp. was found on 
a N. nasua (INTA N° 1727). Amblyomma 
ovale Koch 1844 was collected from dogs, 
tapirs and also a human, and Amblyomma 
scalpttiratum Neumann 1906 was found on 
a tapir (Table 1 ). Amblyomma ovale was 
identified according to Aragao and Fonseca 
(1961b), and A. scalptiiratum following 
Boero and Prosen (1960). One of us (JEK) 
searched the U. S. National Tick Collection 
for additional specimens of both tick spe- 
cies from Ecuador. The search yielded sev- 
eral records, all from the Amazonian region 
of Ecuador (Table 1 ). 

Amblyomma ovale is widespread in the 
Neotropical Region, reaching the southern 
Nearctic (Guglielmone et al.. in press). 
However, its presence in Ecuador has been 
documented only twice. Becklund (1968) 
found a specimen on an ocelot imported to 
U.S.A. from Ecuador, and Guglielmone et 
al. (in press) cites its presence in the coun- 

try. Adults of this tick species primarily in- 
fest carnivores, but findings on tapir and 
man are not infrequent (Guglielmone et al., 
in press), as reflected in the records from 

Amblyomma scalpturatum is usually 
found on tapirs and to a lesser extent on 
anteaters and dogs in Bolivia, Brazil, Co- 
lombia, French Guiana, Guyana, Peru and 
Venezuela (Jones et al. 1972). Evidentally, 
A. scalpturatum can occasionally parasitize 
humans (Aragao and Fonseca 1961a). 
These are the first records of A. scalptura- 
tum from Ecuador, and expand the known 
distribution of this poorly known tick spe- 

We acknowledge the collaboration of 
Angelika Raimann, Director of Centro de 
Rescate Amazoonico, with the work carried 
out by CZ. We also acknowledge the sup- 
port of National Institute of Allergy and In- 
fectious Diseases grant Al 40729 to JEK, 
and INTA and Fundacion ArgenlNTA to 

Literature Cited 

Aragao, H. de B. and F da Fonseca. 1961a. Notas de 
ixodologia. VIII. Li.sta e chave para os represen- 
tantes da fauna ixodologica brasileira. Memdrias 
do Instituto Oswaldo Cioiz 59: 115-130 + 27 

. 1961b. Notas de ixodologia. IX. O complexo 

ovale do genera Amblyomma. Memorias do Insti- 
ttito Oswaldo Craz 59: 131-148 + 4 plates. 

Becklund. W. W. 1968. Ticks of veterinary significance 
found on imports in the United States. Journal of 
Parasitology 54: 622-628. 

Boero. J. J. and A. F. Prosen. I960. Ixodideos de 
"Anla." III. Amblyomma scalpturatum Neumann 
1906. Descripcion del macho y redescripcion del 
henibra. Anales del Instituto de Medicina Region- 
al 5: 115-119. 

Guglielmone, A. A.. A. Estrada-Peiia, A. J. Mangold. 
D M. Barros-Battesti, M. B. l.abruna, ,1. R. Mar- 


























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tins, J. M. Venzal. M. Araza. and J. E. Keirans. 
In press. Ambtyomma aiireolatum (Pallas. 1772) 
and Amblyomma ovale Koch. 1X44 (Acari: Ixod- 
idae): DNA sequences and dislributions. Veteri- 
nary Parasitology. 
Jones. E. K.. C. M. Clifford. J. E. Keirans, and G. M. 
Kohls. 1972. The ticks of Venezuela (Acarina: l,x- 
odoidea) with a key to the species of Ainhlyorninu 
in the Western Hemisphere. Brigham Young Uni- 
versity Science Bulletin, Biological Series 17: 1- 

Corina Zerpa, Cdtedra de Higiene, Epi- 
demiologi'a y Administnicion Sanitaria. Fa- 
cidtad de Ciencias Veterinarias, Universi- 
dad Nacional de Rosario. Ovidido Lagos y 

nita 33. CP 2170 Casilda. Santa Fe, Ar- 
gentina {e-mail: corina.zerpa@hotmail. 
com). James E. Keirans, U.S. National Tick 
Collection. Institute of Arthropodology and 
Parasitology. Georgia Southern University. 
Stateshoro. GA 30460-8056. U.S.A. (e- 
mail: Atilio J. Man- 
gold and Alberto A. Guglielmone, Instituto 
Nacional de Tecnologi'a Agropecuaria. Es- 
tacion E.xperimental Agropecuaria Rafaela. 
Casilla de Correo 22. CP 2300 Rafaela, 
Santa Fe. Argentina (e-mails: amangold® 
rafaela. inta. gov. ar: agugliclmone@ rafaela. 

105(3). 2003. pp. 786-788 


Phylogenetics and the Reconfirmation of Dentatella Allen 
(Ephemeroptera: Ephemerellidae) 

The history of Ephemeroptera taxonomy 
has involved instances of one species de- 
scribed as an adult and another species de- 
scribed as a larva proving to be the same 
.species once rearing associations of the two 
stages have taken place. For some of the 
more recent examples establishing such as- 
sociations, see Whiting and Lehmkuhl 
(1987), McCafferty and Provonsha (1988), 
McCafferty and Silldorff (1998), and Bur- 
ian (2002). Lugo-Ortiz and McCafferty 
(1996a) argued that some such associations 
were inevitable but because of the present 
imperative to document the biodiversity of 
the world, the possibility of such eventual 
associations should not preclude the de- 
scription of species based on either stage. 
In a recent instance (Burian 2002), adults 
of Eiirylophella coxalis (McDunnough) 
(previously unknown as larvae) were 
shown to be associated with the known lar- 
vae of the genus Dentatella Allen, as both 
D. bartoni (Allen) and D. dmiutae Mc- 
Cafferty (previously unknown as adults). 
DeutateUci and the more speciose genus Eu- 
rylophella Tiensuu had previously been 
shown to be sister branches within the 
Ephemerellidae subfamily Timpanoginae 
(McCafferty and Wang 1994. 2000). Buri- 
an's placement of the species in Eiirylo- 
phella and dissolution of Dentatella, how- 
ever, discounted compelling cladistic evi- 
dence (McCafferty 1977, 1978, 2000; 
McCafferty and Wang 1994, 2000) that, 
within a framework of strict phylogenetic 
hierarchical classification, allows this spe- 
cies to reside in a separate genus. 

To support his position, Burian (2002) 
proposed an arbitrary rule that both adult 
and larval stages of lineages must possess 
defining morphological apomorphies in or- 
der to be recognized at the senus level. This 

effectively eliminated the genus Dentatella 
because its divergence with Eurylophella is 
based on larval apomorphies. We reject 
Burian's rule because it is both unrealistic 
and unnecessary to phylogenetic systemat- 
ics, and would prove devastating to the 
higher classifications of metamorphic or- 
ganisms. For example, we estimate that 
Burian's rule would eliminate well over half 
of the mayfly genera, by discounting genera 
that are now based on phenetic data and 
thus taken as hypothetical groupings yet to 
be tested with cladistics, and by discounting 
genera now unknown in one or the other 
stage, including nearly all extinct genera. A 
consideration of groups that have under- 
gone cladistic analysis [for example, in the 
Baetodes complex (McCafferty and Baum- 
gardner 2003), the Biigilliesia complex 
(Lugo-Ortiz and McCafferty 1996b), the 
Hennanella complex (Flowers and Domin- 
guez 1991), the Mirocuhis complex (Sav- 
age and Peters 1982), Neoephemeridae 
(Bae and McCafferty 1998), Potamanthidae 
(Bae and McCafferty 1991 ), Teloganodidae 
(McCafferty and Wang 1997, McCafferty 
and Benstead 2002), and Timpanoginae 
(McCafferty and Wang 1994)] indicates 
that a combination of both adult and larval 
synapomorphies or autapomorphies does 
not exist at the point of generic branching 
in a large proportion of genera. Such does 
not even exist at more basal branchings in- 
volving several families and higher taxa of 
mayflies. For example, a single adult syn- 
apomoiphy common to the Leptohyphidae 
is the basis of that taxon being recognized 
as a non-paraphyletic family separate from 
Coryphoridae (Molineri et al. 2001). Even 
if Burian intended to qualify his rule by 
limiting it only to monospecific genera [the 
largest category of genera in all biota (see 


e.g., Raup 1991)], his rule would still dec- 
imate higher classifications and undermine 
their potential applications. 

Monophyly and branching sequence are 
the essential bases for phylogenetic higher 
classifications (e.g., Hennig 1966). with no 
specification of what life stage or multiple 
life stages of a lineage must contribute the 
apomorphies supporting hypotheses of 
monophyly and no specification of the de- 
gree of synapomorphy required at any 
branch for deHning taxonomic categories. 
Because rates of morphological evolution 
vary considerably between larval and adult 
stages of mayflies (e.g.. see McCaffeily and 
Edmunds 1976. fig. 1) and other metamor- 
phic organisms when different selection en- 
vironments are involved, there is no logical 
reason to expect moiphological apomor- 
phies to be expressed in multiple life stages 
of every lineage and clade. These life stages 
are "'character-bearing semaphoronts," the 
basic comparable elements of biology, in 
the terms of Hennig (1966), and any one 
semaphoront of an evolving lineage may 
sufficiently demonstrate morphological 
character evolution for the purposes of phy- 
logenetics. Burian's rule, insisting on lay- 
ered evidence from both adults and larvae 
for a particular taxonomic category is tan- 
tamount to an arbitrary gap criterion. 
McCafferty (1991) rejected the use of all 
gap criteria, following Wiley's (1981) ad- 
monition that phylogenetic classifiers must 
reject gaps and definitions of taxonomic 
categories based on such criteria. Finally 
and not in the least, imposition of Burian's 
rule and the resultant severe reduction of 
genera would significantly lessen the valu- 
able indications of comparative biology (the 
"explanatory powers") that are inherent in 
the complex hierarchies of phylogenetic 
classifications (Ross 1974. Farris 1979, Wi- 
ley 1981, Ax 1987). 

In keeping with a philosophy of provid- 
ing strictly phylogenetic classifications of 
Ephemeroptera when possible, with a max- 
imum of information content and uncon- 
strained by gap or special criteria, or any 

selective use of such, we are regarding the 
above treated recently associated species as 
DentateUa co.xalis (new combination), and 
thus continue to recognize the cladistic- 
founded genus DentateUa — a highly dis- 
tinctive taxon in the larval stage and sister 
lineage to Euiylophella. These sister genera 
are phylogenetically distinct within the tribe 
Eurylophellini of the subfamily Timpano- 
ginae. In the case of DentateUa. for exam- 
ple, the broadened larval femur is autapo- 
moiphic within the subfamily, and the rel- 
atively enlarged size of operculate gill 4, 
covering much of abdominal tergum 8 is 
autapomorphic within the order. For Euiy- 
lophella. the much elongated larval abdom- 
inal tergum 9 is a unique synapomorphy 
within the order. We know of no evidence, 
out-group or otherwise, that suggests alter- 
native polarities with respect to these char- 

Literature Cited 

Ax. P. 1987. The phylogenetic system; The systemi- 
zation of organisms on the basis of their phylo- 
genesis. Wiley, New York. 

Bae. Y. J. and W. P. McCafferty. 1991. Phylogenetic 
systematics of the Potamanthidae (Ephemerop- 
tera). Transactions of the American Entomological 
Society 117: 1-143. 

. 1998. Phylogenetic systematics and biogeog- 

raphy of the Neoephemeridae (Ephemeroptera: 
Pannota). Aquatic Insects 20: 35-68. 

Biirian. S. K. 2002. Taxonomy of Eurylophella co.xalis 
(McDunnough) with notes on larval habitat and 
behavior (Ephemeroptera: Ephemerellidae). Jour- 
nal of the North American Benthological Society 
21: 602-615. 

Farris. J. S. 1979. The information content of the phy- 
logenetic system. Systematic Zoology 28: 483- 

Flowers. R. W. and E. Dominguez. 1991. Preliminary 
cladistics of the Hermcmella complex (Ephemer- 
optera: Leptophlebiidae. Atalophlebiinae). pp. 49- 
62. In Alba-Tercedor, J. and A. Sanchez-Ortega. 
eds. Overview and strategies of Ephemeroptera 
and Plecoptera. Sandhill Crane Press. Gainesville. 

Hennig, W. 1966. Phylogenetic systematics. University 
of Illinois Press, Urbana. 

Lugo-Ortiz. C. R. and W. R McCafferty. 1996a. New 
species of Leptophlebiidae (Ephemeroptera) from 
Mexico and Central America. Annales de Lim- 
nologie 32: 3-18. 

. 1996b. The Bugilliesia complex of African 



Baetidae (Ephemeroptera). Transactions of the 
American Entomological Society 122: 1 7?- 1 97. 

McCafferty. W. P. 1977. Biosystematics of Dannelia 
and related subgenera of Ephemerella (Ephemer- 
optera: Ephemerellidae). Annals of the Entomo- 
logical Society of America 70: 881-889. 

. 1978. A natural subgeneric classification of 

Ephemerella bartoni and related species (Ephem- 
eroptera: Ephemerellidae). The Great Lakes En- 
tomologist 11: 137-138. 

. 1991. Toward a phylogenetic classification of 

the Ephemeroptera (Insecta): A commentary on 
systematics. Annals of the Entomological Society 
of America 84: 343-360. 

. 2000. A hierarchical classification of the Tim- 

panoginae (Ephemeroptera: Ephemerellidae) and 
description of a new species from Quebec. An- 
nales de Limnologie 36; 157-161. 

McCafferty. W. P and D. E. Baumgardner. 2003. Lii- 
goiops maya. a new genus and species of Ephem- 
eroptera (Baetidae) from Central America. Pro- 
ceedings of the Entomological Society of Wash- 
ington 105: 397-406. 

McCaffeny. W. P and J. R Benstead. 2002. Cladistic 
resolution and ecology of the Madagascar genus 
Maiiohypliella Allen (Ephemeroptera: Teloganod- 
idae). Annales de Limnologie 38: 41-52. 

McCafferty, W. P and G. F Edmunds. Jr 1976. Re- 
definition of the family Palingeniidae and its im- 
plications for the higher classification of Ephem- 
eroptera. Annals of the Entomological Society of 
America 69: 486-490. 

McCafferty. W. P and A. V. Provonsha. 1988. Revi- 
sionary notes on predaceous Heptageniidae based 
on larval and adult associations (Ephemeroptera). 
The Great Lakes Entomologist 21: 15-17. 

McCafferty. W. P and E. L. Silldorff. 1998. Reared 
association and equivalency of Badis adnnis and 
B. caelestis (Ephemeroptera: Baetidae). Entomo- 
logical News 109: 261-265. 

McCafferty. W. R and T.-Q. Wang. 1994. Phylogenet- 
ics and classification of the Timpanoga complex 

(Ephemeroptera: Ephemerellidae). Journal of the 
North American Benthological Society 13: 569- 

. 1997. Phylogenetic systematics of the family 

Teloganodidae (Ephemeroptera: Pannota). Annals 
of the Cape Provincial Museums (Natural History) 
19: 387-437. 

. 2000. Phylogenetic systematics of the major 

lineages of pannote mayflies (Ephemeroptera: 
Pannota). Transactions of the American Entomo- 
logical Society 126: 9-101. 

Molineri, C J. G. Peters, and M. Zuninga de Cardoso. 
2001. A new family. Coryphoridae (Ephemerop- 
tera: Ephemerelloidea). and description of the 
winged and egg stages of Coi-yphonis. Insecta 
Mundi 15(2): 1-6. 

Raup. D. M. 1991. Extinction: Bad genes or bad luck? 
W. W. Norton. New York. 

Ross. H. H. 1974. Biological systematics. Addison- 
Wesley. Reading. Massachusetts. 

Savage. H. M. and W. L. Peters. 1982. Systematics of 
Miroculiis and related genera from northern South 
America (Ephemeroptera: Leptophlebiidae). 
Transactions of the American Entomological So- 
ciety 108: 491-600. 

Whiting. E. R. and D. M. Lehmkuhl. 1987. Raptohep- 
tagenia cruentala, gen. nov. (Ephemeroptera: 
Heptageniidae). new association of the larva of 
previously thought to be Anepennts with the 
adults of HepUigeniii cnientara Walsh. The Ca- 
nadian Entomologist 1 19: 405-407. 

Wiley. E. O. 1981. Phylogenetics: The theory and 
practice of phylogenetic systematics. Wiley. New 

W. p. McCafferty, Luke M. Jacobus. De- 
partment of Entomology. Purdue Universi- 
ty. West Lafayette. IN 47907. U.S.A (e-mail:, and T- 
Q. Wang, 8 Copely Dr.. Andover. MA 
01810. U.S.A. 

105(3), 2003, pp. 789-791 


First Report of Nuptial Feeding in Sawflies, Aneiignwiuis flavipes (Norton) 
(Hymenoptera: Tenthredinidae) 

A deep, transverse concavity or depres- 
sion is found on the seventh tergite in males 
of some sawflies of the subfamily Selandri- 
inae (Tenthredinidae). most of which are 
fern feeders. This stnicture was first recog- 
nized by Malaise (1944; 28, 29, fig. 12) in 
some species of Neostromboceros Rohwer 
and in Aneugmeuus jacobsoni (Enslin) from 
southeastern Asia. Malaise coined the temi 
"sinus se.Micilis" "for this hitherto unnoticed 
secondary sexual organ of unknown purpose 
of certain species of the genus Neostrom- 
boceros Rhw." It is difficult to see the sinus 
sexualis in dried specimens because the ab- 
domen is usually curved up with the anterior 
and posterior surfaces of the depression 
pressed together, thus hiding it. Modifica- 
tions of the abdomen (Figs. 1-2) include the 
fifth and sixth tergites each divided into two 
halves by a medial suture and the hind mar- 
gin of the sixth tergite carinate with two 
short apically projecting spines at the center; 
the seventh tergite deeply concave, cut deep- 
ly into the abdomen and posteriorly raised 
into a carinate posterior margin; and the 
eighth tergite with a large, central, oval, 
opaque or subopaque. shallowly depressed 
area. The functions of these modifications 
have not been documented. 

This sexual modification also occurs in 
males of some New World and Palearctic 
species of Aneiigmeniis Hartig (DRS, per- 
sonal observation), though it has not been 
recorded in the literature. It is present in 
males of Aneiigineiiusfloridella Ross (south- 
eastern U.S.). A. scittellatiis Smith (south- 
western U.S.. northern Mexico), unidentified 
Anengmenus species (Central and South 
America), A. japoniciis Rohwer and A. kio- 
tonis Takeuchi (Japan), and A. corona tiis 
(Klug) and A. fuerstenhergensis (Konow) 
(Europe). We have not seen it in other gen- 

era except for Neostromboceros. We here re- 
port the occurrence of the sinus sexualis in 
Anengmenus fiavipes (Norton), a fern-feed- 
ing sawfly that occurs in the eastern United 
States and Canada and provide evidence that 
the structure serves to transfer glandular se- 
cretions from the male to the female. 

A pair of A. flavipes (Figs. 3-4) were ob- 
served by SAM on low foliage in a mixed 
forest near Tobermory, Ontario, Canada. 
They were first noticed as they chased each 
other on the upper surface of a leaf, after 
which they took the position shown in Figs. 
3-4. with the mouthparts of the female 
firmly affixed to the seventh tergite of the 
male. This position was held for about a 
minute, then broken, then reestablished af- 
ter what appeared to be a brief attempt at 
copulation. It is not known whether copu- 
lation took place prior to the apparent trans- 
fer of glandular secretions from the male to 
the female A. flavipes. 

We interpret this behavior in A. flavipes 
as the transfer of glandular secretions from 
male to female. Some other insects are 
known to transfer nutritional substances to 
the female both before and after mating. 
This behavior is well known in tree crickets 
{Oecanthus spp.). males of which produce 
glandular products in external notal glands. 
The time females spend feeding on the 
males" exudates is significantly coixelated 
with egg production (Brown 1997). Some 
male cockroaches have tergal glands posi- 
tioned similarly to those of A. flavipes, and 
the female feeds on the products of those 
glands in the same fashion (Brossut et al. 
1975). Males of some soft-winged flower 
beetles, such as Anthocomus bipiinctatns 
Harrer (Malichiidae). also have glandular ar- 
eas in a conspicuous depression at the apex 
of the elytra (SAM. personal observation). 

Most sawflies display little or no court- 
ship behavior, as mentioned in some gen- 


Figs. 1 — t. Aiiciit:ininu\ lUni/u s \. Laii.'ral view of the "sinus sexualis." 2. Apex of abdomen, dorsal view; 
a = "sinus sexualis" of 7th tergite; b = central, opaque area of 8th tergite. 3 — I. Nuptial feeding positions, male 
in front, female behind. 

eral discussions (Benson 1950; Anderbrant 
1993; Viitasaari 2002). At most, rapid wing 
vibration has been observed in some spe- 
cies such as Hemitaxonus dubitatiis (Nor- 
ton) (Tenthredinidae: Selandrdiinae) (Gordh 
1975) and Arge aiuuilipes (Klug) (Argidae) 
(Scott 1991). The observations reported 
here represent the first record of the possi- 
ble function of the sinus sexualis in male 
Selandriinae and the first record of nuptial 
feeding in sawflies, and so far as we know, 
in Hymenoptera. We hope this will lead to 
further study of this unusual behavior in 
Aneiigmemis, eventual determination of its 
purpose, and determination of the chemical 
substances involved. 

Literature Cited 

Anderbrant. O. 199,1. Chapter ?. Pheromone biology 
of Sawflies. pp. 1 19-154. //; Wagner. M. R. and 

K. E Raffa. eds. Sawfly Life History Adaptations 
to Woody Plants. Academic Press. Inc.. San Die- 
go. 581 pp. 

Benson. R. B. 1950. An introduction to the natural 
history of British sawflies (Hymenoptera Symphy- 
ta). Transactions of the Society for British Ento- 
mology 10: 45-142. 

Brossut. R.. R Dubois. J. Rigaud. and L. Sreng. 1975. 
Etude biochemique de la secretion des glandes ter- 
gales des Blatteria. Insect Biochemistry 5: 719- 

Brown. W. D. 1997. Courtship feeding in tree crickets 
increases insemination and female reproductive 
life-span. Animal Behaviour 54: L'^69-1382. 

Gordh. G. 1975. Sexual behavior of Hemita.xonous 
[sic] ditbitutiis (Norton) (Hymenoptera: Tenthre- 
dinidae). Entomological News 86: 161-166. 

Malaise. R. 1944. Entomological results from the 
Swedish Expedition 1934 to Burma and British 
India. Hymenoptera: Tenthredinidae. Arkiv for 
Zoologi 35 A: 1-58. 

Scott. J. K. 1991. Arge annuliiu's Klug (Hymenoptera: 
Argidae), a sawfly associated with Riiniex siigil- 



lauis Thunb. (Polygonaceae). Journal of the En- 
tomological Society of South Africa 54: 265-270. 
Viitasaari, M. 2002. Sawflie.s (Hymenoptera. Symphy- 
ta) I. A Review of the Suborder, the Western Pa- 
learctic Taxa of Xyeloidea and Pamphilioidea. 
Treniex Press Ltd.. Helsinki. 516 pp. 

