PROC. ENTOMOL. SOC. WASH. 
95(3), 1993, pp. 327-350 
REVIEW OF THE GENUS CHELIPODA MACQUART OF AMERICA NORTH OF 
MEXICO (DIPTERA: EMPIDIDAE; HEMERODROMIINAE) 
John F. MacDonald 
Department of Entomology, 1 158 Entomology Hall, Purdue University, W. Lafayette, 
Indiana 47907-1158. 
Abstract.— The genus Chelipoda Macquart is reviewed for America north of Mexico. 
Descriptions of C contractu Melander, C. elongata (Melander) (Lectotype designated), C. 
praestans Melander, C. sicaria Melander, C americana (Melander) (new combination), 
Chelipoda limitaria n. sp., and Chelipoda truncata n. sp. are presented. A key to males 
and females, illustrations of male terminalia, known distributions, and comments on 
biology and systematics are included. 
Key Words: Diptera, Empididae, Hemerodromiinae, Chelipoda, Phyllodromia 
Adult flies of the genus Chelipoda Mac- 
quart are very small (body length about 2 
mm) and delicate. They possess strong rap- 
torial fore legs, and their legs and body usu- 
ally are predominantly yellow to yellowish 
brown. The wings are relatively slender since 
they lack an anal lobe. Chelipoda has been 
placed in the subfamily Hemerodromiinae, 
the taxonomy of which was treated by Me- 
lander (1902, 1928, 1947). This paper is 
limited to species of Chelipoda of America 
north of Mexico and includes revised de- 
scriptions of known species, descriptions of 
two new species, a key to male and female 
adults, known distributions, and comments 
on biology and systematics. In addition to 
species treated here, representatives fitting 
the present concept of Chelipoda are re- 
ported from: China (Yang and Yang 1990); 
India, Indonesia, Philippines and Taiwan 
(Melander 1928); Europe (Chvala and Wag- 
ner 1989, Collin 1961, Engel 1956); New 
Zealand (Collin 1928); and South America 
(Collin 1933, Lynch Arribalzaga 1878, 
Smith 1967). 
Materials and Methods 
The present study was facilitated by ex- 
amination of the large numbers of Cheli- 
poda adults added to North American col- 
lections since Melander's (1947) revision. 
The following institutions (acronyms fol- 
lowing) loaned the material upon which this 
work is based: American Museum of Nat- 
ural History, New York (AMNH); Cana- 
dian National Collection, Biological Re- 
sources Division, Agriculture Canada, 
Ottawa (CNC); California Academy of Sci- 
ences, San Francisco (CAS); Cornell Uni- 
versity, Ithaca (CU); Florida State Collec- 
tion of Arthropods, Gainesville (FSCA); 
Helzinki Zoological Museum (HZM); Pur- 
due University Entomological Research 
Collection (PERC); Snow Museum, Uni- 
versity of Kansas, Lawrence (UKL); United 
States National Museum of Natural His- 
tory, Washington, D.C. (USNM); Univer- 
sity of Minnesota, St. Paul (UMSP); Uni- 
versity of New Hampshire, Durham (UNH); 
University of Wisconsin, Madison (UWM); 
and Utah State University, Logan (USU). 
328 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Specimens also came from the author's col- 
lection (MAC). 
Specimen examination took place under 
fiber optic illumination, with study of ves- 
titure facilitated by use of fluorescent light- 
ing. Since coloration and color patterns are 
in part a consequence of light incidence, ex- 
amination of antennae and tarsi under a 
microscope required rotation of specimens. 
Body length was measured from front of 
head (exclusive of antennae) to the apex of 
female cerci and the most distant margin of 
male terminalia (not their apex since they 
project anteriorly over the abdomen). Mac- 
eration of body parts during the early phase 
of the study was conducted in an approxi- 
mately 15% solution of sodium hydroxide 
heated to about 80 C, but much better prep- 
arations were achieved after maceration in 
85% lactic acid heated to about 80 C, as 
described by Gumming (1992). Some spec- 
imens prepared by the latter technique sub- 
sequently were passed through an approx- 
imately 1 5% solution of sodium hydroxide 
heated to about 80 C in order to completely 
remove soft tissue and to effect additional 
clearing of heavily sclerotized male termi- 
nalia. 
Special of Chelipoda possess a number of 
characters of taxonomic value. Among the 
most important is vestiture, the terminol- 
ogy for which is based on McAlpine (1981). 
The basic form of macrotrichia is a seta (= 
hollow, articulated epidermal outgrowth), 
with the following descriptive terms applied 
to it: hair (= long, slender seta); bristle (= 
long, stout seta); and setula (= short, stout 
seta). Setulae on adults of Chelipoda typi- 
cally are black, and may be sharply pointed 
or bluntly pointed; the latter assume a peg- 
like appearance. The color of antennal seg- 
ments and arista, distal tarsomeres and 
postgena is also of taxonomic importance. 