David R. Smith, Systematic Entomology 
Laboratory. PSI. Agricultural Research 

Serx'ice, % National Museum of Natural 
History, Smithsonian Institution, Washing- 
ton. DC 20560-0168. U.S.A. (e-mail: and Stephen A. 
Marshall. Department of Environmental Bi- 
ology, University of Guelph. Guelph. On- 
tario. Canada NIG 2WJ (e-mail: 
samarsha @ uo guelph. ca ) 

105(3), 2003, pp. 792-793 


New Nomenclatural Applications for Certain African Heptageniidae (Ephemeroptera) 

Four species of African Heptageniidae 
have had a dubious history of generic 
placement. These species were originally 
described as Adenophlebia bequaerti Navas 
1930: A. siniiosa Navas 1931; A. torrinen'is 
Navas 1 930; and Afroniiriis njalensis Kim- 
mins 1937. All eventually were placed in 
the genus Notomirus Crass (1947) by De- 
moulin (1956, 1970). Gillies (1963) syn- 
onymized Notonuriis with Compsoneuriella 
Ulmer, a genus originally described from 
southeastern Asia (Ulmer 1939). Gillies 
( 1984) later elaborated his rationale but did 
not account for several other related Asian 
genera, and his synonymy was not univer- 
sally accepted. Subsequently, Braasch and 
Soldan (1986) synonymized Compsoneu- 
riella with Compsoneuria Eaton (1881). 
This synonymy was not followed, and cur- 
rent research shows that Compsoneuriella 
and Compsoneuria belong to different 
clades (McCafferty and Wang, in manu- 
script). In recent lists, these African species 
have continued to be considered in the ge- 
nus Compsoneuriella; however, McCafferty 
and de Moor (1995) indicated that generic 
placement was tenuous. 

Based on my studies of generic charac- 
terization and a broad spectrum of hepta- 
geniid materials from throughout the world, 
the four species are members of the genus 
Thalerosphyrus Eaton (1881). Three of the 
species are thus placed as T. hequaeni. new 
combination, T. njalensis, new combina- 
tion, and T. tortinenis. new combination. 
Such a new combination of the fourth {A. 
sinuosa) species, however, creates a sec- 
ondary homonym with respect to a differ- 
ent. Oriental species that has been known 
as Thalerosphyrus sinuosus (Navas) (origi- 
nally Ecdyonurus sinuosus) based on Ul- 
mer's (1939) recombination. The African 
species is therefore renamed Thalerosphy- 

rus longinosi McCafferty, new replace- 
ment name, for Adenophlebia sinuosus Na- 
vas 1= T. sinuosus (Navas) 1931, nee T. 
sinuosus (Navas) 1933). The new epithet is 
after the Spanish Jesuit, Longinos Navas. 

In addition to examining adults of T. be- 
quaerti from the Democratic Republic of 
Congo and larvae and adults of T. njalensis 
from South Africa now held in the Purdue 
Entomological Research Collection, the de- 
scriptive literature associated with the spe- 
cies in question, for example, that brought 
together by Gillies (1984), has been suffi- 
cient for determining generic assignment. 
The fundamental characteristics associated 
with Thalerosphyrus include in the adults: 
mesothoracic sterna with a parallel-sided 
median depression, penes lacking dorsolat- 
eral spines, forelegs with a strongly elon- 
gated tarsal segment 1, and male hindtarsi 
about two-thirds or more the length of the 
hindtibiae: and in larvae: galealaciniae with 
scattered setae ventrally, mandibles with a 
single apical denticle, caudal filaments with 
whorls of spines well-developed, abdominal 
gills not strongly elongated, and meso- and 
metathoracic nota above the coxae pro- 
duced as acute or subacute posterior pro- 

The African species T. ethiopicus Soldan 
1977 was correctly placed to genus when 
discovered, as was the Madagascar species 
T. josettae Sartori and Elouard 1996. Thus, 
among the Arctogaean family Heptageni- 
idae, only Thalerosphyrus and the well- 
documented Afronurus Lestage (see e.g., 
Schoonbee 1968, Demoulin 1970. Mc- 
Cafferty 2003) can presently be document- 
ed from the Afrotropical Region. 

Literature Cited 

Braasch, D. and T. .Soldan. 1986. Die Heptageniidae 
de Gombak River in Malaysia (Ephemeroptera). 
Reichenbachia 24: 41-32. 


Crass. R. S. 1947. Mayflies (Ephemeroptera) collected 

by J. Omer-Cooper in the Eastern Cape Province. 

South Africa, with a description of a new genus 

and species iNotoiuinis cooperi). Proceedings of 

the Royal Entomological Society of London 16: 

Demoulin. G. 1956. Revision de quckjues Ephenie- 

meropteres decrils du Congo Beige par L. Navas 

II. Bulletin & Annales de la Societe Royale 

d'Entomologie de Belgique 92: 44-52. 
. 1970. Ephemeroptera des fauna Ethiopienne 

et Malgache. South African Animal Life 14: 24- 

Eaton. .A. E. 1881. An announcement of new genera 

of the Ephemeridae. Entomologist's Monthly 

Magazine 18: 21-27. 
Gillies, M. T. 1963. A list of Ephemeroptera in the 

National Museum, Bulawayo. southern Rhodesia. 

Entomologist's Monthly Magazine 98: 232-233. 
. 1984. On the synonymy of Nolonunis Crass 

with Compsoneuriella LHmer (Heptageniidae). pp. 

21-25. In Landa. V., T. Soldan. and M. Tonner. 

eds. Proceedings of the Fourth International Con- 

ference on Ephemeroptera. Czechoslovak Acade- 
my of Sciences Institute of Entomology. Budejo- 

McCafferty, W. R 2003 (2002). Gose's African Ephem- 
eroptera (Baetidae, Heptageniidae). Entomologi- 
cal News 113: 299-308. 

McCafferty. W. R and F C. de Moor 1995. South Af- 
rican Ephemeroptera: problems and priorities, pp. 
463-476. In Corkum. L. and J. Ciborowski, eds. 
Current directions in research on Ephemeroptera. 
Canadian Scholars" Press, Toronto. 

Schoonbee. H. J. 1968. A revision of the genus Afron- 
iinis Lestage (Ephemeroptera: Heptageniidae) in 
South Africa. Memoirs of the Entomological So- 
ciety of Southern Africa 10: 1-62. 

LUmer. G. 1939. Eintagsfleigen (Ephemeropteren) von 
den Sunda-Inseln. Archiv tiir Hydrobiologie. Sup- 
plement 16: 44.^692. 

W. p. McCafferty, Dcpartnicnt of Ento- 
mology. Purdue University, West Lafayette, 
IN 47907. U.S.A. (e-mail: pat^nccajferty® 

105(3). 2003, pp. 794-796 


Bdelloid Rotifers (Rotifera: Bdelloidea) Inhabiting Larval Black Flies (Diptera: 
Simuliidae) and Their Effect on Trichomycete (Zygomycota) Fungal Abundance 

Black fly larvae (Diptera: Simuliidae) are 
restricted to lotic habitats where they an- 
chor themselves with a silken pad spun onto 
solid substrates (e.g., rocks) and filter food 
from the water column (Adler and Mc- 
Creadie 1997). Symbiotic relationships 
have evolved between black flies and other 
organisms, including, bacteria, fungi, nem- 
atodes, viruses, and protozoans (Crosskey 
1990). During laboratory investigations of 
endosymbiotic trichomycete fungi of the 
genus Sinittium (Zygomycota: Trichomy- 
cetes) and the larval black fly SiuniUuni vit- 
tatiini Zetterstedt cytospecies IS-7, we 
found bdelloid rotifers (Rotifera: Bdello- 
idea) in the simuliid larval midgut (Fig. 1 ). 
Here we report the first record of this as- 
sociation and provide evidence that the 
presence of rotifers in the larval midgut in- 
fluences the ability of Sinittium to establish 
in the hindgut. 

Larvae of Siinuliuin vittatum cytospecies 
IS-7 were reared from eggs obtained from 
a parasite-free colony housed at the Uni- 
versity of Georgia (Athens, GA, U.S.A.). 
Approximately three weeks after submer- 
gence of eggs in 500 ml of aged tap water 
maintained at 22°C (Percival® incubator. 
Model: 1-36 VL), 40 larvae each were 
placed in 1-L polypropylene containers 
with 500 ml of aged tap water and moved 
to another incubator All containers were 
aerated with aquarium pumps, and larvae 
were fed daily on a fish food slurry 
(McCreadie and Colbo 1991). 

Fungi were reared on plates of 1/10 Brain 
Heart Infusion agar (Difco® 235-500: 
0037-17) at room temperature (22-25°C) 
with sterile water overlays added to induce 
trichospore production. Trichospores are 
single sporangia, each housing a single spo- 
rangiospore and are the asexual infective 

stage of trichomycete fungi (Lichtwardt 
1986). Once the trichospore enters the black 
fly larval hindgut, the sporangiospore ex- 
trudes, attaches to the cuticle, and produces 
a new thallus. In our experiments, a dosage 
of 4,000 trichospores/ml of rearing water 
was used. 

To determine fungal abundance in hosts, 
larvae 4 days after inoculation were dis- 
sected in a drop of distilled water and the 
mid- and hindguts removed. Under phase- 
contrast microscopy, the hindgut was 
viewed at 400X through a 10 mm X 10 mm 
ocular grid. The number of grid squares that 
contained one or more hyphae were count- 
ed; relative abundance was expressed as the 
percentage of grid squares containing hy- 
phae. During routine dissections in three 
experiments, four different treatment con- 
tainers, out of 36, had larvae with active 
bdelloid rotifers in their midguts. In exper- 
iments 1 and 2, one out of 12 containers in 
each experiment had larvae with rotifers: in 
experiment 3, two out of 12 containers 
housed infected larvae. A total of 186 lar- 
vae were examined from these containers 
and 37 (19.9%) contained rotifers; abun- 
dance of bdelloids ranged from to 24 in- 
dividuals per larval host. Rotifers used a 
telescoping-type locomotion and fed on 
green algae in the simuliid midgut. Al- 
though the identity of the rotifers remains 
unknown, they possess characters consis- 
tent with the family Philodinidae (Wallace 
and Snell 2001). Whether the black fly lar- 
vae acquired bdelloids before or after tri- 
chospore inoculation, is unclear. 

Bdelloid rotifers are free-living inverte- 
brates that inhabit aquatic vegetation, sedi- 
ment of lentic habitats, moist forest soils 
(Wallace and Snell 2001), and even the sur- 
face of freshwater insects and crustaceans 


Fig. 1. A. Bdelloid rotifers inhabiting the peritrophic niatri I I 
7. Bdelloids are located posterior to the food bolus on the left. Stale bar 
partial eversion of internal viscera. Scale bar = 23 ixm. 

I s / ruiii cytospecies IS- 

100 ijini B A bdelloid rotifer with 

(Peiinak 1978). There are two dubious re- 
ports of bdelloid rotifers entozoic in larval 
Citlex (Diptera: Culicidae) and Chironoinus 
(Diptera: Chironomidae) (Marchoux 1898. 
Bartos 1951). Relationships between roti- 
fers and trichomycete fungi are unknown, 
but some parasitic fungi, such as Harpospo- 
riiiin (Deuteromycota: Moniliales), depend 
on rotifers for reproductive success (e.g.. 
Baron 1980). 

The source of the bdelloids in our ma- 
terial is unknown but might be related to 
the anhydrobiotic nature of these rotifers 
(Ricci 1987. 1998). The ability to survive 
dehydration (i.e.. in a dessicated form) 
might indicate an airborne origin from 

within the laboratory or building. Stock cul- 
tures of larvae were always free of rotifers. 
Infections were not a result of epizoic ro- 
tifers since they were contained between 
food boluses in the black fly midgut in three 
different experiments conducted on differ- 
ent occasions. Thus, our reported occur- 
rence of rotifers is not an isolated event. 
Bdelloid rotifers were noted in hindguts of 
several larvae, but none were alive. Also, 
dead bdelloid rotifers have been seen in 
hindguts of field collected S. tuberosum 
(Lundstrom) cytospecies F larvae from Mo- 
bile County. Alabama (Nelder, unpublished 

During experiments in which rotifers 

Table I. Mean relative abundance of Swittiuni meguzygosponim. S. near typlicllimi. and S. 
hindgut of larval Siuuiliiiiu viihituin cytospecies lS-7. 

m Co 

5. inci>azygosponim (experiment 1 )" 
.S. n. typlielluin (experiment 2) 
S. n. lyphelhim (experiment 3) 
5. morhpsiim (experiment 3) 

19.2 (47) 

14.7 (20) 
8.4 (27) 

26.8 (20) 

0.6 (33)* 
1.2 (10)* 
0.0 (19) 
9.9 (10) 

" Relative abundance = percentage of ocular grid squares containing hyphae. For each experiment data were 
analyzed using a (-test on arcsine transformed percents: however, raw data are presented for comparative pur- 
poses. For each species, an asterisk indicates a significant difference (p < 0.05) in hyphal abundance. No test 
could be performed on S. n. typhelhim from the second experiment. 

''Each experiment had a total of 12 replications. 


were found, black flies had been dosed with 
trichospores of either Smittium megazyi^o- 
sporum Manier and Coste, 5. near typhel- 
lum Manier and Coste, or S. morbosiim 
Sweeney. In black fly larvae taken from ro- 
tifer-infected containers, the relative abun- 
dance of both S. megazygosponim and S. 
near typhellum in the hindgut was signifi- 
cantly lower (p < 0.05) than in larvae taken 
from rotifer-free containers (Table 1 ). Tri- 
chospores of these fungi are well within the 
size range of particles fed on by bdelloids 
(i.e., 4-17 |jLm; Gilbert 1985). Accordingly, 
bdelloids might have reduced the number 
of trichospores passing to the hindgut. 
which in turn reduced the number of at- 
tached thalli. 

We suspect the nature of the relationship 
between bdelloid rotifers and black fly lar- 
vae is one of accidental commensalism. Un- 
der this scenario, larval black flies are not 
effected by the rotifers; however, the roti- 
fers use trichospores as a food source. 
Clearly, the exact nature of the relationship 
between black flies and rotifers warrants 
further investigation. 

We thank A. Orstan, T W. Snell, and P. 
H. Adler for comments on this discovery 
and encouraging its documentation. Finan- 
cial support came from the National Sci- 
ence Foundation Grant DEB 0075269 
awarded to J. W. McCreadie and the Uni- 
versity of South Alabama. 

Literature Cited 

Adler, R H. and J. W. McCreadie. 1997. The hidden 
ecology of black flies: Sibling species and ecolog- 
ical scale. American Entomologist 43: 153-161. 

Baron. G. L. 1980. Fungal parasites of rotifers: Har- 
posporium. Canadian Journal of Botany 58: 443- 

Bartos. E. 1951. The Czechoslovak Rotatoria of the 
Order Bdelloidea. Vestnik Ceskoslovenske Zool- 
ogicke Spolecnosti XV: 241-344. 

Crosskey, R. W. 1990. The Natural History of Black- 
flies. Chichester. England: John Wiley & Sons. 
Ltd. 711 pp. 

Gilbert, J. J. 1983. Competition between rotifers and 
Daphnia. Ecology 66: 1943-1950. 

Lichtwardt, R. W. 1986. The Trichomycetes: Fungal 
As.sociates of Arthropods. New York. New York: 
Springer- Verlag. 343 pp. 

Marchoux, E. 1898. Note sur un Rotifere {Pliiloclhui 
parasitica n. sp.) vivant dans le tube digestif de 
larves aquatiques d'insectes. Comptes Rendus des 
Seances de la Societe de Biologic et de ses Filiales 
X.V.5: 749-750. 

McCreadie. J. W. and M. H. Colbo. 1991. The influ- 
ence of temperature on the survival, development, 
growth and chromosome preparation quality of 
the EFG/C, ACD, and AA cytotypes of the Si- 
miilium vemistum/vereciiiu/itin complex. (Diptera: 
Simuliidae). Canadian Journal of Zoology 69: 

Pennak. R. W. 1978. Freshwater hnertebrates of the 
United States. 2nd ed. John Wiley & Sons. New 
York, New York. 805 pp. 

Ricci. C. 1987. Ecology of bdelloids: How to be suc- 
cessful. Hydrobiologia 147: 117-127. 

. 1998. Anhydrobiotic capabilities of bdelloid 

rotifers. Hydrobiologia 387/388: 321-326. 

Wallace, R. L. and T W. Snell. 2001. Phylum Rotifera, 
pp. 195-254. In Thorp, J. H. and A. R Covich. 
eds. Ecology and Classification of North Ameri- 
can Freshwater Invertebrates. 2nd ed. Academic 
Press, New York, New York. 1056 pp. 

Mark P. Nelder and John W McCreadie, 
Department of Biological Scieiice.s. Univer- 
sity of South Alabama. Life Sciences Build- 
ing. Room 124. Mobile. AL 366H8-0002. 
U.S.A. {e-mail: 


105(3). 2003. p. 191 


Depository of the Holotype of Aiilonlini ncikcihurui Williams and Millei 
(Hemiptera: Pseudococcidae) 

In a recent article we described and il- 
lustrated the three mealybug species of the 
Antonina crawi Cockerell complex (Wil- 
liams and Miller 2002). In addition, we in- 
cluded a key to the species of Antonina that 
occur on bamboo and described and illus- 
trated one additional species. Two new spe- 
cies were included: A. macii and A. naka- 
luirui. In the "Type material" section of the 
description of A. nakaharai (page 903), we 
did not explicitly mention the depository of 
the holotype, but indicated in the "Speci- 
mens examined" section on page 906 that 
the series of specimens that included the 
holotype was deposited on BMNH and 
USNM. We did not indicate which of the 
two museums was the depository of the ho- 
lotype. and it is possible (through somewhat 
unlikely) to assume that the holotype was 
not one of the specimens mentioned in the 
"Specimens examined" section. This omis- 
sion was brought to our attention by F. 
Christian Thompson, who indicated that A. 
nakaharai should be considered as a nonien 
nudum until the description meets all cri- 
teria of Chapter 4 of the International Code 
of Zoological Nomenclature (2000); specif- 
ically, that we did not state where the ho- 
lotype was deposited. Although broad in- 
terpretation of the description could suggest 
that the primary type was in one of the two 
museums, it is not clear if this information 
is specific enough to fit the criteria of Ar- 
ticle 16.4.2 requiring "a statement indicat- 

ing the name and location of that [the] col- 
lection" [where the holotype is deposited). 
To be certain that there is no question of 
the validity of A. nakaharai. we here state 
that the holotype is deposited in the Coc- 
coidea portion of the National Museum of 
Natural History Entomological Collection. 
Smithsonian Institution, Beltsville, Mary- 

Acknowledgments. — We are grateful to 
F. Christian Thompson, Systematic Ento- 
mology Laboratory, PSI, Agricultural Re- 
search Service, U.S. Department of Agri- 
culture (SEL), for drawing attention of this 
problem and to David R. Smith of SEL for 
reading and making suggestions about this 

Literature Cited 

Williams. D. J. and D. R. Miller. 2002. Sy.stematic 
studies on the Anioiiinu crawi Cockerell (Hemip- 
tera: Coccoidea: Pseudococcidae) complex of pest 
mealybugs. Proceedings of the Entomological So- 
ciety of Washington 104: 896-911. 

International Commission on Zoological Nomencla- 
ture. 2000. International Code of Zoological No- 
menclature. International Trust for Zoological No- 
menclature. London. U.K., 306 pp. 

Douglas J. Williams. Departnwnt of En- 
tomology. The Natural History Museum. 
Cromwell Road. London SW7 5BD. U.K. 
and Douglass R. Miller, Systematic Ento- 
mology Laboratoiy, PSI. Agricultural Re- 
search Sen'ice. U.S. Department of Agri- 
culture. Beltsville. MD 20705. U.S.A. (e- 

105(3). 2003, pp. 798-SOO 

Book Review 

Scarab Beetles (Coleoptera: Scarabaei- 
dae) of South Carolina. Biota of South 
Carolina Vol. 2. By Phillip J. Haipoot- 
lian. 2001. Clemson University. Clem- 
son. S.C. ISBN 0-9712527-0-X. 157 pp. 
$37.50 (softcover). Available from Public 
Service Room, 96 Poole Agricultural 
Center, Clemson University, Clemson, 
South SC. 29634-0129 or purchase at 

Identifying scarab beetles, especially for 
the generalist or serious hobbyist, is often 
quite challenging. The book by Arnett et al. 
(2002) has simplified generic level identi- 
fications but species level determinations 
remain problematic and require consulta- 
tion of single species descriptions or revi- 
sions widely scattered in the scientific lit- 
erature. For eastern United States, the re- 
gional works of Dillon and Dillon (1972) 
and Downey and Arnett ( 1996) provide the 
only comprehensive reviews of the region, 
but these works are incomplete and out of 
date. The works of Woodruff (1973). 
Woodruff and Beck (1989). and Ratcliffe 
( 1991 ) are also useful for some groups. The 
book by Harpootlian, Scarabs Beetles of 
South Carolina, is extremely useful for 
identifying species from Virginia to Geor- 
gia. For e.vample, of the 260 species and 
subspecies of scarabs that occur or are like- 
ly to occur in Virginia (Evans, in prepara- 
tion), 225 also occur in South Carolina. 

The foundation for the Scarabs of South 
Carolina is the species checklist gleaned 
from the papers of Cartwright ( 1 934, 1 939, 
1950) and Kirk (1969, 1970). Literature re- 
cords not verified by recent collections 
were also included, unless they could be 
shown to be in error The checklist was sup- 
plemented with data from recent mono- 
graphs. Additional data were gathered from 
university, museum, and private collections, 
as well as from the author's own field work. 

The classification used by Harpootlian 

follows that of Downie and Arnett ( 1996), 
except that the tribe Pentodontini and sub- 
families Trichiinae and Valginae are re- 
tained. Lawrence and Newton (1995) el- 
evated several subfamilies within the 
Scarabaeidae to family rank (Geotrupidae, 
Ochodaeidae, Hybosoridae. Glaphyridae, 
Ceratocanthidae, Trogidae), a classifica- 
tion followed by Arnett et al. (2002). As 
is unfortunately customary among North 
American workers, the families Lucanidae 
(stag beetles) and Passalidae (bess bee- 
tles) are not included. Given the relatively 
small number of species in these two fam- 
ilies represented in any region of United 
States, future authors would do well to 
consider launching more inclusive works 
aimed at completely characterizing the en- 
tire superfamily Scarabaeoidea of their re- 

The keys of Scarabs of South Carolina 
are clear, concise, generally well illustrated 
and are easy to use, especially by those not 
intimately familiar with the group. This is 
due largely to the fact that the keys are de- 
signed primarily to identify taxa rather than 
reveal phylogenetic relationships. Sources 
for each key, when applicable, are cited be- 
neath the heading. Couplets are numbered 
to allow the user to easily backtrack 
through the key. Keys are provided to dis- 
tinguish subfamilies, tribes, genera, and 
species of South Carolina scarabs. Line 
drawings are used to illustrate characters 
used in the key. The male genitalia of many 
species are illustrated, particularly those of 
the genera Phyllophaga and Serica. 

Each genus is followed by a list of its 
synonyms, type species, and a brief taxo- 
nomic review. Many genera are illustrated 
with habitus drawings that originally ap- 
peared in Woodruff (1973). Others are il- 
lustrated with reasonably clear black and 
white digital photographs or computer-as- 
sisted line drawinss. The line drawings do 



not always accurately present the overall 
gestalt of the beetle, having rendered some 
of them shorter and wider than they appear 
in real life. An additional set of digital 
black and white habitus photos appears in 
an appendix at the back of the book. The 
overall utility of the book would have in- 
creased considerably had these images ap- 
peared in the appropriate places in the text. 

The species information is easily acces- 
sible. The species are listed alphabetically 
within each genus rather than as they ap- 
pear in the keys. This is an extremely useful 
feature, particularly for retrieving species 
data from larger genera such Aphodius and 
Phyllophaga. Each species is followed by 
its synonymy, a brief description, supple- 
mentary notes on taxonomy and biology, 
overall distribution, and South Carolina re- 
cords. The South Carolina records are listed 
not by county, but rather alphabetically, a 
feature that does not easily facilitate atlas- 
ing efforts. Instead, the reader must refer to 
an appendix that provides an alphabetical 
listing of South Carolina localities, each 
followed by its county and ecoregion. Al- 
though this appendix eliminated much rep- 
etition throughout the text, the information 
would have been more user-friendly had it 
appeared with the species synopses. 

A very useful three-page morphological 
glossary appears at the end of the work, 
clearly defining terms used in the keys, de- 
scriptions and text. The "Literature Cited" 
section appears complete for the taxa cov- 
ered and includes publications up to the 
year 2000. 

Harpootlian's taxonomic detective work 
is illuminating. He has clarified the author- 
ship and dates for several taxa and presents 
a number of new synonyms. For example, 
the authorship of Dynastes is MacLeay, not 
Kirby and Ceratocanthinae is Marti'nez, not 
Cartwright and Gordon. Osinodennu Saint- 
Fargeau (usually cited as Le Peletier) and 
Serville was first proposed as a subgenus of 
Trichiiis. The authorship of Ochodaeiis. Po- 
pillia. Eiiparia, and Plectris are Saint-Far- 
aeau and Serville, 1828, not Serville, 1823 

or 1828. The following synonyms are pre- 
sented: Copris fricator cartwrightii Robin- 
son = C. fricator (Fabricius); Stepluuuica 
thonicica Casey = S. areata (Fabricius): 
Euphoria appalachia Casey = E. scolopa- 
cea Casey, and E. o.xysternwn Casey = E. 
sepiilchralis (Fabricius) (see Hardy 2001); 
and Geohowdetuus Zunino = Geotrupes 

New state records are clearly noted and 
include Onthophagiis gazella (Fabricius), 
Aphodius erraticus (Linnaeus), Martinezicl- 
la dutertrei (Chalumeau), Platyonnis no- 
tialis (Cartwright), and Serica lo.xia Daw- 
son. Two species removed from North 
American lists are noted because they still 
appear in the literature: Dialytes unihralus 
Balthasar (Aphodiinae) is restricted to Mad- 
agascar (an error traced to the 1964 Zoo- 
logical Record, due to misspelling of the 
type locality "Kanada"), and Epicometis 
vestita (Say) (Cetoniinae) is known only 
from Eurasia. 

Scarab systematists, collection managers, 
cataloguers, faunal surveyors, and hobby- 
ists working with scarabs of eastern North 
America will Hnd Scarabs of South Caro- 
lina an indispensable addition to their li- 
braries. The author is to be congratulated 
on a well-done work that is sure to inspire 
others to create additional regional refer- 

Literature Cited 

Arnett, R. H.. Jr.. M. C. Thomas. P. E. Skelley. and J. 
H. Howard. 2002. American beetles. Volume II. 
CRC Press. Boca Raton. FL. 861 pp. 

Cartwright. O. L. 1934. A list of Scarabaeidae col- 
lected at Clemson College, South Carolina. En- 
tomological News 45: 237-240. 268-269. 

. 1939. Corrections and additions to the Clem- 
son list of Scarabaeidae and other records from 
South Carolina. Entomological News 50: 284- 

. 1950. Further corrections and additions to the 

Clemson. South Carolina list of Scarabaeidae. En- 
tomological News 61: 72. 

Dillon. E. S. and L. S. Dillon. 1972. A manual of com- 
mon beetles of eastern North America. Volumes I 
and II. Dover Publications, New York, NY. 894 pp. 

Downie. N. M. and R. H. Arnett, Jr.. 1996. The beetles 
of northeastern North America. A manual of the 



Coleoptera found in northeastern North America. 
Volumes I and II. The Sandhill Crane Press, 
Gainesville, PL. 1721 pp. 

Hardy. A. R. 2001. Studies in the Euphoriina of the 
Americas (Coleoptera: Scarabaeidae). II. Status of 
names in Euphoria, types and synonymies, with 
notes on the South American species. Pan-Pacihc 
Entomologist 77(3): 127-143. 

Kirk. V. M. 1969. A list of the beetles of South Car- 
olina. Part 1 — Northern coastal Plain. Technical 
Bulletin 1033. South Carolina E.xperiment Station. 
Clemson University. 124 pp. 

. 1970. A list of the beetles of South Carolina. 

Part 2 — Mountain. Piedmont and southern coastal 
Plain. Technical Bulletin 1038. South Carolina 
Experiment Station, Clemson University. 117 pp. 

Lawrence, J. E and A. F Newton. 1995. Families and 
subfamilies of Coleoptera (with selected genera, 
notes, references and data on family-group 
names), pp. 779-1006. In Pakluk. J. and S. A. 
Slipinski, eds. Biology, phylogeny, and classifi- 
cation of Coleoptera. papers celebrating the 80th 
birthday of Roy A. Crowson. Muzeum i Instytut 
Zoologii Polska Academia Nauk, Warsaw. 

Ratcliffe, B. C. 1991. The scarab beetles of Nebraska. 
Bulletin of the University of Nebraska State Mu- 
seum 12: 1-333. 

Woodruff. R. E. 1973. The scarab beetles of Florida 
(Coleoptera: Scarabaeidae). Part I. (Coleoptera: 
Scarabaeidae). Part 1. The Liparosticti. Subfami- 
lies: Scarabaeinae, Aphodiinae, Hybosorinae, 
Ochodaeinae, Geotrupinae, Acanthocerinae. Vol- 
ume 8. Arthropods of Florida and neighboring 
land areas. Florida Department of Agricultural and 
Consumer Services. Division of Plant Industry. 
220 pp. 

Woodruff, R. E. and B. M. Beck. 1989. The scarab 
beetles of Florida (Coleoptera: Scarabaeidae). Part 
II. The May or June beetles (genus Phyllophaga). 
Volume 13. Arthropods of Florida and neighbor- 
ing land areas. Florida Department of Agricultural 
and Consumer Services. Division of Plant Indus- 
try. 225 pp. 

Arthur V. Evans. Research Associate, 
Department of Entomology, National Mu- 
seum of Natural History and Department of 
Recent Invertebrates. Virginia Museum of 
Natural History. Home address: 1600 Not- 
toway Ave.. Richmond, VA 23227 (e-mail: 


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105(4), 2003. pp. 801-808 





Thomas J. Henry and Julieta Brambila 

(TJH) Systematic Entomology Laboratory. PSI. Agricultural Research Service. U.S. 
Department of Agriculture, '^'c National Museum of Natural History. Smithsonian Insti- 
tution. RO. Box 37012. MRC-168. Washington. DC 20560-0168. U.S.A. (e-mail: thenry®; (JB) Division of Plant Industry. Florida State Collection of Arthropods, 
Florida Department of Agriculture and Consumer Services. Gainesville. FL 32614-0737, 
U.S.A. (e-mail: brambij@doacs. state. fl. us) 

Abstract. — The first report of the prostemmatine nabid Alloeorliyiicluis triiiuiciilu (Stein) 
in the United States is given based on specimens collected in two counties of Florida. We 
diagnose the genus AUoeorhynchiis. redescribe and provide photographs of the adult male 
and female of A. thwaciila. and give other diagnostic information to help separate this 
newly discovered immigrant from other North American Nabidae. Also given are the Hrst 
report of Phorticus coUaris (Stal) in Florida and additional Florida records for Pugasa 
confusa Kerzhner. 

Ke\ Words 

Insecta. Hemiptera. Nabidae. damsel bug. Alhteorliyiulnis triimicuhi. Pa- 
gasa coiifusii. Phorticus collaris. new records. United States. Florida 

The family Nabidae is a relatively small 
group of predatory bugs, commonly re- 
ferred to as damsel bugs: Kerzhner (1996) 
estimated 21 genera and 500 species world- 
wide. Hanis' (1928) monograph of the 
North American fauna remains the most 
useful reference to identify members of this 
family, despite being more than 70 years 
old. More recently. Henry and Lattin (1988) 
summarized the literature in their catalog of 
the Nabidae of Canada and the United 
States, which included two subfamilies, 
four tribes. 10 genera, and 34 species. Since 
then. Asquith and Lattin ( 1991 ) established 
the new genus Omanonabis for the western 
U.S. species O. lovettii (Harris) and dis- 
cussed the taxonomic position of Nabis 
eda.x Blatchley known from California. 
Kerzhner (1993) described the two new 

species Lasioinerus ondobata and Pagasa 
confusa from Central America. Mexico, and 
the United States; resurrected Hoplistoscelis 
pallescens Renter from synonymy under H. 
sordida Renter and removed the latter and 
H. deiuipes Hanis from the U.S. list; gave 
species status to Pagasa fiisca van nigripes; 
and synonymized the subgenus Parapagasa 
Hussey under Lainpropagasa Renter. Ker- 
zhner ( 1996) reduced Omanonabis to a sub- 
genus of Nabis Latreille. Blinn (1996) re- 
ported Phorticus collaris Stal in North Car- 
olina and Tennessee, a species previously 
known only from west of the Mississippi 
River, and Wheeler (2001) gave additional 
records and field notes for Lasioinerus an- 
dabata in Florida. 

During ongoing faunal surveys in Flori- 
da, we have discovered specimens of three 



poorly known prostemmatine Nabidae. In 
this paper, we give the first records for Al- 
loeorhyiichiis triinaciila (Stein) in Florida 
and the United States, redescribe and pro- 
vide photographs of the adult male and fe- 
male, and furnish other diagnostic infor- 
mation to help separate this newly recog- 
nized immigrant from other North America 
Nabidae. Also given are the first report of 
Phorticiis collaris in Florida and additional 
Florida records for Pagasa confiisa. 

Acronyms used for depositories cited in 
the paper are as follows: ABS (Archbold 
Biological Station, Lake Placid, Florida); 
FSCA (Florida State Collection of Arthro- 
pods, Gainesville, Florida): VGC (Vince 
Golia Collection, Boynton Beach, Florida): 
and USNM ((United States] National Col- 
lection and Natural History, Washington, 

Genu>i Alloeorhynchiis Fieber 1860 

Alloeorhychus Fieber 1860: 43 (n. gen.), 
1861: 159 (first included species); Stal 
1865 (3): 40 (descrip.), 1873: 107; 
Champion 1900: 300 (descrip.); Distant 
1904: 393 (descrip.); Kerzhner 1981: 113 
(descrip., genitalia); Henry and Lattin 
1988: 518 (cat., distr.); Froeschner 1999: 
141 (cat.). Type species: Pirates flavipes 
Fieber, 1836. Subsequent monotypy by 
Fieber 1861. 

Alloeorrhynchus [sic] (Alloeorrhynchus 
[sic]): Renter and Poppius 1909: 33 (des- 
crip., key); Barber 1922: 103 (diag., key); 
Harris 1928: 12 (descrip., key). 

Discussion. — The prostemmatine genera 
Alloeorliynchus Fieber, Pagasa Stal, and 
Phorticiis Stal are known to occur in the 
United States (Harris 1928, Henry and Lat- 
tin 1988). The genus Phorticus may be rec- 
ognized by the dull body texture, four-seg- 
mented antenna (supplementary segment II 
greatly reduced or absent), and the presence 
of a ventral median keel on the first visible 
abdominal segment. Both Alloeorliynchus 
and Pagasa can be distinguished from 
Phorticiis by their overall shiny body tex- 

ture, Hve-segmented antenna (presence of a 
distinct supplemental segment II), and the 
lack of a distinct median keel on the first 
abdominal segment. Pagasa is best recog- 
nized by the long supplementary antennal 
segment II (Fig. 1) that is half or more the 
length of antennal segment I, the relatively 
slender pro- and mesofemora (Figs. 2—3) 
that lack a stout tooth at the middle of each, 
and the overall uniformly shiny black body 
coloration. Alloeorhynchus is distinguished 
from Pagasa by the short supplementary 
antennal segment II (Fig. 4) that is much 
shorter than half the length of antennal seg- 
ment III, the angularly widened pro- and 
mesofemora (Figs. 5-6) with each angle 
bearing a stout tooth, and the overall pale 
or multicolored body, often with yellow, or- 
ange, or red on the pronotum. Only the sub- 
genera Alloeorhychus and Psilistiis Stal are 
recognized in Alloeorhynchus. and only the 
former is known from the New World. 

Alloeorhynchus trinuiciila (Stein) 1857 
(Figs. 4-11) 

Prosteninia triniacula Stein 1857: 76 (n. 

Alloeorhynchus triniacula: Stal 1873: 109 
(distr.); Champion 1900: 300 (distr.); 
Froeschner 1999: 141 (cat.). 

Alloeorrhynchus [sic] (Alloeorrhynchus 
[sic]) triniacula: Renter and Poppius 
1909: 40 (descrip., distr.); Barber 1922: 
104 (diagnosis in key); Harris 1928: 16 
(descrip., distr.): Blinn 1996: 216 (note). 

Diagnosis. — This species (Figs. 7-11) 
can be recognized by the stout, shiny body 
having the head, three marks on posterior 
lobe of pronotum (one at each posterior an- 
gle and a triangular one at base of midline), 
scutellum, hemelytra, ventral area of thorax, 
lateral line on abdomen, and a spot on each 
of connexival segments 3—6 black; the 
bright reddish-orange pronotum, except for 
three black marks on posterior lobe; and the 
pale testaceous to pale brownish-white ab- 
domen with the lateral line on each side and 
the genital sesments black. 


Figs. 1-6. Aniennae and femora of arul Alltuorlixiit liii.\ spp. 1-3, Piif;iisii fiiscci (S). 1. Antennal 
segments I-III (arrow indicates long antennal segment ID. 2. Prof'eniur. 3, Mesofemur. 4-6. Alloeorhynclnis 
niinacula ( d ). 4. Antennal segments I-III (arrow indicates short antennal segment II). 5. Profemur (arrow 
indicates angiilate area at middle bearing stout spine). 6, Mesofemur (arrow indicates angulate area at middle 
bearing stout spine). 



Description. — Male (Figs. 9-10): Length 
4.99 mm. width across widest area of hem- 
elytra 1 .44 mm, width across widest area of 
connexivum 1.90 mm. Head: Width 0.75 
mm, vertex 0.31 mm; uniformly shiny 
black. Rostniin: Length 1.40 mm, extend- 
ing to mesocoxae; fuscous. Antenna: Seg- 
ment I, length 0.51 mm; IL 0.95 mm; III, 
0.83 mm; IV, badly curled; segment I tes- 
taceous, becoming infuscated on apical 
half; segment II-IV brown to fuscous. Pron- 
otum: Length 1.30 mm, basal width 1.54 
mm; pronotum shiny red to reddish orange, 
except for black collar on anterior lobe and 
three black marks on posterior lobe, one at 
each posterior angle and along lateral edge 
and a triangular-shaped one at base of mid- 
line. Scutelhim: Uniformly dull black, with 
three large, deep punctures; clothed with 
long, pilose setae about 2 X length of setae 
on hemelytra. Hemelytron: Macropterous; 
uniformly shiny black, including mem- 
brane; thickly clothed with erect, simple se- 
tae about half the length of setae on scutel- 
lum. Ostiokir area: Evaporative area dull 
black, becoming paler on lower half; auricle 
mostly shiny fuscous or black, elongate, 
horizontal, extending nearly to posterior 
edge of metapleuron. Ventral surface: Tho- 
rax uniformly dull black; abdomen pale tes- 
taceous or pale brownish white with lateral 
line, genital capsule and adjacent segments, 
and a spot on each of connexival segments 
3-6 black. Legs: Overall pale testaceous, 
apex of procoxa becoming fuscous or 
black; pro- and mesofemora pale fuscous or 
black on apical halves, more so on outer 
face, metafemora fuscous or black on apical 
third; tibiae black at base and apex; tarsi 
and claws fuscous. Parameres symmetrical, 
reduced (see Harris 1928: 96, fig. 10). 

Female (Figs. 7-8): Similar to male in 
coloration but larger in overall size; hem- 
elytron sometimes submacropterous. not 
quite extending to apex of the abdomen. 
Length 5.95 mm, width across widest area 
of hemelytra 1.68 mm; width across widest 
area of connexivum 2.38 mm. Head: Width 
0.78 mm, vertex 0.30 mm. Rostrum: 

Length 1.54 mm. Antenna: Segment 1. 
length 0.58 mm; II. 1.14 mm; III, 1.00 mm; 
IV, ca. 1.16 mm (curled). Pronotum: 
Length 1.43 mm, basal width 1.76 mm. 

Distribution. — Previously known from 
Brazil, Guatemala, Mexico, and Panama 
(Harris 1928). Florida represents the first 
record for the United States. 

Specimens examined. — MEXICO: Ta- 
maulipas: 1 9, Adolfo Lopez Mateos, El 
Chamalito camino al Paraiso, 30-III-2001, 
400 m, L. Cervantes and N. Pefialoza 
(FSCA), on ground below Ficus cotinifolia 
H. B. & K. [Moraceae] (FSCA); Veracruz: 
1 6. Lake Catemaco. 1-15 VII 1963, D. R. 
Whitehead (USNM): Veracruz: 1 cJ, 1 9, 
Actopan La Mancha, 26-XI-1998 & 19- 
VIIl-2001, L. Cervantes, on the ground be- 
neath Ficus sp. (USNM). UNITED 
STATES: Florida: 1 9, Alachua Co.. 
Gainesville. NE 31 Ave. at 9th St., 10-VIl- 
1995, J. Eric Cronin, on Gaura angustifolia 
Herb. Willd. Ex Steud. [Onagraceae] 
(FSCA); 1 9, Highlands Co.. Archbold 
Biol. Sta.. 1 Feb. 1999. Mark Deyrup. taken 
in yellow bowl trap on main grounds in dis- 
turbed oak hammock with ferns (ABS); 1 
9. Highlands Co.. Archbold Biological Sta- 
tion. 12-11-1999. L. Riopelle and Mark 
Deyrup, taken in yellow bowl trap on main 
grounds in disturbed hammock with ferns 
(ABS); 1 9, Highlands Co., Archbold Biol. 
Sta., 16 August 2000, Mark Deyrup, taken 
on walkway by main building (FSCA); 1 
9, Highlands Co., Archbold Biol. Sta., 1 
Feb. 2001, Mark Deyrup, taken in yellow 
bowl trap at bayhead by Lake Annie 
(USNM); 1 9, Palm Beach Co., Boca Ra- 
ton, Route 441, 24 Aug. 1988, Vince Golia. 
at mercury vapor light (VGC); 2 J, St. Lu- 
cie Co., Ft. Pierce, City Park on Florida Av- 
enue between SW 11 & 12 St., in leaf litter 
under Ficus lutea Vahl [Moraceae], 18-IX- 
2001, S. E. Halbert. G. B. Edwards. K. Hib- 
bard. & J. Brambila (USNM); 1 9. St. Lu- 
cie Co., Ft. Pierce. City Park. 12-VII-2001. 
Ken Hibbard. Park, on soil below Ficus lu- 
tea. FSCA #E2001-2822 (USNM). 

Discussion. — Only two species of Al- 


Figs. 7-8. Photographs o( AllDedrliyiuinis iriiuacuta. adult 9. 7. Dorsal aspect. 8, Lateral aspect. 

loeorhyuclms are known from the United 
States. In addition to A. tnmocula repotted 
from Florida in this paper, A. nigrolohiis 
Barber is known from Arizona and Texas 
(Barber 1922). Alloeorhynclms trimaciila 
can be separated from A. nigrolobits by the 
larger size (4.50-6.00 mm), the red to red- 
dish-orange pronotum with three fuscous 
spots on the hind lobe (one at each posterior 
angle and one at base of middle), and the 
uniformly black hemelytra. Alloeorhynchiis 
nigrolobits may be distinguished by the 
much smaller size (less than 4.00 mm), the 
bicolored pronotum with the anterior lobe 
pale brown and the posterior lobe black, 
and pale testaceous hemelytra. 

Alloeorhynchiis trimaciila has been col- 
lected in several localities of Florida in leaf 
litter under Ficiis luteci where lygaeoid 

nymphs occur. Only one specimen was tak- 
en on a plant, Gaiira angiistifolia. Luis Cer- 
vantes (personal communication) has ob- 
served this nabid feeding on lygaeoid 
nymphs in leaf litter under Ficiis trees in 
Mexico. In Florida, two males were kept in 
captivity for 4 and 6 weeks, respectively. 
on a diet of rhyparochromid nymphs and 
water. When rhyparochromid nymphs were 
no longer available, one male survived for 
two weeks on laboratory-cultured, flightless 

We consider A. trimaciila a recent im- 
migrant in Florida. Despite our searches 
through the Florida State Collection of Ar- 
thropods and the National Museum of Nat- 
ural History, no material collected earlier 
than 1999 was found. Also, Florida is rea- 
sonably well collected, so if this species had 


Figs. y-10. Photographs of Alloeor/iyiichn.s niimiciila. adiill 6. 9. Dorsal aspect. 10. Lateral aspect. 


been present much earliei". it slunild have 
been found by other collectors. In addition, 
the distribution of A. trimocula in Florida 
is considerably disjunct from its previously 
known range, providing further evidence 
that it is adventive. 


Photograph of adult AUoeorhynclnis Hi 
ris; on dead leaf. 

Pliorticus collaris Stal 1 873 

Pluiriiiii.s collaris Stal 1873: 109 (n. sp.); 
Champion 1899: 301 (distn); Henry and 
Lattin 1988: 520 (cat.); Blinn 1996: 216 
(descrip., distr.). 

This species was described from Texas 
(Stal 1873) and later reported from Teapa, 
Mexico (ChaiTipion 1899). More recently. 
Blinn (1996) redescribed and illustrated P. 
collaris and gave new county records for 
Texas (Brazos and Hidalgo counties) and 
the first eastern U.S. records from North 
Carolina and Tennessee. Phorticus collaris. 
the only species of the genus known from 
the United States, can be distinguished from 
species of Alloeorhyncluis and Paficisa by 


the dull body texture, four-segmented an- 
tenna, and presence of a median keel on the 
ventral surface of the first abdominal seg- 
ment. The specimen recorded below repre- 
sents a new state record for Florida. 

Specimen examined. — UNITED STATES: 
Florida: 1 9, Leon Co., Tall Timbers Re- 
search Station, 23 June 1993. Vince Golia, at 
mercury vapor light (VGC). 

Pagasa confiisa Kerzhner 1993 
(Figs. 1-3) coiifiisa Kerzhner 1993: 43 (n. sp.). 

This relatively newly recognized species, 
described from Costa Rica, Guatemala, 
Mexico, Panama. Puerto Rico, and the 
United States (Connecticut to Florida, and 
west to California), was long confused with 
P. fiisca. from which it can be distinguished 
by the pale legs and greatly reduced male 
parameres (Kerzhner 1993. figs. 23-25). 
The genus Pagasa may be separated from 
Alloeorhynchus by the overall shiny black 
body, longer antennal segment II (Fig. 1 ), 
and the relatively slender pro- and meso- 
femora lacking a stout tooth (Figs. 2-3). 
From Pliorticiis. Pagasa may be separated 
by the shiny black body, five-segmented an- 
tenna, and absence of a median keel on the 
first abdominal segment. Though previous- 
ly reported from Florida, the only recorded 
specimen was taken in 1911 from Newber- 
ry [Alachua Co.|, Florida. Below we pro- 
vide several new county records based on 
more recently collected material. 

Specimens examined. — UNITED STATES: 
Horida: 2 6. Highlands Co., Archbold Biol. 
Stn., 27 May 1999, M. Deyrup, in yellow 
bowl trap neai" Lake Annie dock (FSCA, 
USNM): 1 6. Highlands Co., Archbold Biol. 
Stn., 28 Aug. 2()()0, M. Deyrup, in yellow 
bowl trap in marshy iuea neai- Lake Annie 
dock (ABS). 


We thank Michele Touchet (Systematic 
Entomology Laboratory. ARS. USDA. ^'c 
National Museum of Natural History. 

Washington. DC [SEL]) and Jeffrey Lotz 
(Division of Plant Industry. Florida Depart- 
ment of Agriculture and Consumer Servic- 
es. Gainesville. FL) for the photographs of 
A. trimaciihi [MT. dorsal and lateral views; 
JL. living specimen on leaf|: and Luis Cer- 
vantes (Instituto de Ecologia. Xalapa. Ve- 
racruz. Mexico) for sharing his field obser- 
vations and specimens. We also thank Ken 
Hibbard (Division of Plant Industry. Florida 
Department of Agriculture and Consumer 
Services. Fort Pierce. FL). Vince Golia 
(FSCA Research Associate. Boynton 
Beach. FL). Eric Cronin (FSCA Research 
Associate. Gainesville. FL). and Mark Dey- 
rup (ABS) for collecting and sharing their 
specimens of A. trimaciila. Susan Halbert 
(FSCA). J. D. Lattin (Oregon State Univer- 
sity. Corvallis. OR). S. Scheffer (SEL). D. 
R. Smith (SEL). and A. G. Wheeler. Jr. 
(Clemson University. Clemson. SC) kindly 
reviewed the manuscript and offered sug- 
gestions for its improvement. 

Literature Cited 

A.squith. A. and J. D. Lattin. 1991. Omanonabis: A 
new genus of Nabidae (Heteroptera) from western 
North America, with a review of Ncihis eda.x 
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Barher. H. G. 1922. Two new species of Reduviidae 
from the United States (Hem.). Proceedings of the 
Entomological Society of Wa.shington 24: 103- 

Blinn. R. L. 1996. Phorticus collaris Stal: a nabid new 
to eastern North America (Heteroptera: Nabidae). 
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103: 216-218 (1995). 

Champion. G. C. 1897-1901. Insect: Rhynchota (He- 
miptera-Heteroptera). Vol. II. /)( Godwin and Sal- 
vin. eds. Biologia Centrali-Americana. London. 
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\vi + 345-416]. 

Distant. W. L. 1903-1904. The fauna of British India, 
including Ceylon and Burma. Rhynchota. II: .wii 
+ 242 pp (1903); 243-503 (1904). 

Fieber. F. X. 1860-1861. Die europaischen Hemiptera. 
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old's Sohn. Vienna. 1860: i-vi + 1-112; 1861: 

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the study of Panamanian biodiversity. Memoirs of 


the American Entomological Institute No. 61, 393 

Harris, H. M. 1928. A monographic study of the he- 
mipterous family Nabidae as it occurs in North 
America. Entomologica Americana 9 (new series): 

Henry, T J. and J. D. Lattin. 1988. Family Nabidae 
Costa, 1853. The damsel bugs, pp. SOS-.'i^O. In 
Henry, T J. and R. C. Froeschner, eds. The Het- 
eroptera, or true bugs, of Canada and the United 
States. E. J. Brill, Leiden and New York, 958 pp. 

Kerzhner, I. M. 1981. Heteroptera of the family Na- 
bidae. In The Fauna of the U.S.S.R. Academy of 
Sciences USSR, Zoological Institute. New Series, 
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. 1993 (1992). New and little-known Nabidae 

from North America (Heteroptera). Zoosystema- 
tica Rossica I: 37-45. 

. 1996. Family Nabidae A. Costa, 1853 — Dani- 

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ger, eds. Catalog of the Heteroptera of the Pa- 

laearctic Region, Vol. 2. Cimicomorpha I. Ponsen 
and Looijen, Wageningen, The Netherlands. 

Reuter O. M. and B. R. Poppius. 1909. Monographia 
Nabidarum orbis terrestris. Acta Societatis Scien- 
tiarum Fennicae 37(2): 1—62. 

Stal, C. 1865-1866. Hemiptera Africana. I: 1-256 
(1865): 2: 1-181 (1865); 3: 1-200 (1865): 4: 1- 
275 (1866). 

. 1870-1876. Enumeratio Hemipterorum: Bi- 

drag till en foreteckning ofver alia hittills kanda 
Hemiptera, jemte systematiska meddelanden. 
Parts 1-5. Kongliga Svenska Vetenskaps-Akade- 
miens Handlingar 1870, part 1. 9(1)1-232; 1872, 
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1874, part 4, 12(1): 1-186; 1876, Part 5. 14(4): 

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105(4), 2003. pp. 809-815 




Kelly B. Miller and Michael Balke 

(KBM) Department of Entomology. Cornell University. Ithaca, NY 14853. U.S.A. 
(e-mail:; (MB) Zoologische Staatssammlung. Miinchhausenstrasse 
21. D-81247 Miinchen. Germany and Department of Entomology, The Natural History 
Museum. Cromwell Road. London SW7 5BD. U.K. (e-mail: michb(s'nhm. 

Abstract. — Copelatiis aphroditae Balke, new species, is described based on a single 
female from Baltic amber (Eocene), and C. predaveteriis Miller, new species, is described 
from five specimens preserved in Dominican amber (late early Miocene to mid-Oligo- 
cene). The former does not fit into previously defined species groups and is assigned to 
the new C. aphroditae-group. Copelatiis predaveteriis is assigned to the C. trilohatiis- 
group which currently contains at least 20 extant species and has, in aggregate, a Pan- 
iropical distribution. The discovery of these specimens is interesting since aquatic beetles 
are relatively rare in fossil tree resins. Also, they are very similar to derived extant Co- 
pelatiis. suggesting relatively prolonged stasis in the morphology of this very speciose 
genus (>430 described species). Based on these specimens. Copelatiis is at least 50 mil- 
lion years old and at least some of the species groups are perhaps 30 million years old. 


k-ater beetle, Baltic and Dominican amber, phylogeny. fossil insects 

Despite considerable interest in amber- 
preserved insect specimens, few diving bee- 
tles (Dytiscidae) have been reported from 
amber. This seems reasonable since aquatic 
beetles are seemingly unlikely candidates 
for preservation in tree resins. Nevertheless, 
the unusual occurrence of a few adult and 
even some laivae of aquatic adephagan 
beetles in amber has been mentioned in- 
cluding specimens of Gyrinidae (Hatch 
1927: Helm 1896; Keilbach 1982; Menge 
1856; Motschulsky 1856; Weit.schat and 
Wichard 1998; Wichard and Weitschat 
1996) and Dytiscidae (Berendt 1845; Helm 
1896; Keilbach 1982; Poinar 1992; Poinar 
and Poinar 1999; Spahr 1981a, b; Weidner 
1958: Weitschat and Wichard 1998: Wi- 
chard and Weitschat 1996; Wunderlich 
1986), though many of these refer to the 

same specimens. The only species of Dy- 
tiscidae described from amber so far is 
Glesseria rostrata Koch and Berendt, 1845. 
This species is based on a single larva in 
Baltic amber and was originally assigned to 
Thysanura. not Coleoptera, albeit with 
some doubt (Handlirsch 1908; Koch and 
Berendt 1854). 

Diving beetles are a comparatively well- 
studied family taxonomically, and the group 
is becoming better known phylogenetically 
(e.g., see Miller 2001 ). Despite this and de- 
spite the usefulness of fossils for providing 
unique character combinations and deter- 
mining minimum ages of divergences, rel- 
atively little effort has been placed on the 
study of fossil dytiscids. Because diving 
beetles are aquatic, the chance of fossiliza- 
tion of specimens in sediments is probably 


higher than for many other insects. The ht- 
erature does, in fact, contain many refer- 
ences to fossiUzed dytiscids (mainly im- 
pression fossils), but these are often de- 
scribed in only limited detail with very little 
evidence offered for their taxonoinic place- 
ment. The numerous dytiscid (and other hy- 
dradephagan) fossils available in collec- 
tions are a potentially large, untapped 
source of information about the evolution 
of the group. Though amber specimens of 
Hydradephaga are certainly much more rare 
than impression fossils, at least several 
comparatively well-preserved adult speci- 
mens of Copelatiis Erichson are available 
for study and are described herein. These 
are the first species of Dytiscidae described 
from adult specimens in amber. 

Copelatiis is one of four genera of Co- 
pelatinae and is the most speciose genus of 
Dytiscidae (433 described species: Nilsson 
2001) with most of its members occurring 
in the tropics. The genus has never been 
completely revised and is in need of a de- 
tailed cladistic analysis with broad taxon 
and character sampling in addition to con- 
siderable species-level monographic work. 
Extant species in the group are usually sep- 
arable on the basis of male genital structure. 
as well as coloration and fine stiiictural de- 
tails of the cuticle, all of which are difficult 
or impossible to examine in amber pre- 
served specimens. Nevertheless, the species 
described here appear to possess external 
character state combinations sufficient for 
diagnosing them from extant taxa. Here, we 
describe two new species which belong to 
the extant genus Copelatiis. C. aphroditae 
Baike from Baltic amber, and C. predavet- 
eriis Miller from Dominican amber. 

Material and Methods 

Figures. — Line drawings of C. predav- 
eteriis and C. aphroditae were created with 
the aid of drawing tubes on a Wild M3C 
and a Leitz MZ12 dissecting microscope, 

Preparation and condition of speci- 
mens. — Specimens of C. predavetenis were 

prepared with methods described by Gri- 
maldi ( 1993). They were examined by plac- 
ing the amber piece on a drop of glycerine 
on a microscope slide. A drop of glycerine 
was then placed on the dorsal surface of the 
amber piece and a coverslip placed over the 
drop. The entire piece containing C. 
aphroditae was placed in a small tray with 
glycerine and then examined. 

Measurements. — Measurements of C. 
predavetenis and C. aphroditae were made 
using ocular scales in a Wild M3C and a 
Leitz MZ12 dissecting microscope, respec- 
tively. Some of the specimens are distorted 
by compression or stretching in the amber, 
but measurements do not correct for this in 
any way. For this reason, measurements are 
unavoidably inaccurate and give only a 
general idea of the actual dimensions of 
most of the specimens. Total length (TL) 
and greatest width (GW) are provided along 
with the ratio TL/GW to provide an indi- 
cation of shape. 

Copelatiis aphroditae Balke, new species 

(Fig. I) 

Holotype. — 9, in collection A. Herrling, 
Bramsche, Germany, labeled # 1153. The 
piece is clear, yellowish golden. The beetle 
is partly covered with a whitish film of air. 
Besides the beetle, there are two pieces of 
sclerotized, cuticular material in the amber, 
which could not be identified. Also, there 
are few "Sternhaare." which are thought to 
be minute parts of oak trees and which are 
characteristic of the Baltic Amber (Weit- 
schat and Wichard 1998). 

Diagnosis. — With 19 discal striae on el- 
ytron anteriorly: posteriorly with ten striae 
visible. Submarginal stria absent (Fig. 1); 
size small (total length 5.0 mm): body in 
dorsal view approximately parallel-sided 
(Fig. 1). 

Description. — Measurements: TL = 5.0 
mm; GW = 2.2 mm; TL/GW = 2.27. 

Dorsal side (Fig. 1 ): Elongate, lateral 
margins continuously curved between pro- 
notum and elytron. Body approximately 
parallel-sided in dorsal view. Head (partly 


Figs. 1-4. 1. Copelanis aphroditae. holotype 9. dorsal habitus. 2 — 1. C. prcilnveteni 
ventral aspect. 3-4. Holotype 9, habitus. 3. Dorsal. 4. Ventral. 

Male protibia 


crushed) with distinct microreticulation 
(MR) of honeycomb-like cells; with few 
larger punctures, their diameters equal to 
tliat of meshes. Pronotum with MR of reg- 
ular, slightly longitudinally-oriented cells, 
with few larger and smaller punctures: dis- 
cally with short strioles which increase in 
length and depth laterally. Pronotum with 
lateral bead complete, extending nearly to 
anterior angle. Elytron with 19 longitudinal 
striae, with only 10 striae visible on poste- 
rior portion of elytron; without submarginal 
stria. With MR and relatively dense punc- 
tation between the striae. 

Ventral side: Prosternal process moder- 
ately convex, apex narrowly rounded. Me- 
tasternum with MR of horizontally oriented 
cells; metacoxa with MR of cells in diag- 
onal orientation, few larger punctures and 
with diagonal strioles. Metacoxal lines 
closely approximated posteriorly. Ventrites 
\-5 chagreened and with few larger punc- 
tures; with oblique strioles on ventrite 1 be- 
coming increasingly transverse posteriorly. 
Last visible sternite without lateral striae. 

Etymology. — Named after Aphroditae. 
Greek god of love. 

Occurence. — Baltic Amber, Eocene. 

Comments. — This species does not fit 
well into other, currently-recognized spe- 
cies groups, and we assign it to a new Co- 
pelatiis cipliroclitae-group based on the 
presence of nineteen discal impressed striae 
on the elytron and the lack of a submarginal 
elytral stria. 

Copelatus predaveteriis Miller, 
new species 

(Figs. 2-4) 

Type specimens. — Holotype, 9 , AMNH 
DR- 14-6 15, labeled, "AMBER: Oligo- 
Miocene Dominican Republic AMNH no. 
DR- 14-6 15 Locality :/Copelatus trilobatus- 
group Det AN H\\^'>on-96l HOLOTYPE Co- 
pelatus predaveteriis Miller 2000."" 

Paratypes (4): 1 6. AMNH, same amber 
piece as holotype, same labels; 1 i. AMNH 
DR- 14-974, labeled. "AMBER: Oligo-Mio- 
cene Dominican Republic AMNH no. DR- 

14-974 Locality:": 1 9, AMNH DR- 14-973. 
labeled. "AMBER: Oligo-Miocene Domin- 
ican Republic AMNH no. DR- 14-973 Lo- 
cality:/Amber: Oligo-Miocene Dominican 
Republic: mines at or near La Toca AMNH 
(no cat. no. yet)/ DYTISCIDAE COPELA- 
TUS SR det. PJ Spangler": 1 9. AMNH 
DR- 1 4-6 16. labeled, "AMBER: Oligo-Mio- 
cene Dominican Republic AMNH no. DR- 
14-616 Locality :/Copelatus trilobatus-group 
Det AN Nilsson-96." 

Although male specimens of Dytiscidae 
are generally selected as holotypes. a fe- 
male specimen was selected since this in- 
dividual is easily visible dorsally and ven- 
trally, and the piece of amber in which it is 
included also contains a male specimen, 
which is currently less visible, but which, 
in the future, will be easily associated with 
the female holotype. The only other male 
specimen is the least distorted of the spec- 
imens and is also easily visible. However, 
it is on the extreme edge of the amber piece 
where this piece is broken, and some of the 
specimen is exposed to the air. Parts of the 
legs and antennae are in the piece that is 
broken off. Rather than selecting this spec- 
imen, which is in more tenuous condition, 
a specimen was chosen that is well within 
a relatively strong piece of amber and is 
entirely intact. 

Diagnosis. — Eleven discal and one sub- 
marginal striae present on elytron, first stria 
not shortened, of similar length to others 
(Fig. 3); size small (total length 3.8-4.4 
mm); male pro- and mesotarsal claws un- 
modified (Fig. 2): male without strioles on 

Description. — Measurements: TL = 
3.82-4.40 mm; GW = 1.84-2.36 mm; TL/ 
GW = 1.79-2.21. 

Dorsal side: (Fig. 3). Elongate oval, lat- 
eral margins continuously curved between 
pronotum and elytron. Male apparently 
without distinct strioles on pronotum or el- 
ytra. Pronotum with lateral bead complete, 
extending nearly to anterior angle. Elytron 
with 1 I discal and 1 submarsinal striae; all 



discal striae elongate and distinct, not short- 
ened (Fig. 3). 

Ventral side: (Fig. 4). Prosternal process 
moderately convex, apex nanowly rounded 
(Fig. 4). Male pro- and mesotarsal claws 
not modified (Fig. 2). Male pro- and me- 
sotarsomeres 1-3 moderately expanded, 
ventrally with 4 rows of adhesive setae 
(Fig. 2). 2 rows on tarsomere 1. 1 row on 
2, and 1 row on 3, most proximal row with 
3 adhesive setae, other rows with 4 adhe- 
sive setae. Male pro- and mesotibiae not 

Etymology. — This species is named pre- 
(lavetenis from the Latin preda. meaning 
"predator" and veterus, meaning "of a for- 
mer time" in reference to the carnivorous 
habits and ancient existence of this species. 

Occurrence. — Dominican amber. Oligo- 
cene-Miocene. Dominican Republic. 

Comments. — This species is a member 
of the Copelatiis nilohciTus-group (sensii 
Gueorguiev (1968)) based on the presence 
of eleven discal and one submarginal im- 
pressed striae on the elytra. Within this 
group, it may be distinguished from most 
of the species by its smaller size. Other spe- 
cies in the group similar to C. predaveterus 
in size are C concolonms Balfour-Browne. 
C. gschwendtneri Guiqnot. C. perfectiis 
.Sharp. C. trilobatus. Regimbart and C. itii- 
dcciiiistriatiis Aube, and members of most 
of these species are still generally larger 
than C. predaveterus. Copelatiis predavet- 
erus has unmodified male prostarsal claws, 
has the first elytral stria elongate (not short- 
ened) and males apparently lack strioles on 
the pronotum. though fine details of cutic- 
iilar surfaces are not easy to discern on 
these specimens. Based on these features. 
C. predaveterus may be most similar to the 
extant C. perfectiis. 


The very speciose genus Copelatiis has 
l">een divided into informal species groups 
based on the number and type of longitu- 
elinal striae present on the elytron. These 
characters have yet to be adctjuately tested 

as synapomorphies, but given the extreme 
size of the genus and lack of a phylogenetic 
hypothesis it is convenient to recognize 
these informal groupings of species for ease 
of communication (Balfour-Browne 1939). 

Based on the number of elytral striae, 
Copelatiis aphroditae is rather isolated 
within Copelatiis. The maximum number of 
elytral striae in Copelatus is typically 13 or 
less, with the extant South American C. 
ahonnenci Guignot being the only known 
exception thus far This species has 22 dor- 
sal striae plus one submarginal stria 
(Gueorguiev 1968; Guignot 1939). It has 
been historically placed in a group of its 
own, the C ahonnenci-group. which was 
defined as having 14-23 striae (Guignot 
1939). This is somewhat confusing since C. 
ahonnenci is the only species of the group 
and it has 23 striae. Therefore, we redefine 
the C. abonnenci-group as those Copelatus 
with a total of 23 elytral striae. We assign 
Copelatus aphroditae to a new species 
group, the C aphroditae-group, which is 
characterized by having 19 dorsal striae an- 
teriorly on the elytron, and no obvious sub- 
marginal one. 

Copelatus predaveterus belongs to the 
group with eleven sutural and one submar- 
ginal elytral striae (Copelatiis-group 13 sen- 
su Sharp ( 1882) and trilohatus-group sensu 
Gueorguiev (1968)). Other members of the 
C. trilobatus-group are tropical and occur 
in South America, Africa, southeast Asia 
and Australia. New World members occur 
primarily in northern South America and 
their habits are essentially unknown. How- 
ever, other Copelatinae, such as Aglymbiis 
bromeliarum Scott and A. himaculatus Re- 
sende and Vanin, are known from Brome- 
liaceae (e.g. Resende and Vanin 1991). The 
unusual situation of having aquatic beetles 
entrapped in tree resin could be explained 
by their living in bromeliads, tree holes or 
other similar arboreal aquatic habitat. How- 
ever, this is certainly speculative, and there 
are other possible explanations such as en- 
trapment during dispersal or intenment in 
resins deposited in the water The fact that 


the dytiscids from amber from very dispa- 
rate localities and time periods are conge- 
neric suggests that Copelatiis may show a 
greater potential for becoming interred in 
amber, possibly because of specialized hab- 
itat preferences. Other inclusions in the Do- 
minican amber pieces are terrestrial includ- 
ing several collembolans and a terrestrial 

The age of Dominican amber has been 
controversial, but recent estimates place 
much of it at 15-20 million years (late Ear- 
ly to early Middle Miocene) (Iturralde-Vi- 
nent and MacPhee 1996) with others plac- 
ing it at as much as 23-30 million years 
(Lower Miocene to mid Oligocene) (Gri- 
maldi 1995). Baltic amber appears to be 
much older, and recent authors place it at 
some 40-55 million years (Eocene) (Mich- 
elsen 2000; Weitschat and Wichard 1998). 

Little has been proposed regarding the 
phylogeny of Copelatinae. The striate Co- 
pehitiis species appear to be monophyletic 
(Balke 2001). but relationships within this 
clade are not yet known. However, an up- 
coming molecular phylogenetic study of 
Copelatinae (Balke, Ribera & Vogler. in 
preparation) will likely help clarify the phy- 
logeny of this diverse group. When attempt- 
ing to propose potential historical explana- 
tions for diversity, it is crucial to reliably 
calibrate at least some nodes of cladograms 
used to infer divergences. Indirect, biogeo- 
graphic evidence suggests a Gondwanian 
origin of Copelatinae. earlier than perhaps 
120 mya (Balke 2001). Here, we present for 
the first time evidence for an origin of the 
more derived, striate forms of the genus 
predating perhaps 50 mya, a conclusion that 
will aid substantially any future studies of 
the evolutitin of this old and diverse group 
of beetles. 

Other Fossil Copelatus 

Our new species are the first Copelatinae 
species described from amber and one, C. 
iiphroditcie, is the oldest fossil copelatine 
known. Three other fossil species in this ge- 
nus have been described from well-pre- 

served impression fossils from the middle 
Miocene (10-15 mya) in Northern Cauca- 
sian layers (Riha 1974): 

Copelalus stovropoUtaiuis RiTia, 1974: 

eleven elytral striae (C. nii>r(>liiiealiis- 

Copelatus pouomarenkoi Rilia. 1974: six 

elytral striae plus one submarginal stria 

(C. iriiius group). 
Copelatus fossilis RiTia, 1974: ten elytral 

striae plus one submarginal stria (C. er- 

ichsoni group). 

Copelatus species similar to these do not 
cunently occur in this area. The closest ex- 
tant relatives of these species are probably 
Afrotropical and/or Oriental. 


KBM thanks especially Dr. D. A. Gri- 
maldi (American Museum of Natural His- 
tory) for providing the opportunity to de- 
scribe these specimens, for preparing the 
material and for much useful information 
about amber. KBM also thanks Dr Q. D. 
Wheeler for advising him on this and other 
projects. MB thanks Dr Fritz Hieke and Dr 
E. Pietrezeniuk (Natukundemuseum Berlin, 
Paleontology) for advice on the Berlin am- 
ber collections. Dr. W. Weitschat (Ham- 
burg) and Mr Carsten Crohn (Glinde) are 
thanked for the incredible opportunity to 
study the material under their care. Finan- 
cial support of the FAZIT foundation 
(Frankfurt/Main, Germany) is greatly ac- 

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105(4). 2003. pp. 816-831 


Oliver S. Flint. Jr. 

Department of Systematic Biology. Section of Entomology. National Museum of Nat- 
ural History, Smithsonian Institution, Washington, DC 20013-7012. U.S.A. (e-mail: 

Abstract. — Based on the study of types and, mostly, new material collected between 
1970 and 1980, nine species of Macrostenuim Kolenati are recognized on the island of 
Sri Lanka. The species Macronemo miiltifarium (Walker), M. nebiilositni (Hagen), and M. 
sepiiltiim (Hagen) are redescribed based on the Hrst material collected since the original 
types in the 1850s, and are formally transferred to Macrostemiim (all n. comb.). Ma- 
crostemiim splendiduin (Hagen) and M. pseiidoneiira (Brauer) are redescribed, and M. 
ethelda (Banks) from southern India is reduced to a subspecies of the latter (n. status). 
Macrostciiuiin indistinctiim (Banks), a species widespread in southeastern Asia, is newly 
recorded from the island and M. scunvcipci Malicky is placed in its synonymy (n. syn.). 
Macrostemiim barnardi. M. krombeitii. and M. malickyi are newly described. All species 
are described, the male genitalia figured (M. kroiubeiiii and M. imdtifariiim are known 
only from females), and photos of the wing patterns presented. The species Macronema 
oblicpiiini Hagen is transferred to the genus Marilia in the Odontoceridae (n. comb.). 

Key Words: Mcicrostemitm. Hydropsychidae, Sri Lanka, new species, descriptions, il- 
lustrations, wings 

Sri Lanka (previously Ceylon) lies off 
the southeastern tip of India. Considering 
its proximity to India, it seems to have a 
surprising high level of endemism in its in- 
sect fauna. This level of endemism may be 
more due to the comparative lack of study 
of the south Indian fauna in comparison to 
the Sinhalese. Of the nine species treated 
here, only two are known from the conti- 
nent, one of which is represented by differ- 
ent subspecies on the continent and the is- 
land, the other being found throughout 
southeastern Asia as far south as Sumatra. 

The genus Macrostemum Kolenati, as 
presently recognized, is widespread over 
the World, being known from North and 
South America, Africa, and Asia. Most spe- 
cies presently placed in Macrosteirniin were 

originally described in Macronema Pictet. 
In 1982 Flint & Bueno divided Macronema 
into two generic entities, by resurrecting 
Macrostemiim. Macronema is restricted to 
the Neotropical Realm. The Sinhalese spe- 
cies were mostly described in Macronema. 
and all were placed in this genus for most 
of the 20th century. 

Walker (1852) described the first species 
now in this genus as Hydropsyche miillifar- 
ia. and all subsequent mention of this spe- 
cies has been based on the type. Hagen in 
1858 and 1859 described 7 species in Ma- 
cronema, of which three, M. splendidiim, 
M. nebiilosiim. and M. sepiiltiim now lie in 
Macrostemiim. Macronema ceylaniciim has 
been transfened to Pseiidoleptonema Mo- 
sely and M. vitrina to Oestropsyche Brauer. 


both in the Hydropsychidae; M. annnlicor- 
ne was moved to Anisocentropiis Mc- 
Lachlan in the Calamoceratidae; A7. oIjU- 
quum has remained in Macrosteiuimi till 
now. I studied the holotype of the latter 
many years ago and realized that it was not 
a hydropsychid. During the preparation of 
this paper 1 restudied the holotype, includ- 
ing its cleared male genitalia. It is an Odon- 
toceridae. and is here placed in the genus 
Marilia Miiller (n. comb.), pending further 
revision of that genus and its possible dis- 
memberment. I have not seen another ex- 
ample of this species in any material avail- 
able to me. Brauer described M. pseiido- 
neiira from Sri Lanka in 1865. a species 
rediscovered and well characterized subse- 
quently. In 1911 Banks described M. iiulis- 
tinctum from eastern India, it is here re- 
corded from Sri Lanka and the recently de- 
scribed M. saowapa, is synonymized with 
it. Three new species have been discovered 
in the material collected under the auspices 
of the Smithsonian's "Ceylon Insect Pro- 
ject" directed by Dr K. V. Krombein. and 
are described herein. 

Several of the old localities and labels 
need some clarification. Hagen in his 1858 
publication listed the types of M. splendi- 
ciiiiii and M. iiebiilosiim as from Rainbodde. 
Further, on p. 487 he stated (translated) that 
the species labelled Rainbodde are all col- 
lected at this spot in the mountains, 3,500 
to 4,000 feet above the sea. In his second 
paper (Hagen 1859) he corrected this spell- 
ing to Rambodde for the type of M. sepiil- 
titni. This locality is now accepted as Ram- 
boda and is at 7°04'N, 80°42'E in the Dis- 
trict of Nuwara Eliya. 

The paratype of M. baniardi in the Nat- 
ural History Museum. London bears a 
handwritten label stating "Ceylon," but on 
the back a handwritten "N'pitia." This has 
been interpreted (Barnard 1980, p. 66) as 
Nawalapitiya, a town in the central high- 
lands at 7°03'N, 80°32'E in the District of 

A series of M. luhulosuDi was collected 
at Kabaragala, Niilomalai, by S. & P. B. 

Karunaratne. Unfortunately there are two 
localities named Kabaragala, one a popu- 
lated place at 6°03'N, 80°18'E. which 
would place it in the lowlands in the Galle 
District. The other an estate near Adam's 
Peak at 6°56'N. 80°26'E. which is about 6.5 
km SSE of the Kitulgala Resthouse and 
right on the border between the Kandy and 
Kegalla Districts (it is labelled Kan. Dist.) 
at about 300 m. There is no Niilomalai in 
the gazetteer Based on the other two col- 
lections of M. nebiiloswn, it seems that the 
lowland site is quite improbable for this 
species, and that the estate site is the one 
intended. Another P. B. Karunaratne site, 
Karambaketiya off Memure. Knuckles 
Range, is also problematic. I can not find 
Karambaketiya in the gazetteer, however, 
there is a Mimure (the label, handwritten, 
is Memure) about 7 km NNE of Corbets 
Gap and below Knuckles Peak at about 600 
m. I place this locality for M. splendidnm 
in the Kandy District in the vicinity of Mi- 
inure at 7°26'N, 80°50'E. 

Fernand Schmid collected extensively on 
Sri Lanka (then Ceylon) in the first 4 
months of 1954. The results of this field 
work were published in a large work in 
1958. Many years later the Smithsonian 
purchased the vast majority of this collec- 
tion, including the types. They are labelled 
Ceylan, a 2 or 3 letter initial for the prov- 
ince, locality, date and F. Schmid. However, 
in his work (1958) on pages 4-24 he also 
gave the name of the water course, eleva- 
tion, and ecological setting at each of these 
localities. I have included, in brackets, these 
elevational and water course data for his lo- 
calities whenever cited. 

All descriptions are from adults, none 
having their immature stages positively as- 
sociated. Acronyms used are given in Ac- 

Macrosteiniini haniardi Flint, 
new species 

(Figs. 1-6, 43) 

This species is most similar to M. pseii- 
doiH'iini on the basis of coloration. The two 


Figs. 1-12. Male genitalia. 1-6, Macrostemum barnanli. 1, Lateral. 2. Dorsal. 3. Inferior appendage, pos- 
terosentral. 4. Phallus, lateral. 5, Tip of phallus, lateral. 6, Same, ventral. 7-12, M. imlistiiutitm. 7, Lateral. 8, 
Dorsal. 9. Inferior appendage, posteroventral. 10, Phallus, lateral. 1 1, Tip of phallus, lateral. 12. Same, ventral. 


can be easily distinguished by the complete, 
transverse, pale band near the base of the 
forewing and the three, trianguloid, pale 
marks from the anterior margin beyond in 
M. banianli. In contrast M. pseudoneura 
has a transverse, pale, band nearer the apex 
and this band is developed from the inner- 
most of the three, pale, trianguloid marks. 
In addition there are many more smaller 
differences in maculation. The male geni- 
talia suggest a closer relationship to M. iie- 
biilosiiDi and M. splemUduni based on the 
similarity in the structure of the tenth ter- 
gum and phallic apex. The basolateral ridge 
of the tenth tergum in M. hanuinU is 

Male. — Length forewing 9-10 mm. Col- 
or fuscous, with cream-colored and clear 
marking. Head golden yellow, cuticle in- 
fuscate laterally between dorsal warts; max- 
illary palpus with 4 basal segments infus- 
cate, apical segment golden; antenna with 
scape, pedicel and outer face of first flagel- 
lar segment, golden; following 3-4 seg- 
ments fuscous, then stramineous, becoming 
infuscate apically. Cuticle of meso- and me- 
tanota and pleura, black. Legs golden yel- 
low; tibia of foreleg darkened for apical 
half. Forewing (Fig. 43) with 3 pale-yellow 
marks from anterior margin, a series of 
small spots in costal cell basally, and a sin- 
gle, complete, transverse, pale-yellowish 
band at about Vs length of wing; apex with 
wash of golden setae anteriorly, posteriorly 
and apically with extensive pale yellowish 
or clear marks on a fuscous background; 
hindwing infuscate apically. Abdomen fus- 
cous. Genitalia: Ninth segment with antero- 
lateral margin broadly produced, rounded, 
posteroventral margin produced, rounded 
and shelflike above inferior appendage; 
posterior margin with scattered, elongate 
setae, becoming numerous dorsolaterally. 
Tenth tergum with dorsobasal setal wart, 
with distinct basolateral ridge, apex angu- 
late; in dorsal aspect with apex produced 
laterad. Inferior appendage elongate, paral- 
lel-sided; in posteroventral aspect with api- 
cal segments not quite Vi of total length. 

Large basal section of phallus Vs of total 
length of phallus, curving sharply into 
stem; apex broadly rounded ventrally, dor- 
sally barely elevated; apex nearly vertical 
with a small dorsal lobe and large ventral, 
liplike lobe. 

Female. — Length forewing 8—9 mm. 
Coloration as in male. Mid tibia and tarsus 
flattened, broadened, concave on inner face. 

Material examined. — Holotype, i: SRI 
LANKA, Ratnapura Dist., Weddagala, 8-13 
Feb 1977, K.V. Krombein et al., collected 
at light [NMNH]. Paratypes: Same data, 2 
6. \ 9; Gilimale, 17-18 Jun 1976, Krom- 
bein et al., light trap, 2 6\ Gilimale, 17-18 
May 1975, Wood & Petty, 1 c? (in alcohol); 
Gilimale, Induruwa Jungle, 5-1 Feb 1977, 
Krombein et al., blacklight trap, 1 9 ; 2 mi 
lea. 3 km] S of Weddagala, Sinharaja Jun- 
gle, 8-12 Feb 1977, Krombein et al., 3 9. 
Galle Dist., Kanneliya Section, Sinharaja 
Jungle, 2-5 Oct 1980, Krombein et al., col- 
lected at blacklight. 1 <5 ; Hiniduma, 22 Oct 
1973, M. & B. Robinson, collected at black 
light, 1 9. Ceylon, N|awala]piti[y|a, Mc- 
Lachlan Coll BM 1938-674, 1 9 (BMNH). 

Etymology. — I dedicate this species to 
Dr Peter C. Barnard of The Natural History 
Museum, London in gratitude for all his 
help and friendship during my many visits 
to that Museinn.'iiiKm inctistimtuiu (Banks) 
(Figs. 7-12. 44) 

Macroiiomi iiulisiiiuta Banks 1911:106 
I forewing]. 

McicroncDUi hrisi Navas 1930:5 [fore- 
wing). — Fischer 1963:178 [catalogue]. 

Macrouemo fiilvesceris Martynov 1935:186 
\S. wings[.— Fischer 1963:187 [cata- 

MacioiH'ina iiulistiiictiini: Fischer 1963:189 

Macrostenuiin saowapa Chantaramongkol 
and Malicky 1986:528 [<5, wings, 
head].— Malicky 1998:777 \6. wingsj. 
New synonym. 

Macrostemuiu liulisliiuiuin: Malicky 1998: 


777 [S . wings, synonymy of M. 
and M. fulvescens]. 

Malicky (1998) redescribed the species, 
placed M. brisi and M. fulvescens in syn- 
onymy, and extended its known distribu- 
tion. I have studied a male paratype of M. 
suowapa from the Malicky collection, com- 
paring its structures and genitalia with a 
male from Padukka, and the male type of 
M. indistinctiini from Bengal, India. The 
male genitalia are in very close agreement, 
offering no notable differences. Unfortu- 
nately the wings of the M. saowapa para- 
type are now virtually decolored and offer 
no clues in terms of pattern; the pattern as 
originally reconstructed is compatible with 
M. imiistinctitin in spite of some differenc- 
es. All the Sri Lankan material here record- 
ed is quite uniform in coloration and agrees 
perfectly with the type of M. iiutistinctiini. 
In all the material from Sri Lanka before 
me, there is no other species approaching 
M. indistiiictum in structure or appearance: 
I, therefore, believe that M. sciowupci is a 
synonym of it. 

The species is known from India, China, 
Thailand, Laos, Malaysia, Sumatra, and 
now Sri Lanka (Malicky 1998). Our collec- 
tion contains numerous captures of the spe- 
cies, but only one contains any males. Most 
captures were made at night at light traps: 
perhaps the males are generally active dur- 
ing the day with the females night active. 
This is a species of low elevations, most 
recorded elevations being 150 m or less, but 
one lot was taken at 900 m. 

Male. — Length of forewing 1 1 mm. Col- 
or tawny, wings with clear markings. Color 
of head stramineous, browner dorsally: an- 
tenna stramineous. Thoracic cuticle brown: 
legs stramineous. Forewing (Fig. 44) basi- 
cally tawny with clear marks, outlines rath- 
er indistinct: 3 clear marks from anterior 
margin, basalmost mark extending along 
anterior margin to wingbase: with mem- 
brane posteriad to clear marks darkened, 
with various, rather ill defined, pale marks 
apically and posteriorly. Hindwing clear 

with slight apical darkening. Genitalia: 
Ninth segment with anterolateral margin 
broadly produced, posteroventral margin 
produced, shelflike above inferior append- 
age: posterior margin with scattered, elon- 
gate setae, becoming numerous dorsolater- 
ally. Tenth tergum with dorsobasal setal 
wart, apex rounded: in dorsal aspect with 
translucent, mesal lobe subapically. Inferior 
appendage elongate, parallel-sided: in pos- 
teroventral aspect with two segments of 
nearly equal length. Basal section of phallus 
relatively broad, about 45% of total length 
of phallus: apex broadly rounded ventrally, 
shaiply, and obliquely elevated dorsally: 
apex nearly vertical with small indentation 
at midheight. 

Female. — Length of forewing 8-1 1 mm. 
Coloration as in male. Midtibia and tarsus 
strongly flattened, broadened and concave 
on inner face. 

Material examined. — Sri Lanka, 6 para- 
type M. saowapa (HMPC). [India], Pusa, 
Bengal, Oct 07, A. Mujtaba. S holotype 
I 1765, M. indistinctwn Banks (MCZ). 

SRI LANKA: Colombo Dist., Labuga- 
ma, 400 ft [ca. 120 m], 24 Aug 1973, G. 
Ekis, collected at blacklight, 9 9 : Bayaga- 
ma, sea level, 28-29 Aug 1973, G. Ekis. 
collected at black light, 2 9; Padukka, 6 
Aug 1972, R B. Karunaratne, at light, 3 S. 
1 9. Galle Dist., Kanneliya Jungle, 11-16 
Jan 1975, Krombein et al., blacklight. 119; 
Kanneliya, 22-24 May 1975, Wood & Pet- 
ty, collected in Malaise trap, 2 9 ; Kanne- 
liya, 6-15 Aug 1975, Panawatta, II 9; 
Kanneliya, 500 ft [ca. 150 m], 21-22 Apr 
1973, Baumann & Cross, at blacklight, 1 
9 ; Kanneliya Jungle, Udugama, 400 ft [ca. 
120 m], 6-12 Oct 1973. Krombein et al.. at 
blacklight, 1 9: Kanneliya Jungle, II mi. 
[ca. 18 km] E. Udugama, 11 Oct 1973, 
Krombein et al., at blacklight. 4 9 ; Sinha- 
raja Jungle, Kanneliya section, 13-16 Jul 
1978, Krombein et al., blacklight trap, 1 9: 
Sinharaja, above Enselwatta, 3,000 ft [ca. 
900 m], 20 Apr 1973, Baumann & Cross, 
1 9: Hiniduma, 22 Oct 1973, M. & B. Rob- 
inson, collected at black lisht. 1 9. Ham- 


bantota Dist., Badagiria Tank. 6.5 mi |ca. 
10.5 km] N Hambantota. 75 ft. (ca. 2.^ m). 
27 Oct 1970. O. S. Flint. Jr.. 1 9. Kalutara Agalawatta, 24 Jul 1975. Huang et 
al., light trap, 1 9. Kegalle Dist. Kelani 
Ganga, Kitulgala. 500 ft [ca. 150 m]. 12 
Mar 1973, Baumann & Cross, at blacklight, 
5 9. Monaragala Dist., Menik Ganga. Sella 
Kataragama, 150 ft. (ca. 45 m). 24 Oct 
1970, O. S. Flint, Jr., 1 9. Ratnapura Dist.. 
Walawe Ganga, Embilipitya, 100 ft. (ca. 30 
m), 21 Oct 1970, O. S. Flint. Jr., 2 9; dam 
site, Udawalawe, 250 ft. (ca. 75 m). 19 Oct 
1970, O. S. Flint, Jr.. 5 9: Gilimale. 17-18 
Jun 1976. Krombein et al.. light trap, 1 9; 
Gilimale, lumber mill, I 15 ft [ca. 35 m], 
20-25 Oct 1976, Hevel et al., blacklight, I 
9; Sinharaja Jungle, 2 mi. [ca. 3 km| S 
Weddagala, 8-12 Feb 1977, Krombein et 
al.. 2 9. 

Macrostemiim krombeini Flint, 
new species 

(Fig. 45) 

The holotype of this species was first be- 
lieved to be a variant of M. sepultum. but 
with the discovery of several new collec- 
tions unquestionably M. sepultum. this now 
is untenable. The color pattern suggests a 
relationship to M. sepultum. but the pattern 
is reduced to only a single small spot on 
the anterior margin of the forewing and an 
elongate mark from the posterior margin. 
The apparent black stripes along the front 
and hind margins of the forewing. so ap- 
parent in Fig. 45, are an artifact of poor 
spreading; they are due to the wing being 
folded at these spots. 

The specimen was collected by net in a 
heavily forested, bushy area by a very small 
streamlet. My field note indicates that a red- 
bodied macronematine was collected, the 
red now has faded to a tawny color. 

Male. — Unknown. 

Female. — Length of forewing 10 mm. 
Color generally fuscous with white marks. 
Head tawny frontally and dorsally, with 
frontal cuticle between eyes and central 
wart with bluish sheen: antenna with scape. 

pedicel, and first flagellar segment ventrally 
tawny, first flagellar segment dorsally and 
next 4 flagellar segments infuscate. remain- 
ing segments stramineous (missing beyond 
segment 15). Thoracic cuticle tawny: legs 
tawny, tarsi stramineous; midtibia slightly 
broadened and slightly concave mesally, 
tarsus unmodified. Forewing (Fig. 45) ba- 
sically fuscous with creamy-white marks; 1 
small spot at midlength from anterior mar- 
gin. 1 elongate stripe from posterior margin 
at '/3 length. Hindwing infuscate, a very 
small paler spot at stigma. 

Material examined. — Holotype, female: 
SRI LANKA, Kandy Dist.. Peradeniya 
[near top of hill behind University |, 1,700 
ft [ca. 520 m], 13 Nov [not Oct as labelled] 
1970, O. S. Flint, Jr. (NMNH). 

Etymology. — I dedicate this species to 
Dr. Karl V. Krombein of the National Mu- 
seum of Natural History. Washington, in 
gratitude for all his efforts originating and 
directing the survey of the Sinhalese insect 

Macrostemiim malickyi Flint, 
new species 

(Figs. 13-18, 46, 48) 

This distinctive new species is. in mac- 
ulation. most like M. sepultum. in that both 
have black wings, marked with a few, dis- 
crete, white spots. However, M. malickyi 
has four such spots, and M. sepultum has 
six. There are only two anterior marginal 
spots in M. malickyi and the outermost pos- 
terior mark extends from the wing margin 
anteriad into the gap between the two an- 
terior marks, a pattern quite different from 
that of M. sepultufu. The outline of the tip 
of the phallus in M. malickyi is also unique 
among the insular species. 

Male. — Length of forewing 8.5 mm. Col- 
or fuscous with white marks. Head strami- 
neous: frontal cuticle between eyes and 
central wart and posteriad on vertex, dark- 
ened: antenna with scape, pedicel and basal 
third of flagellum, stramineous, apical V2 of 
flagellum. infuscate. Thoracic cuticle shin- 
ing fuscous: legs stramineous except meta- 


Figs. 13-24. Male genitalia. Mi-li. Mainystemiim malickyi. L3, Lateral. 14, Dorsal. 15. Inferior appendage, 
postero ventral. 16. Phallus. lateral. 17. Tip of phallus, lateral. 18. Same, ventral. 19-24, M. nebulosiim. 19, 
Lateral. 20. Dorsal. 21. Inferior appendage, posteroventral. 22. Phallus, lateral. 23. Tip of phallus, lateral. 24, 
Same, ventral. 

tibia darkened. Forewing (Fig. 46) fu.scous 
with 4 white spots, 2 of which extend pos- 
teriad from costal margin (posterior seg- 
ment of outermost narrowly separated from 
main mark in holotype) and 2 extending an- 

teriad from posterior margin with outermost 
extending between 2 anterior spots, other 
more basad. Hindwing infuscate with an- 
terior margin and stigma white. Genitalia: 
Ninth seszment with anterolateral mariiin ir- 



regularly produced with large, rounded an- 
teroventral lobe, posteroventral margin 
nearly vertical; posterior margin with scat- 
tered, elongate setae, not much denser dor- 
solaterally. Tenth tergum with dorsobasal 
setal wart diffuse, apex rounded; in dorsal 
aspect with apex rounded, slightly curved 
laterad. Inferior appendage elongate, paral- 
lel-sided; in posteroventral aspect with two 
segments of nearly equal length. Basal sec- 
tion of phallus about Vj total length of phal- 
lus, relatively high and at right angle to 
stem of phallus; apex broadly bulging ven- 
trally before apex, and slightly elevated 
dorsally; apex with large, lip-like process 

Female. — Length of forewing 7 mm. 
Coloration as in male, except posterobasal 
white mark of forewing much smaller (Fig. 
48). Midtibia and tarsus unmodified. 

Material examined. — Holotype. male: 
SRI LANKA. Galle Dist., Udugama, Kan- 
neliya Jungle, 400 ft [ca. 120 m], 6-12 Oct 
1973, K.V. Krombein et al., at black light 
(NMNH). Paratype: Same data, 1 ? 

Etymology. — 1 dedicate this species to 
Dr. Hans Malicky, Lunz am See. Austria, 
in gratitude for his help with this project 
and to honor all his work on the South- 
eastern Asian fauna. 

Macrosteiniiiii luullifariiim (Walker), 
new combination 

(Fig. 47) 

Hydropsyche luultifaria Walker 1852:1 15. 

Macroneina multifarium: Ulmer 1907b:96 
[wings in color]. — Betten and Mosely 
1940:202 [redescription. wings]. — Fi- 
scher 1963:191 [catalogue]. 

This species, as well as several others of 
its congeners, has not been found since its 
original description. 1 have now seen four 
more examples, one of which has been 
compared directly with the holotype, and 
found identical in appearance. The holotype 
now lacks most of its abdomen, and mine 
are all females, thus the male genitalia are 

unknown. The midlegs of these females are 
not at all flattened, as in many other species. 

The new sites are all in the wet. south- 
western corner of the island, at elevations 
probably between 50 and 150 meters. 

Male. — Unknown. 

Female. — Length of forewing 10-11 
mm. Color generally fuscous with cream 
colored marks. Head stramineous to tawny 
frontally, labrum fuscous, shining black 
dorsally between eyes; antenna with scape, 
pedicel, and first flagellar segment ventrally 
stramineous, first flagellar segment dorsally 
and next 10 flagellar segments infuscate, re- 
maining segments tawny. Thoracic cuticle 
shining fuscous; legs stramineous; midtibia 
and tarsus unmodified. Forewing (Fig. 47) 
basically fuscous with cream-colored 
marks; 3 yellow marks from anterior mar- 
gin, with various other pale marks apically 
and posteriorly. Hindwing infuscate with 
white stigma and basally directed streak in 
costal and subcostal cells. Abdomen appar- 
ently orangish dorsally. 

Material examined. — Ceylon, William 
Templeton, holotype (BMNH). 

SRI LANKA: Galle Dist., Sinharaja Jun- 
gle, Kanneliya section, 2-5 Oct 1980, 
Krombein et al., collected in Malaise trap, 
2 9. Ratnapura Dist., Sinharaja Jungle, 8 
Sep 1979, P. B. Karunaratne et al.. in Mal- 
aise trap, 2 ? . 

Macrosteinuiu nchiilosiiiii (Hagen), 

new combination 

(Figs. 19-24, 49-50) 

Macronema iiehiilosiim Hagen 1858:485. — 
Ulmer 1907a:61 [redescription, wing tips, 
wings in color]. — Ross 1952:34 [lecto- 
type]. — Fischer 1963:192 [catalogue]. 

This species is very poorly known, being 
recorded from only the original type series, 
a female paratype of which was figured in 
color by Ulmer ( 1907a). I here present pho- 
tographs of the wings of the male lectotype 
and a female paralectotype from the MCZ, 
figure the male genitalia, and record several 
new collections. The species appears to be 


one limited to higher elevations and con- 
sequently to higher rainfall. The types from 
Ramboda were stated by Hagen ( 1 859. p. 
487) to be from 3,500 to 4.000 feet (1.065- 
1,220 m) above sea level, and the Adam's 
Peak collection was made at 6.000 ft. 
(1.825 m.). 

Male. — Length of forewing 14-17 mm. 
Color pale brown with yellowish cast. Head 
flavescent with black mark between dorsal 
warts; antenna fuscous; maxillary palpus 
flavescent. Mesonotum flavescent with fus- 
cous markings anteriorly and laterally, leav- 
ing rectangular flavescent area centrally; 
legs flavescent (Adam's Peak series with 
most of head dorsally fuscous with pale 
warts, thoracic notae with cuticle entirely 
fuscous). Forewing (Fig. 49) with pale 
spots poorly contrasting with ground color, 
only dark stigmal spot contrasting (even 
this spot in series from Adam's Peak is 
barely noticeable). Hindwing clear. Genita- 
lia; Ninth segment with anterolateral mar- 
gin produced and angulate, posterolateral 
margin nearly vertical; posterior margin 
with few setae laterally, with dense brush 
of long setae dorsolaterally. Tenth tergum 
with dorsobasal setal wart small, apex 
rounded in lateral view; in dorsal aspect 
with apex slightly flared laterad. Inferior 
appendage elongate, with each segment 
slightly constricted near base; in postero- 
ventral aspect with basal segment nearly 
two-thirds of total length. Basal section of 
phallus relatively narrow, slightly more 
than one-third of total length of phallus; 
apex slightly rounded ventrally. and 
obliquely elevated dorsally; apex with dis- 
tinct, lip-like projection. 

Female. — Length of forewing 14 mm. 
Coloration either as in male, or forewing 
with darker ground color (Fig. 50). Midtibia 
and tarsus unmodified. 

Material examined. — Ceylon. [Ramboda. 
Nuwara Eliya Dist.|, Nietner, 6 lectotype, 
2 paralectotype (MCZ). 

SRI LANKA: Kandy Dist., Kabaragala, 
Nillomalai, 22-23 Mar 1975, S. & R B. Ka- 
runaratne, 6 6 . Nuwara Eliya Dist.. Adam's 

Peak, 6 mi [ca. 9% km] S Maskeliya, 6,000 
ft [ca. 1,825 m], 17 Feb 1970, Davis & 
Rowe, 6 cJ, 1 2. 

Macrostemum pseudoneura pseiidoneitra 


(Figs. 25-30. 51) 

MacnmcDici pseudoneura Brauer 1865: 
420.— Ulmer 1907b:95 [redescription, 
forewing pattern]. — Schmid 1958:108 
[distribution], — Fischer 1963:194 [cata- 
logue [. 

Miuroswuniiii pseudoneura: Chantara- 
mongkol and Malicky 1986:528 [cJ. fore- 
wing pattern. head[.— Malicky 1998:778 
[6. forewing pattern]. 

This has been the most frequently en- 
countered species of the genus in Sri Lanka, 
and often comes to light in large numbers. 
The Sri Lankan specimens were captured at 
lower elevations, most recorded elevations 
being between 30 and 150 m. only one col- 
lection being made at 305 m and another 
230 m. 

Martynov (1935) recorded this species 
from Mysore State in southern India and 
presented a drawing (Fig. 93) of its wing 
pattern. This pattern agrees with the type of 
Macronema ethelda Banks, also described 
from Mysore State. I have studied the type 
of M. ethelda and compared it to the ma- 
terial available of M pseudoneura from Sri 
Lanka. The continental example has a well- 
marked pale streak in the 2nd R, cell. This 
mark is lacking in the insular material, and 
in addition there are slight differences in 
their genitalia, especially the apex of the 
phallus. Considering the overall similarity 
in the color pattern between the two re- 
gions, it seems best to consider them sub- 
species, pending further study with more 
continental material. I therefore recognize 
M. ethelda as Macrostemum pseudoneura 
ethelda (Banks), new .status. 

Male. — Length of forewing 8-9 mm. 
Head stramineous, infuscate dorsally; fron- 
tal cuticle between eyes and central wart 


Figs. 25-36. Male genitalia. 25-30. Macnisteiniim lysvidloiifiini. 25. Lateral. 26. Dorsal. 27. Interior ap- 
pendage, posteroventral. 28, Phallus, lateral. 29. Tip of phallus, lateral. 30. Same, ventral. 31-36. A7. .\i-piilliiiii. 
31. Lateral. 32. Dorsal. 33. Inferior appendage, posteroventral. 34, Phallus, lateral. 35, Tip of phallus, lateral. 
36. Same, ventral. 


dark with bluish sheen; antenna and max- 
illary palpus stramineous. Pronotum stra- 
mineous; meso- and metanota and pleura, 
fuscous; legs stramineous. Forewing (tig. 
5 1 ) with 3 white marks from anterior mar- 
gin, innermost extending as oblique, trans- 
verse, white band completely across wing; 
series of smaller white marks apically and 
basally; wash of golden setae between most 
pale marks, leaving a narrow brown band 
suiTounding most marks; hindwing infus- 
cate apically. Abdomen fuscous. Genitalia: 
Ninth segment with anterolateral margin 
broadly produced, nearly vertical; postero- 
ventral margin slightly produced and slight- 
ly oblique; posterior margin with scattered, 
elongate setae, becoming more numerous 
dorsolaterally. Tenth tergum with dorsobas- 
al setal wart small, apex obtusely angulate 
in lateral view; in dorsal aspect with apex 
rounded, sides narrow, rodlike. Inferior ap- 
pendage elongate, parallel-sided; in poster- 
oventral aspect with apical segment only 
sightly shorter than basal. Basal section of 
phallus almost Vs of total length of phallus, 
relatively high and angled almost 90° to 
stem; apex broadly rounded ventrally, and 
obliquely elevated dorsally; apex oblique 
with small apicoventral, protruding angle. 

Female. — Length of forewing 8 mm. 
Coloration as in male. Midtibia and tarsus, 
strongly flattened, broadened and concave 
on inner face. 

Material examined. — SRI LANKA: Am- 
parai Dist., Ekgal Aru tank, 100 ft [ca. 30 
m], 19-23 Feb 1977, Krombein et al., 
blacklight, 2 9 ; Inginiyagala, 250 ft [ca. 75 
m], 21-24 Nov 1976, Krombein et al., 
black light. 2 9 . Anuradhapura Dist., Wild- 
life Society Bungalow, Hunuwilagama, 
Wilpattu, 200 ft [ca. 60 m], 10-19 Mar 
1970, Davis & Rowe, 1 (5,8 9; Irrigation 
Bungalow, Padaviya, 180 ft [ca. 55 m], 27 
Feb-9 Mar 1970, Davis & Rowe, 9 9 ; Pa- 
daviya, 180 ft [ca. 55 m], 2-8 Nov 1970, 
O. S. Flint, Jr, 11 d, 5 9; Padaviya, 180 ft 
[ca. 55 m], 19 May 1976, Krombein et al.. 
blacklight trap, 1 9; Sluiceway, Padaviya, 
170 ft [ca. 50 m]. 3 Nov 1970, O. S. Flint, 

Jr, 17 (5, 17 9; Balaya Wewa, nr. Padaviya, 
150 ft [ca. 45 m], 4-5 Nov 1970. O. S. 
Flint, Jr, 2 9. Badulla Dist., 5 mi [ca. 8 km] 
E Mahiyangana, I Apr 1971, R & R Span- 
gler, at black light, 2 9. [Batticaloa Dist.]. 
Rukam Wewa, [90 ft, ca. 25 m], 14 Mar 
1954, F Schmid, [at light|. 1 9. Colombo 
Dist.. Gampaha Bot. Gardens, 16 Apr 1980, 
Mathis et al., 1 6; Labugama. 400 ft [ca. 
120 m], 24 Aug 1973. G. Ekis, collected at 
black light, 1 S: [Wak Oya], Labugama, 
[173 ft. ca. 53 m], 7-9 Jan 1954, F Schmid, 
[by net], 1 6. [Kelani Ganga], Kaduwela, 
[50 ft, ca. 15 m], 3 Apr 1954, E Schmid, 
[at light], 4 9. Galle Dist., Kanneliya, 500 
ft [ca. 150 m], 21-22 Apr 1973, Baumann 
& Cross, at blacklight, 24 c5 , 3 9; Kanne- 
liya. 27 Apr 1980, Mathis et al., 3 6: Kan- 
neliya, 6-15 Aug 1975, Panawatte, 2 9; 
Kanneliya. 22-24 May 1975. Wood & Pet- 
ty, collected in blacklight trap. 4 J . 3 9 ; 
Kanneliya. 200 ft [ca. 60 m]. 15-17 Oct 
1976. Krombein et al., blacklight, 4 d, 4 
9 ; Kanneliya Section, Sinharaja Jungle, 2- 
5 Oct 1980, Krombein et al., collected at 
blacklight, 8 d, I 9; same, but 13-16 Jul 
1978. blacklight trap, 16.19: Kanneliya, 
Sinharaja Jungle. 9-10 Nov 1977, Kiom- 
bein et al., collected in Malaise trap, 1 S\ 
Kanneliya Jungle, 300 ft [ca. 90 m], 28 Jul 
1973, G. Ekis, collected at blacklight, 8 <?, 

9 9; Kanneliya Jungle, 11-16 Jan 1975, 
Krombein et al.. blacklight, 2 9; Udugama, 
Kanneliya Jungle, 400 ft [ca. 120 m]. 6-12 
Oct 1973, Krombein et al., at black light, 4 
(5,79; same, but in Malaise trap, 1 9 ; 1 1 
mi |ca. 18 km] E Udugama, Kanneliya Jun- 
gle. 1 1 Oct 1973, Krombein et al., at black 
light, I (5, 5 9; Hiniduma, 22 Oct 1973. 
M. & B. Robinson, collected at black light, 
1 9. Kandy Dist.. Roseneath. Kandy, 25 
Mar 1971. P & R Spangler, in Malaise trap, 

10 (5; Hasalaka, 16-19 Feb 1977, Krom- 
bein et al., blacklight trap, 2 cj, 2 9; Aluth- 
nuwara, Hasalaka, 19 Apr 1964, [collector 
unknown], at light. 1 9. Kegalle Dist, La- 
vant Estate, nr. Yatiyantota, 80 ft [ca. 25 m], 
19 Nov 1970, O. S. Flint, Jr, 2 (5; Kitulgala. 
150 ft [ca. 45 m|, 30 Sep 1970, O. S. Flint. 



Jr. 1 (5, 2 9: Kitulgala. 31 May 1975. Wood 
& Petty, collected in blacklight trap. 12 c5. 
4 9; Kitulgala Resthouse. 150 ft [ca. 45 m]. 
3-5 Feb 1979. Krombein et al.. blacklight 
trap. 26 6, 32 9; same, but 24-26 Oct 
1977. 12 <5. 5 9; same, but 10-12 Oct 
1980. coll. in UV trap. 4 (5. 8 9: Kclani 
Ganga. Kitulgala. 500 ft |ca. 150 m|. 12 
Mar 1973. Baumann & Cross, collected at 
black light. 1 d. 1 9. [Kelani Ganga), Ki- 
tulgala. [750 ft. ca. 230 m]. 2 Mar 1954. F. 
Schmid. [at light], 1 9. [Monaragala Dist.]. 
Bibile. 4 May 1974. Gans & Prasanna. 1 9. 
Ratnapura Dist.. Panamure, 500 ft [ca. 150 
m]. 15-21 Oct 1970, O. S. Flint, Jr, 1 d; 
Uda Walawe. 300 ft [ca. 90 m|, 1 Aug 
1973. G. Ekis. collected at blacklight. 10 9; 
Giiimale, 17-18 Jun 1976. Krombein et al.. 
light trap, 1 9; Giiimale. Induruwa Jungle, 
collected near Kaluganga, 5-7 Feb 1977, 
Krombein et al.. blacklight trap, 2 9; Kalu 
Ganga. Induruwa Jungle. 1.000 ft [ca. 305 
m). 23 Mar 1973. Baumann & Cross. 1 6. 
[tributary to Kalu Ganga. near Kiriella[. 
Ratnapura. [80 ft. ca. 25 m). 3 Feb 1954. F 
Schmid. [at light]. 1 9. Vavuniya Dist.. Ir- 
rigation canal. Parayanalankulam. 25 mi 
|ca. 40 km) NW Medawachchiya. 100 ft 
|ca. 30 m|. 20-25 Mar 1970, Davis & 
Rowe, 1 9. 

MacrosU'iuiiin sepiiltiiin (Hagen), 
new combination 

(Figs. 31-36, 52) 

Mdcroiu'inci scpitliiiiu Hagen 1859:209. — 
Ulmer 1907a:60 [redescription, wings, 
wings in color]. — Ross 1952:35 [lecto- 
type). — Fischer 1963:197 [catalogue]. 

As with M. nchulosiini. this species has 
been known only from the original type se- 
ries, a male paratype of which was figured 
in color by Ulmer (1907a). I here present 
photographs of the wings of a female par- 
alectotype from the MCZ, figure the male 
genitalia, and record several new collec- 
tions. The original series was from the wet. 
central highlands, at 1.065-1.220 m. the 
new material was l\>inid at elevations of 

around 1 50 m in the wet. southw estern cor- 
ner of the island. 

Male. — Length of forewing 8-8.5 mm. 
Color fuscous with white marks. Head stra- 
mineous frontally. shining black otherwise; 
frontal cuticle between eyes and central 
wart with bluish sheen; antenna with scape 
and pedicel tawny, first 3-5 flagellar seg- 
ments slightly darkened, remaining seg- 
ments stramineous. Thoracic cuticle shining 
fuscous; legs fuscous with tarsi strami- 
neous. Forewing (Fig. 52) fuscous with 6 
white spots, 3 lying on the costal margin, 
and 3 toward posterior margin. Hindwing 
infuscate with anterior margin and stigma 
white. Genitalia: Ninth segment with an- 
terolateral margin produced and angulate, 
posteroventral margin produced, slightly 
oblique; posterior margin with few, elon- 
gate setae, becoming denser dorsolaterally. 
Tenth tergum with dorsobasal setal wart 
large with many short setae, apex rounded 
in lateral view; in dorsal aspect with apex 
produced laterad. Inferior appendage elon- 
gate, parallel-sided; in posteroventral aspect 
with apical segment only about -A total 
length. Basal section of phallus relatively 
narrow, only '/, of total length of phallus; 
apex broadly rounded ventrally and dorsal- 
ly; apex nearly vertical with a small lip-like 
lobe below midheight. 

Female. — Length of forewing 8 mm. 
Coloration as in male. Midtibia and tarsus 

Material examined. — Ceylon, [Rambo- 
da], Nietner. 6 lectotype, 9 paralectotype 

SRI LANKA: Galle Dist.. Kanneliya 
Jungle, Udugama, 400 ft [ca. 120 m|, 6-12 
Oct 1973. Krombein et al., at blacklight. 3 
6.29: Kanneliya, 22-24 May 1975, Wood 
& Petty, collected in Malaise trap, 1 6. 

Macrostcnmiu spk'iii/idiim (Hagen) 
(Figs. 37-42, 53-54) 

Miicroiu'iiia splendidiim Hagen 1858: 
484.— Ulmer 1907b:87 [redescription, 
forewing. wings in color]. — Ross 1952: 
35 I lectotype].— Schmid 1958:108 [dis- 


Figs. 37 — \2. M;ile genitalia. Macni.stciniiin splcinluliiii]. 37. Lateral. 3X, DorsaL 39. Inferior appendage, 
po.steroventraL 40. Phallii.s, lateral. 41. Tip of phallus, lateral. 42. Same, ventral. 

tiibiition. variation]. — Fischer 1963:197 
Macrostemiim spleiulidimi: Malicky 1998: 
778 [(?, wing pattern]. 

This species has been taken a number of 
times, but never in a large series. As re- 
marked by Schmid (1958), the species is 
quite variable in color, no two of the spec- 
imens before me appearing exactly the 

The species is known from the central 
highlands at elevations approaching 1,000 
m. and in the southwestern quadrant of the 
island at elevations from almost 1,000 m 
down to .^."S m. All of the known sites, how- 
ever, are in high rainfall areas with precip- 
itation averaging 100 inches (2,500 mm) 
per year or higher. 

Male. — Length of fore wing 11-15 mm. 
Color goldenyellow marked with fuscous. 
Head goldenyellow, cuticle black between 
dorsal warts; antenna with scape, pedicel 

and outer face of first flagellar segment, 
golden following ca. 12 segments fuscous, 
remaining segments pale. Cuticle of meso- 
and metanota and pleura black; mesoscu- 
tellum golden yellow. Legs goldenyellow. 
Forewing (Fig. 54) with 3 golden-yellow 
marks from anterior margin, posteriorly and 
apically with extensive golden-yellow 
marks on a fuscous background. Hindwing 
infuscate with white stigma and basally di- 
rected streak in costal and subcostal cells. 
Abdomen fuscous, with golden-yellow 
middorsal stripe. Genitalia: Ninth segment 
with anterolateral margin broadly produced, 
rounded, postero ventral margin produced 
ventrally, oblique; posterior margin with 
scattered, elongate setae, becoming very 
numerous dorsolaterally. Tenth tergum with 
dorsobasal setal wart small, apex with small 
excision at midheight; in dorsal aspect with 
apex shallowly emarginate, apicolateral 
margin produced and rounded. Inferior ap- 



Figs. 43-48. Forewings. 43. Macrosteiuiiin IninimJi. S . Kanneliya. 44, M. iiulisiiiiciitm. 9. Udawalawe. 4.i. 
M. kroinheini. 9 holotype. Peradeniya. 46. A/, imilukxi, 6 holotype, Udugama. 47. A/, iimllifuriiim, 9 compared 
with holotype, Sinharaja. 4S. A/, nialickyi. 6 paratypc, Lldiigania. 

pendage elongate, parallel-sided; in pos- 
teroventral a.speet with apical segment V3 of 
total length. Basal section of phallus about 
V3 of total length of phallus: apex broadly 

rounded ventrally. and obliquely elevated 
dorsally; apex nearly vertical with large, 
ventral, lip-like projection. 

Female. — Length of forewing 11-15 


Figs. 49-54. Forewings. 49, Mucrosleiimin iie/'u/nsiiiii. i lectotype. Rainboda. 50. M. nchulosum. 9 para- 
lectotype. Ramboda. 51. A/, pseudoneura. 6, Padaviya. 52. M. sepidtiiin. 9 paralectotype. Ramboda. 53. M. 
splemiuliim. 9 paralectotype. Ramboda. 54. M. spleiididum. S, Tunmodera. 

mm. Coloration as in male (Fig. 54). Mid- 
tibia and tarsus flattened, broadened, con- 
cave on inner face. 

Material examined. -Ceylon. [Ramboda]. 
Nietner. S lectotype, 9 paralectotype 

SRI LANKA: Colombo Dist.. Tunmod- 
era, 200 ft [ca. 60 m], 17 Nov 1970. O. S. 
Flint, Jr., 1 (J, 2 9. Galle DisL, Kanneliya, 
500 ft [ca. 150 m], 21-22 Apr 1973, Bau- 
mann & Cross, at blacklight, 1 6 . [Kandy 
Dist.|. Karambaketiya off Memure (Mimu- 



ie|. Knuckles Range. 8 Mar 1970. P. B. Ka- 
runaratne. at light. 2 6 . Matale Dist.. Hunas 
Falls. Elkaduwa. 3.000 ft |ca. 91.'^ m). 5 Apr 
1973. Baumann & Cross. 1 9. Matara Dist.. 
Deniyaya. 1.000 ft [ca. 305 m]. 20 Apr 
1973, Baumann & Cross, at black light. 1 
9. Ratnapura Dist.. Gilimale. lumber mill. 
115 ft [ca. 35 m). 20-25 Oct 1976. Hevel 
et al.. blacklight. 1 6. Bultota Pass. 3.000 
ft |ca. 915 m|. 18-19 Apr 1973. Baumann 
& Cross, at black light. 1 6: |Kirinda Ela]. 
Wattepanguwa. [1.500 ft. ca. 450 m]. 18 
Feb 1954. F Schmid. [at light [. 1 9. 


I thank Smithsonian's "Ceylon Insect 
Project" for funding my collecting trip to 
the island in 1970. These monies also al- 
lowed numerous other workers from the 
Museum and other institutions to visit and 
collect on the island; most of these workers 
collected caddisflies in addition to their tar- 
get groups. These collections provided the 
most valuable items from the island and the 
backbone of the material reported here. 

Dr. R C. Barnard. The Natural History 
Museum. London. England (BMNH). Dr. R 
D. Perkins. Museum of Comparative Zo- 
ology. Cambridge, MA, USA (MCZ), and 
Dr. Hans Malicky, Lunz-am-See, Austria 
(HMPC), all loaned valuable material from 
their respective collections. All material not 
marked otherwise, is deposited in the col- 
lection of the National Museum of Natural 
History, Washington. DC, USA (NMNH). 

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105(4). 2003, pp. 832-846 


Gary L. Miller and Douglass R. Miller 

Systematic Entomology Laboratory. Plant Science Institute. Agricultural Research Ser- 
vice, U.S. Department of Agriculture, Bldg. 005, BARC-West. 10300 Baltimore Avenue, 
Beltsville, MD 2070.5 U.S.A. (GLM e-mail:; DRM e-mail: 

Abstract. — We provide a compilation of 147 species of soft scales that are considered 
cither pests or represent a threat to United States agriculture. Included for each species, 
where applicable, is reference to origin and date of introduction if applicable, establish- 
ment in the United States, pest or threat status in the United States along with a validation 
citation, principal hosts, and biogeographical region of origin. 

Ke\ Words: Coccidae, soft scales, invasive species, biological control, quarantine, ag- 
riculture, forestry, horticulture, ornamentals, fruit trees 

Invasive (non-native) species of insects 
represent an increasing concern to the Unit- 
ed States. If non-native species become 
pests, the consequences include loss of pro- 
duction, diminished product quality, pro- 
duction cost increases, flexibility decreases 
in production/management decisions, in- 
creased risk of human disease, and dam- 
aging environmental and aesthetic effects 
(Huber et al. 2002). Various insects of ag- 
ricultural concern have been identified as 
potentially damaging if introduced or rein- 
troduced into the United States (Huber et 
al. 2002). Invasive species and potentially 
dangerous species of mealybugs (Pseudo- 
coccidae) have been recently examined as 
they pertain to agriculture in the United 
States (Miller et al. 2002). The Coccidae or 
soft scales, like all scale insects, are plant 
feeders. A few species of soft scales are 
valuable to man as biological control agents 
of noxious weeds, however, many are pests 
of economially important plants. 

Because invasive species of insects rep- 
resent a major concern to U.S. agriculture. 

we have investigated several parameters 
concerning invasive soft scales. Objectives 
of this paper are: I) To develop a prelimi- 
nary world list of the pest soft scales; 2) 
provide a list of pest soft scales introduced 
to the continental United States; 3) to de- 
termine which species in the previous two 
objectives are either introduced or native to 
the continental United States; 3) examine 
data provided by the United States Depart- 
ment of Agriculture. Animal and Plant 
Health Inspection Service — Plant Protec- 
tion and Quarantine (USDA. APHIS-PPQ) 
concerning the most commonly intercepted 
soft scales at the United States ports-of-en- 
try; and 5) using all of this information, try 
to predict which soft scales are the most 
likely candidates for future invasions into 
the continental LInited States. 

Materials and Methods 

Executive Order 13112 established the 
National Invasive Species Council and pro- 
vided a definition of an invasive species as 
"... a species that is 1 ) non-native (or 


alien) to the ecosystem under consideration 
and 2) whose introduction causes or is like- 
ly to cause economic or environmental 
harm or harm tt) human health." This def- 
inition not only includes species alien to the 
United States but also encompasses native 
species. The dehnition also has an econom- 
ic or potentially economic component. By 
this definition, the tulip tree scale, Toiinie- 
yella liriodeiuhi (Gmelin), would be an ex- 
ample of an invasive species in the United 
States even though it likely is native. Our 
definition is based on Miller et al. (2002) 
and is more simplistic. They considered in- 
vasive species to be those that are non-na- 
tive [also introduced, nonidigenous, exotic, 
alien or invasive (Huber et al. 2002)] to the 
United States regardless of economic harm. 

We have used a broad definition of the 
term "pest"" to create a table of pest soft 
scales of the world (Table 1 ). If a soft scale 
is described in the literature as either a pest, 
causing damage, requiring control, or of 
economic importance, we have included it 
in the list. A pest species as defined by 
some authors (e.g., Ebeling 1959, Pfeiffer 
1997) was regarded as any record of a soft 
scale on certain economic hosts. Ebeling's 
(1959) justification was based on his con- 
sideration that some species not of econom- 
ic importance become major pests through 
adaptation or by being transferred to re- 
gions of lower environmental resistance. 
However, Pfeiffer"s (1997) inclusion of Eii- 
k'canium lespedezoe Danzig as a pest of de- 
ciduous fruit trees is probably an error be- 
cause the only known host record of this 
species is Lespedezci bicolor Tuicz. (Danzig 

Our perspective for this paper has fo- 
cused on the impact or potential impact of 
a pest soft scale on agriculture in the con- 
tiguous United States. For example, Cero- 
pkites psidii (Chavannes) is known only 
from Psidiiim spp. Therefore, because gua- 
vas are not widely grown in the contiguous 
United States, it is considered to have rel- 
atively minor pest potential in the United 
States even thouuh it mav be far more im- 

portant in areas of the world where guava 
is of greater economic importance. Con- 
versely, Ceroplastes japoniciis Green, oc- 
curs on many different agricultural plants 
that are economically important in the Unit- 
ed States and it is therefore considered a 
major threat. The term "threat" is used for 
species that are considered pests but do not 
occur in the United States. Determination 
of the date of introduction in the U.S. was 
established either from literature records or 
from the oldest collection record in the Na- 
tional Entomological Ctillection of the Na- 
tional Museum of Natural History, in Belts- 
ville, Maryland. In at least one case (e.g., 
Eriopcltis festucae Boyer de Fonscolombe), 
the first literature record in the U.S. is Patch 
(1905) but the oldest collection record is 
1899. Obviously, these dates are estimates 
of the date when a species first invaded the 
United States. 

It is difficult to determine the zoogeo- 
graphic area of origin for some species. 
Some distributional records for Ceroplastes 
spp. are from Qin et al. (1998) and Coccus 
spp. are from Gill et al. (1977). It is not 
always clear whether an invasive species is 
from the Old or New World. In some cases, 
we have simply made a supposition based 
on the current distribution of the species, 
the distribution of what appears to be its 
closest relatives, and the natural distribution 
of its primary host plants. We have used the 
same criteria to determine if a particular 
species is native to the United States. Our 
use of the terms polyphagous, oligopha- 
gous, and monophagous has been slightly 
modified for the cunent paper and are here- 
by defined for those species that have great- 
er than 10 host-plant families, 3-10 host- 
plant families, and 1-2 host-plant families 


Table 1 provides information on 147 spe- 
cies of soft scales. The table includes one 
species [Prococcus acutissimus (Green)] 
that has been introduced and established 
into the United Stales but is not considered 


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Soft Scale Introductions in the U.S.: 1870-1999 


Fig. 1 . Soft scale 




12 3 4 5 6 7 

Number of Introduced Species 

introductions in the United Slates Ironi 1870-1999. 

9 10 

a pest. Therefore, we estimate that there are 
146 species that are either pests or represent 
threats to U.S. agricuhure. Of the 66 soft 
scales considered pests in the U.S., 25 are 
either native or possibly native species. 
Therefore, 41 of the soft-scale pests in the 
country are invasive. Based on Coccidae in- 
formation presented in ScaleNet (Ben-Dov 
2002), there are 105 species of soft scales 
in the United States; thus, the invasive com- 
ponent of the soft-scale fauna in the United 
States is approximately 39%. This percent- 
age is much higher than the 13% of inva- 
sive species in the mealybug fauna of the 
U.S. (Miller et al. 2002). With the exception 
of the 1960's and the 1980"s, at least one 
species of soft scale (Fig. 1 ) has been in- 
troduced every decade since the 1870"s. 
The greatest number of introduced soft 
scales (nine species) occurred between 
1900 and 1909. Two periods, the 1890"s 
and the 1920's, witnessed the introduction 
of six species. More than 73%- of the in- 
vasive soft scales were introduced in the 

Hrst 69 years of record keeping (1870- 

A summary of the region of origin of all 
invasive soft scales in the U.S. is as fol- 
lows: Palearctic Region, 13; Neotropical 
Region, 11; Oriental Region, 9; Afrotropi- 
cal Region, 5; Nearctic Region outside of 
the U.S., 3; and Australasian Region, 0. Of 
all of these species in the U.S., only Pro- 
coccus aciitissimits (Green) from the Ori- 
ental Region is not considered a pest. 

Examination of the region of origin for 
pest soft scales worldwide provides the fol- 
lowing results; Palearctic, 46; Nearctic, 29; 
Oriental, 25; Neotropical, 23; Afrotropical, 
18; and Australasian, 6. Host characteristics 
of these pests include 38% polyphagous. 
33% oligophagous, and 29% monophagous. 
Based on the characteristics of zoogeo- 
graphic regional distributions of the highest 
number of soft-scale pests (Palearctic, Neo- 
tropical, and Oriental Regions, respective- 
ly), and greatest frequency of host plants 
(polyphagous and oligophagous), a list of 



the species most likely to invade the United 
States was determined. Those species likely 
to invade from the Palearctic Region in- 
clude Eidecaniiim doiiglasi (Sulc). Lichten- 
sia vibunii Signoret, Palaeolecaniiim hitii- 
berciiUitum (Signoret). Parthenolecaniiini 
nifuhini (Cockerel!). Piilvinaria aurantii 
Cockerell, Piilviiuiria regalis Canard, and 
Rhodococcus turanicus (Archangelskaya). 
Those species likely to invade from the 
Neotropical Region include Ceroplastes 
hergi Cockerell. Ceroplastes grandis Hem- 
pel, Philephedra hroadwayi (Cockerell). 
Pidviiiaria ficus Hempel. and PulvinarUi 
flarescens Brethes. Those species likely to 
invade from the Oriental Region include 
Ceroplastes actiniformis Green. Ceroplas- 
tes japoniciis Green. Ceroplastes pseiido- 
cerifents Green. Coccus discrepans 
(Green), Didesinococcus unifasciatiis (Ar- 
changelskaya). Drepanococciis chiton 
(Green). Megapiilvinaria maxima (Green), 
Protopulvinaria longivalvata Green, and 
Pidviiiaria polygonala Cockerell. 

USDA, APHIS-PPQ records from the 
past five years also were searched to deter- 
mine which intercepted species of soft 
scales pose the greatest threat. A list of the 
top seven species most frequently inter- 
cepted at U.S. ports-of-entry are Ceroplas- 
tes japoniciis Green, Coccus moestus De 
Lotto, Philephedra hroadwayi (Cockerell), 
Protopulvinaria longivalvata Green, Pulvi- 
luiria polygonata Cockerell. Tillancoccus 
mexicaiius Ben-Dov. and Udinia catori 
(Green). Comparison of the two lists re- 
veals the following four species common to 
both: Ceroplastes japoniciis, Philephedra 
hroadwayi. Protopulvinaria longivalvata. 
and Pidvinaria polygonata. While the pos- 
sibility exists that other soft scales could 
invade the U.S.. we suggest that the four 
aforementioned species are the most plau- 
sible candidates as the next invasive soft 
scales into the United States. 


Our data indicate that the decade starting 
in 1900 had tlie laraest number of soft scale 

insect introductions into the U.S. Although 
Miller et al. (2002) speculated that high 
number of mealybug interceptions may be 
linked to detection strategies and proce- 
dures developed in conjuction with the 
Plant Quarantine Act in 1912, the number 
of soft scale introductions is not consistent 
with this hypothesis. 

The introduction of even a single species 
is of concern to U.S. agriculture. Huber et 
al. (2002) hypothesized that if a pest can 
enter the United States, over time there is a 
strong likelihood for establishment. As a re- 
sult, they believed more appropriate and 
cost-effective quarantine procedures must 
be developed. We think one step in this pro- 
cedure is to identify those species of insects 
which pose the greatest threat. 


We are grateful M. Gimpel (University of 
Maryland, College Park, MD) for assis- 
tance with Coccidae species records for the 
United States and J. Cousins (USDA, 
APHIS-PPQ. Riverdale. MD) for Coccidae 
interception records at U.S. ports-of-entry. 
We also thank P. Lambdin (Dept. of Ento- 
mology and Plant Pathology. The Univer- 
sity of Tennessee. Knoxville. TN), T Henry 
(USDA. ARS. Systematic Entomology 
Laboratory. Washington. DC), and R. 
Ochoa (USDA, ARS, Systematic Entomol- 
ogy Laboratory, Beltsville, MD) for helpful 
suggestions and comments with the manu- 

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105(4). 2003. pp. 847-858 


Britt a. BliN"! ard 

Biology Department. Dauby Science Center, Ursuline College, Pepper Pike, OH 44124, 
U.S.A. (e-mail; bbunyard^' 

Abstract. — For this study, 134 species from 30 families of Basidiomycete fungi and 19 
species from 1 1 families of Ascomycete fungi were collected from different sites in north- 
eastern Ohio. Adult flies were reared from 87 separate fungal collections (basidiocarps or 
ascocarps = "mushrooms"). During this study, mycophagous species from a number of 
families were found (Tipulidae, Mycetophilidae, Sciaridae, Cecidomyiidae, Phoridae, Pla- 
typezidae, Chloropidae, Drosophilidae); the two most common fly species were Drosoph- 
ila falleni Wheeler and Leiicophenga varia Walker, both Drosophilidae. Less commonly 
found were Drosopliila tripitiictata Loew, Drosophila piitrida Sturtevant, and Mycodro- 
sophila ckiytouae Wheeler and Takada. Frequently, several species of Drosophilidae were 
found cohabiting the same mushroom. Unless factors are in operation to prevent com- 
petition (niche paititioning, predation, parasitism), mycophagous Diptera may pose a chal- 
lenge to the Competitive Exclusion Principle. Preliminary evidence suggests parasitism 
and predation by other species of arthropods may play a role in reducing competition. 

AVv Words: Competitive Exclusion Principle, fungi-feeding flies, mushrooms 

The sporophores of macrofungi (i.e., 
mushrooms of Basidiomycetes and Asco- 
mycetes) are analogous to vascular plants 
in a number of ways: they are immobile, 
frequently contain chemical toxins, have 
few physical defenses, and have members 
that may be ephemeral or perennial in lon- 
gevity (Hanski 1989). Fungi are found in 
virtually every ecological niche, and the 
sporophores of many groups of macrofungi 
serve as hosts of mycophagous Diptera. De- 
spite the frequency and diversity of Diptera 
that inhabit mushrooms, few studies have 
been concerned with mycophagous species. 
Most such ecological studies were conduct- 
ed several decades ago (Buxton 1960, Pie- 
lou 1966, Pielou and Mathewman 1966, 
Pielou and Verma 1968. Valley et al. 1969, 
Papp 1972, Shorrocks and Wood 1973) and 
often gave only anecdotal accounts of adult 

flies occurring on mushrooms (Patterson 
1943, Valley et al. 1969, Graves and Graves 
1985). not verifying true mycophagy. Still 
other studies have included flies as myco- 
phages, where only one or two adults 
emerged from fungal material (Buxton 
I960. Valley et al. 1969. Hackman and 
Meinander 1979, Graves and Graves 1985, 
Yakolev 1993), or had emerged solely from 
decaying mushrooms (thus, possibly only 
scavenging) (Frouz and Makarova 2001) 
and therefore did not establish a strong eco- 
logical association (i.e., food substrate, site 
of overwintering, etc.) with fungal sporo- 
carps. Still other studies have focused on 
the evolution of tolerance of the amanitin 
toxin (present throughout the Basidiomy- 
cete genus Amanita Persoon) tolerance 
(Jaenike et al. l983,Jaenike 1985) or mech- 
anisms for aggregation of adult flies on 
mushrooms (Jaenike et al. 1992). 



Overlooked by nearly all studies has 
been the ecology of mushroom-feeding spe- 
cies, and little is known about larval stages, 
feeding preferences, seasonality, or geo- 
graphic range for numerous species (Bux- 
ton 1960; Graves and Graves 1985: Bun- 
yard and Foote 1990a, b). The larval stages 
of many of the rarer species of mycopha- 
gous flies have never been described. Most 
mycophagous flies are probably generalists 
and not specific to any species of fungus, 
as fungal hosts are considered too patchy 
and/or ephemeral, or are scavengers, feed- 
ing on all sorts of decaying organic material 
(Jaenike 1978a, b). Oligophagous and spe- 
cialist species are uncommon in nature and 
in the literature; many of the records re- 
porting monophagy are likely artifacts of 
insufficient sampling (Hanski 1989). It has 
been postulated that mycophagous Diptera 
probably arose from ancestral detritivores 
(Bruns 1984). One of the largest groups of 
mycophagous Diptera, the family Droso- 
philidae, is considered to have evolved my- 
cophagy more than once from a common 
ancestor that was a detritivore or fed on 
saprophytic yeasts (Courtney et al. 1990). 

For this study I attempted to determine: 
a) the families of Diptera that are truly my- 
cophagous. b) the existence of associations 
among families of Diptera with families of 
Basidiomycete host fungi, c) seasonality 
among mycophagous families of Diptera, 
and d) evidence of seasonality within a 
common mycophagous family, the Droso- 

Materials and Methods 

Sites were selected to obtain a diversity 
of mushroom species, as well as biotic and 
abiotic conditions, and consisted of mature 
forest, mixed mesophytic forest, urban for- 
est, and urban residential zones in north- 
eastern Ohio (Cuyahoga, Geauga, and Por- 
tage counties). Two of the study sites have 
been described previously: Towner's Woods 
near Kent. Ohio (Portage Co.) (Bunyard 
and Foote 1990a) and The West Woods, 

near Newbury, Ohio (Geauga Co.) (Bun- 
yard, in press). 

Fungi were collected throughout the 
growing season from March to November. 
2001. Many fungal species were collected 
more than once and/or on different dates or 
sites. Fungal material was placed in paper 
bags to prevent larvae of one sporophore 
from entering another sporophore. Different 
species were kept in separate bags; conspe- 
cifics from different sites also were kept 
separately. Fungal specimens were identi- 
fied using keys in Lincoff 1984, Arora 
1986. Phillips 1991. Smith and Weber 
1996, Bessette et al. 1997. To avoid inci- 
dental occurrences of Diptera with the fungi 
(for example, resting or hiding in crevices) 
only adults which actually emerged from 
larvae occuixing within the fungus were 

For rearing adult flies from fungal hosts, 
special rearing chambers were constructed 
as previously described (Bunyard and Foote 
1990a) and consisted of the bottom of a pe- 
tri dish ( 10 X 100 mm) to which had been 
added moistened vermiculite. The upper 
portion of the rearing chamber consisted of 
rigid clear plastic tubing (90 mm diameter) 
cut to various lengths. To the top end of 
each tube was glued a fine polyester mesh 
material. Fungal sporocarps were placed on 
the vermiculite substrate, and the upper por- 
tion of the chamber placed securely over 
the fungus, into the petri plate. The rearing 
chambers allowed the fungal specimens to 
remain in a somewhat natural condition. It 
was necessary to moisten the vermiculite 
substrate periodically to prevent desiccation 
of fungal material. As the sporocarps de- 
cayed, the substrate absorbed any excess 
moisture produced. 

Following emergence, adult Diptera were 
kept alive for at least 24 hr to allow for 
exoskeleton hardening (to facilitate identi- 
fication) and then killed in alcohol. Adult 
flies were dried and pinned for identifica- 
tion. Preserved specimens are in the Kent 
State University collection of Diptera. 



During this study 134 species from 30 
families of Basidiomycete fungi and 19 
species from 1 1 families of Ascomycete 
fungi were collected from different sites in 
northeastern Ohio. Adult flies were reared 
from 87 separate fungal collections (Table 
1). A few fungal species seemed to host 
only a single fly species per mushroom, but 
most were found to support more than one 
species of Diptera (Table 2). Five families 
of Diptera that include known or suspected 
mycophagous species (Tipulidae. Phoridae, 
Platypezidae, Chloropidae. Drosophilidae) 
were reared from fresh mushroom collec- 
tions (Table 2), confirming mycophagy (as 
opposed to scavengening rotting material). 
Additionally, members of the Mycetophili- 
dae. Sciaridae. and Cecidomyiidae also 
were seen (data not shown). Five other dip- 
teran families that were reared from fungi 
(Psychodidae. Ceratopogonidae. Stratio- 
myiidae. Anthomyiidae. Sarcophagidae) 
likely are larval predators or scavengers oc- 
curring only in decaying mushrooms (Table 
2). In some cases a single member of a dip- 
teran family emerged from fungal material 
but was not counted, as it was unclear if 
any strong ecological association with fungi 

All Diptera reared in this study emerged 
only from Basidiomycete species (Table 1 ). 
No evidence for monophagy was seen by 
any of the dipteran taxa. In general, the 
larger the sporocaips. the greater the num- 
ber of individuals, as well as diversity, of 
Diptera utilizing the mushroom host were 
seen, supporting previous studies (Bruns 
1984, Hanski 1989). Previous demonstra- 
tion of seasonality of mycophagous Diptera 
has been considered a function of season- 
ality of mushroom hosts (Hanski 1989) and 
was demonstrated here (Fig. 1 ). The highest 
number of emergences for all mycophagous 
taxa was seen in spring and fall; this cor- 
related to the highest numbers of mushroom 
fruitings (Fig. 1 ). 

More rearings of Drosophilidae were re- 
corded than those of any other dipteran 
family (Fig. I ). The highest number of 
emergences for drosophilid species was 
seen in spring and fall. Thus, seasonality of 
mycophagous Drosophilidae likely is due to 
seasonality of hosts. Most species of Dro- 
sophilidae are considered yeast-feeders 
(Patterson 1943). with the food material 
serving as a culture medium for the yeast. 
It is thought that all extant species of this 
family came from a common ancestor that 
was detritivorous (Courtney et al. 1990) and 
became selective for rotting substrates sup- 
portive of yeast growth, especially fruits. 
Phylogenetic evidence suggests that my- 
cophagy has arisen more than once within 
the family (Courtney et al. 1990). Today, 
most species of drosophilids feed on decay- 
ing fruit material, some are scavengers, and 
a few feed on fungi. All the species reared 
in this study (Drosophila fiilleni Wheeler, 
D. piithda Sturtevant, D. tripiinctata Loew, 
Mycodrosophila claytonae Wheeler and 
Takada, Leiicophengci vcirici Walker) are 
known to be mycophagous. During this 
study, the two most commonly reared spe- 
cies overall were D. falleni and L. varia 
(Fig. 2). These two species commonly co- 
inhabited basidiocarps, occasionally with 
three other less common drosophilids: D. 
tripitnctata. D. piitridu. and/or M. clayto- 
iHie. These findings support those of pre- 
vious studies (Grimaldi and Jaenike 1984). 
Some drosophilid species have been shown 
to dominate (D. tripuiutata) when compet- 
ing with other species (D. falleni and D. 
putrida) (Worthen 1989), although this 
clearly was not seen here. Likewise, Leii- 
cophenga species may dominate in smaller 
species of mushrooms (Worthen et al. 
1998). Several species of mycophagous 
species of Drosophilidae that were reared 
from fungi previously (Bunyard and Foote 
1990a) were not obtained in this study, in- 
cluding D. diincani Sturtevant, D. giittifeni 
Walker, D. testacea von Roser, and M. dim- 
idiata Loew. Little is known about the life 
history of D. diiiuani. Likewise, D. giitti- 


Table i. Species of macrofungi. serving as hosts for iiiyi. 

ophagous Diplera. arranged alphabeliLally by fungal 


? agaric sp. . 

Agaricaceae Agariciis arvensis Schaeffer 

AauiicKs ciiigiisnis Fries 
Agaricus hitorqiiis Quelel 
Agariciis cctmpeslris Linnaeus 

Agiiricus sp. 
Amanitaceae Amciniui flavoruhescens Atkinson 

Amiinita iiiiiscurin Persoon 
Amanila nihescens Persoon 

Bolbitiaceae Agrocybe praecox Persoon 

Boletaceae Boletus bicolor Peck 

Bolcnis eilulis Bulliard 



Boletus sp. 
Cortinarius sp. 

Galerina autumniiUs Peck 
Psathxrella delineata Peck 

Hygrophoraceae Hygrophorus iiuirgiiuitus Peck 

Pluteaceae Pluteus ceninus Schaeffer 

Polyporaceae Bominrzewia herkeleyi Fries 

Gi'ifola frontlosLi Fries 

Drosophihi falleni 
Tricimba lineella 
Drosophila falleni 

Phitxpeza sp. 

Didsopliila falleni 
Leucophenga varia 

Drosophila falleni 
Drosophila putriJa 

Leucophenga varia 

Platypeza sp. 

Drosophila falleni 
Leucophenga varia 
Drosophila falleni 
Leucophenga varia 

Drosophila falleni 
Drosophila putrida 
Drosophila falleni 
Drosophila putrida 
Leucophenga varia 
Drosophila falleni 
Leucophenga varia 

Drosophila falleni 
Drosophila putrida 
Leucophenga varia 

Drosophila falleni 
Leucophenga varia 

sp. #1 

Drosophila falleni 
Drosophila falleni 

Tricimba lineella 
Drosophila falleni 
Leucophenga varia 
Leucophenga varia 
Drosophila falleni 

Platypeza sp. 

Drosophila falleni 
Mvcodrosophila claytonae 

Drosophila falleni 























































Hihle I C'wntiniicd 

Diplera Spe, 


lAictiponis \itlfitretts Erie's 

Polyporus arculanus Bataille 
Tvromyces chioneus Eries 
Russula aeruginea Lindblad 
Russula hrevipes Peck 

Russula emetica Schaeffer 

Russula nuiriae Peck 

Russula pcbioleucoidcs Kauffman 

Russula sp. 

Strophariaceae Hypholoma fasciculare Hudson 

PhoHota malicola Kauttman 
Pholiora sp. 
Pholiora sp. 

Tricholomataceae Collybia acfnara Eries 

Mycena leaiana Berkeley 

Omphalorus alearius Schweinitz 
Pleurotus dryinus Persoon 
Pleurolus ostreatus Jacquin 

Tricholomopsis platyphylla Persoon 

Xerula fuifuriuea Peck 

Tricimba lineella 
Drosophila fallen! 
Drosophila falleni 
Limonia rara 
Drosophila falleni 
Drosophila putrieta 

Ptecticus sp. 
Liirumia triocellata 
Drosophila falleni 
Plecticus sp. 
Drosophila pulrida 
Leuinphenga varia 
Drosophila falleni 
Leucophenga varia 

Drosophila falleni 
Leucophenga varia 

Drosophila falleni 
Leucophenga varia 
Limonia triocellata 

Drosophila falleni 

Drosophila falleni 
Leucophenga sp. 
Drosophila falleni 
Leucophenga sp. 
Drosophila putrida 

Drosophila falleni 
Drosophila tripunctata 
Leucophenga varia 
Mycodrosophila claytonae 

Limonia triocellata 
Drosophila falleni 
Leucophenga varia 

sp. #] 
sp. #2 

Drosophila falleni 
Limonia Iriocellala 













































feia apparently i.s a rare mycophagous spe- 
cies known from only a few records (Pat- 
terson 1943). Bunyard and Foote (1990a) 
provided the only record for this species in 
Ohio. In a study of its life history, ovipo- 
sitional preferences, and larval feeding hab- 
its it was found to be polyphagous for fruits 

and other vegetation, but with a strong pref- 
erence for mushroom tissue (Bunyard and 
Foote i990b). 

Phoridae, a family of small flies, also is 
described as one of the most common in- 
habitants of fungal sporocarps (Hackman 
and Meinander 1979). Larvae are frequent- 


Table 2. Species of mycophagous Diptera. arranged phylogenetically by family, reared from mushroom 






LInioniii riirti Osten Sacken 
Limonia tiiocellala Osten Sacken 
Liinonia iriocellara Osten Sacken 
Limonia iriocellatci Osten Sacken 
Liinoiua Iriocellala Osten Sacken 

Pteclicus sp. 
PlecticHs sp. 


P/arypeZii sp. 

Tricimba lineella Fallen 
Tricimhci lineella Fallen 

,:phila fallcni Wheeler 






















































Tyromyces chioneus 
Russula brevipes 
Pholiota sp. 
Pteiirotiis oslreatus 
Xenila furfiiracea 
PhoHola sp. 
Pholiota sp. 
Russula brevipes 
Russula emelica 
Agaricus arvensis 
Agaricus augustus 
Agaricus bitorquis 
Agaricus campestris 
Amanita flavoruhescens 
Agrocybe praecox 
Boletus bicolor 
Cortinarius sp. 
Galerina autumnalis 
Pluteus cervuuis 
Pluteus cervinus 
Bondarzewia berkeleyi 
Bondarzewia berkeleyi 
Russula brevipes 
Russula ochroleuca 
Russula sp. 
Pleurotus dryiims 
Pleurotus ostreatus 
Tricholomopsis plalxphylUi 
Agaricus ar\-ensis 
Agaricus campestris 
Pluteus cen'inus 
Agaricus arvensis 
Psathyrella delineata 
iMetiporus sulfureus 
Agaric sp.? 
Agaricus arvensis 
Agaricus augustus 
Agaricus bitorquis 
Agaricus sp. 
Amanita flavoruhescens 
Amanita miiscaria 
Amanita rubescens 
Agrocybe praecox 
Boletus bicolor 
Boletus edulis 
Boletus sp. 
Cortinarius sp. 
Psathyrella delineata 
Pluteus cer\'inus 
Bondarzewia berkeleyi 
Grifola frondosa 
Laetiporus sulfureus 
Polyporus arcularius 
Russula aerugiiu'o 



T.ihk' 2. Continued. 

Fungus F.1 

Fungus Spci 






OrosophiUi /iiirridci Sliirtevant 

Drositp/iilii tiipiinchiui Loew 
LcHcophcnaa vana Walker 

LfKcophenga sp. 

Mycodrosophila claytonoe Wheeler 
and Takada 

Sp. #1 
Sp. #1 
Sp. #2 














































RiissnUi emetica 
RiisMila iichroU'iu-a 
Riissiila sp. 

Hypholoina fasviciilcire 
PholioUi sp. 1 
Collyhia acenata 
Mycena leaiana 
Pleuronts ostreurus 
Tricholomopsis platypliyUn 
Xeriila furfiiracea 
Agariciis bitorquis 
Amanita muscaria 
Anianiut nibescens 
BoU-liis hic»t«i 
RussnUi hre\'ipc.\ 
RkssuUi inariue 
Ompluiloliis oleariiis 
Pletimlus ostreaUis 
Ai>ai-icus augiislus 
Agariciis campestiis 
Agariciis sp. 
Amanita flavonthescens 
Amanita nibescens 
Agrocyin' praecox 
Boletus bicolor 
Boletus eilulis 
Psatliyrclla dcliucata 
Hygrophorus marginatus 
Russula mariae 
Russtila ochroleiica 
Russula sp. 
Pholiota imilicola 
Pleumlus ostrcatus 
Triclioiomtipsis pUityplivlla 
Collyhia acenata 
Mycena leaiana 
Bondarzewia berkeleyi 

PIcurotus ostrcatus 
Boletus sp. 
Xeriila fiirfuracea 
Keriila fiirfuracea 
Agariciis arvensis 
Agaricus campestris 
Boletus eilulis 
Pliiteiis ccn'inus 

ly found in decaying vegetation and fungi; 
.some species (especially of the genus Me- 
gaselia Rondani) are serious pests of com- 
mercial mushroom farms: a few species are 
known to be parasitic on other insects. Dur- 

ing this study, phorid flies emerged from 
more sporocaip collections than any other 
group except the Drosophilidae (Fig. 1). 

Two species of Tipulidae. Liinonia rarci 
Osten Sacken and L. iriocelhihi Osten 


May June July Aug 

Fig. I. Emergence numbers by month for families of Diptera. 

""* Anthomyiidae 

""* Chloropidae 

"• Drosophilidae 

'^ Phoridae 
^ ■ ■ Plalypezidae 

'^ Sarcophagidae 

~^ Tipulidae 

Sacken, were reared in this study (Table 2). 
Most larvae of Tipulidae feed in decaying 
plant materials and frequently are aquatic or 
semi-aquatic, although several species are 
terrestrial. Tipulids have been reared from 
fungi previously (Alexander 1920, Bruns 
1984). Liiiionia triocellata is a known con- 
sumer of decaying organic material (B. 

Foote, personal communication) and has 
been reared from senescent as well as fresh 
mushrooms (Bruns 1984). However, no in- 
formation is available for the feeding sub- 
strate of larval L. rara. Several adults of L. 
ram were reared from sporocarps of the 
soft moist bracket fungus Tyroinyces chi- 
oneus Fries. As the infested sporocarps 

— ♦— 


- D. putricla 


- D. tripunctata 


L.varia. L. sp. 

-M. claytonae 

Fig. 2. 

May June July Aug 

Emergence numbers by month for species of Drosophilidae. 


were newly emerged, this may indicate a 
strictly mycophagous (as opposed to scav- 
enging on decaying fungal or other organic 
material) feeding habit for this species. No 
species of Tipulidae cunently are consid- 
ered to be strictly mycophagous. 

Tricimba lineella Fallen (Chloropidae) 
was reared on a number of occasions (Table 
2). Tricimba lineella previously has been 
recorded from rotting plant material (Gri- 
maldi and Jaenike 1983) and from macro- 
fungi (Bunyard and Foote 1990a). One oth- 
er species (Gaurax atripalpus Sabrosky) 
has been recorded from fungi, probably 
overwintering in polyporaceous species 
(Valley et al. 1969, Bunyard and Foote 

The family Platypezidae was reared from 
fresh sporocarps mostly towards the end of 
the growing season (Fig. 1). The associa- 
tion was especially significant between this 
family and the basidiomycete family Agar- 
icaceae, as platypezids were reared almost 
exclusively from species of Agariciis (Table 
2). Species of Agariciis produce some of 
the largest sporocarps and would seem an 
ideal host, but they seem to be rarely uti- 
lized by other mycophagous Diptera (Bux- 
ton 1960, Hackman and Meinander 1979). 
Members of the Platypezidae (the "flat- 
footed" or "smoke flies") comprise a small 
family (71 species in 18 genera in North 
America) of uncommon flies of wooded ar- 
eas (Kessel 1987). Adult platypezids are 
noted by a fairly large head and character- 
istic flattened hind tarsi and tibiae. Al- 
though all platypezid species are thought to 
be mycophagous, the life cycles and larvae 
for many species remain unknown or have 
never been seen (Kessel 1987). 

Unidentified adults from the families An- 
thomyiidae and Sarcophagidae were reared 
from a number of large, mostly decaying. 
Basidiomycete sporocarps (Fig. 1) suggest- 
ing scavenging behavior and not strict my- 
cophagy. Both families are comprised of 
large bodied species that commonly are 
scavengers of decaying organic material. 

Curiously, several species of mycopha- 

gous Diptera were conspicuously absent 
from this study. For example (besides those 
already mentioned above), Leiomyza laevi- 
gata Meigen (Asteiidae) is a rare species 
known from only a handful of studies (Sa- 
brosky 1957. Papp 1972). Bunyard and 
Foote (1990a) confirmed its mycophagous 

Frequently, larvae of more than one fam- 
ily — often from several families — occupied 
the same sporocarp. Likewise, more than 
one species from a single family frequently 
emerged from the same sporocarp. Because 
fungal sporocarps are a limited and ephem- 
eral food source, it would seem logical that 
inter- and intraspecific competition pres- 
sures should exist. The Competitive Exclu- 
sion Principle (Hardin 1960) states that two 
species cannot coexist in a single limiting 
resource. So. how can we explain so many 
closely related species occupying the same 
niche? Previously, a few studies have pro- 
vided possible explanations as to how spe- 
cies of mycophagous Diptera avoid signif- 
icant interspecies competition. In particular, 
most studies have focused on the common- 
ly seen mycophagous species of Drosophil- 
idae. Biotic pressures (predation, parasit- 
ism) may reduce competition and thus al- 
low drosophilid species to cohabit (Worthen 
1989, Worthen et al. 1995, Jaenike 1998). 
Predation by ants and beetles, and parasit- 
ism by nematodes were the focus. During 
this study, numerous predacious beetles fre- 
quently were seen feeding (presumably on 
fly larvae) on and within mushroom tissues, 
especially the larger Basidiomycete species 
that were associated with rearings of mul- 
tiple species of drosophilids and other taxa. 
Adult parasitic Hymenoptera frequently 
emerged from the sporocarps. Presumably, 
parasitic wasps also could impinge upon the 
fly species to reduce their numbers, and 
thus lessen interspecific competition pres- 

Abiotic pressures also may reduce com- 
petition and allow different fly species to 
cohabit. Worthen and Haney (1999) found 
that when abiotic pressures (heat, desicca- 


tion) are strong, other dipterous species 
may dominate (e.g., D. pittrida is more 
common in small mushrooms that are more 
subject to desiccation and was found to be 
more tolerant of chronic and acute bouts of 
drought or high temperatures). Our findings 
strongly support this; D. putrida was seen 
as a dominant species only in the months 
of August and September (Fig. 2). 

Most of the attention of this paper — and 
the inteiTelationship between Diptera and 
fungal host — has been approached from the 
fly's point of view. It may be just as inter- 
esting to consider the host's (fungus) role 
in this symbiosis. Typical levels of my- 
cophagy frequently resulted in the complete 
destruction of the sporocarp. One would 
think this to be detrimental to the host. If 
this is true, it is logical to expect the host 
to fight back (the "Red Queen" effect |Van 
Valen 1973]). Few studies have attempted 
to determine if any macrofungi are unsuit- 
able to any groups of flies. Of course, by 
virtue of size alone, those fungi that form 
particularly small sporocarps will escape or 
have reduced mycophagy. Similarly, a few 
fungal species avoid damage by their phys- 
ical makeup: many species of bracket fungi 
(mostly within the family Polyporaceae) 
have a hard, woody texture that is difficult 
for many arthropods to consume (Courtney 
et al. 1990). Basidiomycete and Ascomy- 
cete fungi produce a wide array of toxic 
metabolites, although the defensive prop- 
erties of these have been poorly investigat- 
ed. Insecticidal properties have been ex- 
plored (Mier et al. 1996), although the au- 
thors carried out their study by feeding 
mushroom extracts to arthropods in a com- 
pletely artificial fashion. Clearly, the ability 
to detoxify secondary metabolites of mush- 
rooms is widespread throughout taxa of ar- 
thropods. This ability obviously has 
evolved more than once among families of 
Diptera. Furthermore, groups of basidio- 
mycete taxa (e.g., Amanita spp., Galerina 
Earle spp.) considered deadly to most ani- 
mals, including non-mycophagous Diptera. 

have no detrimental effect on mycophagous 
species (Jaenike et al. 1983, Jaenike 1985). 

Mycophagous Diptera may be of some 
benefit to their fungal host. Stinkhorns 
(Phallales: Basidiomycotina) benefit from 
scavenging flies that disperse their basid- 
iospores. Hodge et al. (1997) discussed the 
carriage of fungal material by adult Dro- 
sophila. Ascomycete fungi (including 
members of the genus Balansia) are known 
to benefit from symbiotic associations with 
insects. Recently, Diptera have been shown 
to disperse spermatia (a type of fungal 
spore) from one fungus to other individuals, 
thus facilitating fertilization (a sort of "pol- 
lination") (Bultman et al. 1998, 2000). 

Clearly, this study indicates a great need 
for additional investigations into the inter- 
relationships between fungal host and my- 
cophagous Diptera. Likewise, there are 
many uncertainties regarding the life his- 
tories for many species of mycophagous 
flies, as well as the interrelationships among 
the mycophagous Diptera. 


Sincere thanks are given to Dr. Benjamin 
Foote for identification of dipteran material 
and for critical review of this manuscript. 
Thanks also are extended to Dr. Tom Bruns 
and two anonymous reviewers for their 
helpful comments. This work was support- 
ed in part by a grant from the Geauga 
County (Ohio) Park District. 

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105(4). 2003. pp. 859-875 




A. R. Deans and M. Huben 

(ARD) Department of Entomology, 320 Morrill Hall. University of Illinois. 505 S. 
Goodwin Ave., Urbana, IL 61801, U.S.A. (e-mail: (MH) 27 Winter 
St., Arlington. MA 02474, U.S.A. (e-mail: 

Abstract. — The genus-level taxcMiomic history of Evaniidae is discussed and three new 
genera are described: Venievaiiia Huben and Deans from the Indian subcontinent with 
V. urbaniisorum Deans, n. sp.. Decevania Huben from Central and South America with 
D. pcina (Enderlein), n. comb., and D. striatigcna (Kieffer), n. comb., and Rothevcntia 
Huben from South America with R. valdiviaiui (Philippi). n. comb. The genus Acuiitlti- 
lu'vuiiia Bradley is resunected based on distinctive leg. mesosomal. and mouthpart mor- 
phology, raising the total number of valid genera to twenty. The following species are 
transfened from Evania: Acanlliinevania clavaticornis (Kieffer 1911). n. comb., ,4. Icii- 
cocnis (Kieffer 191 1), n. comb., A. sehcans (Westwood 1851), n. comb., and A. szepli- 
gctiiiiui (Hedicke 1939) n. comb. Sixteen additional genera are also recognized as valid: 
Afrevaiiia Benoit. Bnichevauia Turner, Bnichygaster Leach, Evania Fabricius, Evaniella 
Bradley, Evaniscus Szepligeti, Hyptia Illiger, Micrevania Benoit. Papatuka Deans, Par- 
cvania Kieffer, Prosevania Kieffer, Sennieomyia Bradley, Szepligetella Bradley, Tluiii- 
iiiatevania Ceballos. Trissevania Kieffer, Zeiixevania Kieffer A key to the genera of the 
world is provided. 

Key Words: ensign wasps, Evaniidae, Acaiitliiiievaiiia. Veriwvania. Roilicvaiiia, Dece- 

Despite the frequent collection and rec- and inadequate monograph as the sole 
ognizable habitus of ensign wasps, there source for species or genus identification, 
iiave been very few concentrated efforts di- Several characteristics of evaniid biology 
rected at improving our understanding of make them an interesting group of insects 
them. The most obvious impediment to fur- for future research. All verirtable rearings 
ther research is the current state of evaniid indicate that evaniid larvae develop as sol- 
classification. Since the early 1900"s (the itary egg predators within cockroach oothe- 
height of evaniid taxonomic activity) the cae (Roth and Willis 1960, Cameron 1957. 
amount of ensign wasp research has dwin- Crosskey 1951). This represents a possible 
died. Only one new extant genus has been method of non-chemical control for pestif- 
described since 1953 (Deans 2002). and the erous cockroaches (Thoms and Robin.son 
last comprehensive taxonomic treatment 1987, Edmunds 1953), but few studies have 
was Hedicke's (1939) world catalog. How- tested the effectiveness of these wasps, 
ever, this catalog does not include keys, and Evaniids also form multifaceted mimicry 
one must resort to Kieffer's (1912) outdated complexes, particularly in South America, 


with distantly related species possessing the 
same possibly aposematic and/or disruptive 
color schemes of black, brown, orange, and 
red. No published research, however, has 
explored this phenomenon. 

Several recent papers have contributed to 
our understanding of the evolutionary his- 
tory of the Evaniidae. Basibuyuk et al. 
(2000a, b, 2001) describe extinct genera 
and how they shape our understanding of 
modern evaniids. Dowton and Austin 
(2001 ), Dowton et al. ( 1997), and Ronquist 
et al. (1999), among many others, discuss 
the yet unresolved relationship of Evaniidae 
to the rest of the Hymenoptera. 

The purpose of this paper is to describe 
three new genera, two from South America 
(Decevania and Rotherania) and one from 
the Indian subcontinent (Venievania). and 
to resunect Acaiithiiievania Bradley. We 
also provide an illustrated key to the genera 
of the world. This is not meant to be a full 
revision but instead is intended as a starting 
point for researchers or budding evaniid 
taxonomists interested in determining spec- 
imens to genus as the genera are currently 
defined. Ongoing detailed morphological 
and molecular analyses by ARD into the 
generic limits, phylogenetic relationships of 
the genera, as well as how the Evaniidae 
relate to the rest of the Hymenoptera may 
improve our understanding of this enigmat- 
ic family. 

Taxonomic history 

The taxonomic instability within the 
Evaniidae has been well established, and 
most publications ( 1900-present) include 
one or two sentences bemoaning this fact. 
Prison (1922) divided the problems into 
several categories, which still apply today. 
First, nearly all species were described from 
only one or two specimens, allowing no 
discussion of variation. Color patterns can 
vary within a particular species, and it is 
possible that these color morphs have been 
described as separate species. Second, they 
are sexually dimoiphic (usually in antennal 
morphology, body coloration, facial sculp- 

ture, and/or metasomal morphology) and 
difficult to associate; some described spe- 
cies may actually represent the opposite sex 
of other species. Third, most holotypes are 
difficult to find. Kieffer described most of 
the evaniid species but rarely designated 
holotypes or mentioned their depository. 
Compounding this problem is the fact that 
nearly all the original descriptions are 
vague and could actually apply to multiple 
closely related species. Prison (1922) also 
noted that the number and limits of the eva- 
niid genera were disputed and difficult to 

Table 1 summarizes the history of genus- 
level classification for the Evaniidae based 
on the largest, most complete monographs. 
Schletterer (1889a, b) provided the first 
comprehensive treatment of world species, 
collapsing the three known genera into a 
single genus, Evania Fabricius. Bradley 
(1908) added three new genera and reas- 
signed species non visa. He based his tax- 
onomic scheme on several wing venation 
and non-wing characters (e.g., metasoma 
shape, antenna, leg, tarsal claw, and mouth- 
part morphology, shape of the "furculum"" 
between hind coxae). Kieffer (1912). in an 
unfortunate step backwards, reclassified the 
evaniids solely on wing venation charac- 
ters. He synonymized Acanthiiievcinia, 
Evciniella, and Szepligetella under Evania, 
and Semaeomyia under Brachygaster. Hed- 
icke's (1939) world catalog largely fol- 
lowed Kieffer's (1912) classification but in- 
cluded all the genera and species described 
between 1912-1939. 

Several synapomorphies define the Eva- 
niidae. The most recognizable are the high 
insertion of the metasoma (Figs. 1-2) and 
the thin tubular shape of the petiole (Pigs. 
3-5) giving these wasps their characteristic 
habitus. Most species also have 1 1 flagel- 
lomeres (except Decevania spp. have 8; 
Pig. 1 ) and hind wings with greatly sepa- 
rated jugal lobes (Pigs. 6-8) (except some 
Evaniella and Prosevania spp. <2.5 mm 
long). All evaniid rearings have been from 


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cockroach egg cases, and it is assumed that 
this represents a synapomorphic Hfestyle. 

Key TO World Genera of Evaniidae 

The Evaniidae would benefit enormously 
from a worldwide family-level revision. 
The limits between certain genera are not 
well defined, and we are aware of a few 
specimens that still do not key to the correct 
genus. These rare exceptions are discussed 
under each genus. 

Wing venation varies greatly within the 
family and provides the most useful char- 
acteristics for distinguishing genera. Unfor- 
tunately, ensign wasps are prone to wing 
venation aberrations (usually missing or ex- 
tra veins; see Mani and Muzaffer 1943) 
which may lead one to inappropriate cou- 
plets. Also helpful are head shape/sculptur- 
ing, tarsal claw morphology, metasoma 
shape, and female genitalia morphology. 
Qualifying terms are used to estimate the 
number of specimens displaying a particu- 
lar character based on material examined 
(—7,000 specimens): sometimes (<50%), 
often (50-75%), or usually (75-99%