Relative elongation of the thorax, based on 
the length of the notopleural suture relative 
to the length of the scutum, is important 
and among Nearctic species sorts into two 
forms: 1) thorax relatively long and slender. 
in which the notopleural suture is ca. % the 
length of scutum; and 2) thorax relatively 
short and compact, in which the notopleural 
suture is ca. V2 the length of scutum. Length 
of the female cercus relative to its basal width 
helps in diagnosing some species. 
Male terminalia are distinctive for each 
species. Interpretation and terminology in 
general follows McAlpine (1981) and, spe- 
cifically, the recent interpretation of geni- 
talic homologies outlined in Gumming and 
Sinclair (1990). Maceration is required to 
reveal internal features, but at leasi some of 
the diagnostic features are visible on many 
intact dry-mounted males, especially those 
collected into alcohol and then either pre- 
pared by critical point drying, or by passing 
through cellosolve and then xylene (Sa- 
brosky 1966). Both lateral and dorsal views 
of male terminalia are illustrated and each 
includes details of internal structures. 
Genus Chelipoda Macquart 
Chelipoda Macquart 1823: 148. Type spe- 
cies: Tachydromia mantispa, misidenti- 
fication (orig. des.) (= vocatoria Fallen, 
1816). 
Phyllodwmia Zetterstedt 1837: 31. Type 
species: Empis melanocephala Fab. (des. 
Rondani, 1856: 150). 
Chiromantis Rondani 1856: 148 (also as 
Chyromantis, p. 148) (preocc. Peters, 
1854). Type species: Tachydromia voca- 
toria Fallen 1816 (orig. des.). 
Lepidomya Bigot, 1857: 557, 563. Type 
species: Tachydromia mantispa Panzer, 
1806 (= Empis melanocephala Fab., 
1794). Junior synonym of Phyllodromia 
Zetterstedt, 1837. 
Thamnodromia Mik 1886: 278. (unneces- 
sary name change for Phyllodromia Zett.). 
Litanomyia Melander 1902: 231. Type spe- 
cies: Sciodromyia mexicana Wheeler & 
Melander (Goquillett, 1903: 252). 
LepidomyiaKertesz, 1909: 1 17. (unjustified 
emendation of Lepidomya Bigot, 1857). 
VOLUME 95, NUMBER 3 
329 
Diagnosis. —Adult flies in the genus Che- 
lipoda are very small (body length including 
terminalia usually 1.8 to 2.2 mm, with fe- 
males slightly larger than conspecific males), 
delicate, usually yellow to yellowish brown 
flies that possess strong raptorial fore legs. 
Differing from those of other Nearctic Hem- 
erodromiinae, adults of Chelipoda possess 
an antennal arista at least twice as long as 
the flagellum, an unbranched R4 , 5 vein, and 
bristles on the laterotergite. Characteristic, 
but not necessarily diagnostic features of 
Chelipoda, are the male terminalia that pro- 
ject anteriorly over the abdomen and the 
lack of a sclerotized ovipositor in females. 
Description. — Nearctic species o^ Cheli- 
poda agree with the detailed generic descrip- 
tion presented by Collin (1961). One minor 
modification to Collin's description per- 
tains to the fore tibia being ridged beneath. 
Examination of slide-mounted fore legs re- 
vealed that the "ridge" actually is formed 
by a contiguous series of black setulae, each 
bent at a right angle toward the apex of the 
fore tibia. 
Remarks.— Two arrangements of wing 
venation occur among species allied with 
Chelipoda and have provided a prior basis 
for distinguishing two species groups, given 
either generic or subgeneric rank. Species 
possessing crossvein dm-cu, and thus a 
closed cell dm, have been placed in Cheli- 
poda Macquart whereas species lacking 
crossvein dm-cu, which results in an open 
cell dm, have been placed in Phyllodwmia 
Zetterstedt. This latter taxon has been treat- 
ed as a genus by Steyskal and Knutson 
(1981), Collin (1 96 1 ) and Melander ( 1 947). 
Although treating Phyllodwmia as a genus, 
both Collin (1961) and Melander (1947) 
commented on the doubtful generic impor- 
tance of crossvein dm-cu. For the same rea- 
son, Tuomikoski (1966) considered Phyl- 
lodromia as a subgenus of Chelipoda. 
However, since no phylogenetic analysis has 
demonstrated that ''Phyllodromia'" and 
"'Chelipoda'''' arc monophyletic groups, for- 
mal designation of Phyllodwmia as a sub- 
genus is not considered warranted, and all 
species treated here are placed in Chelipoda, 
without subgeneric designation. Two Pale- 
arctic species, C albiseta Zetterstedt and C. 
vocatoha Fallen, once included on species 
lists from the eastern United States, appar- 
ently do not occur in North America. 
The identification key relies upon vesti- 
ture and male terminalia. Important vesti- 
ture applicable to both males and females 
includes: 1) the presence of either one or 
two ventral rows of some 16-20 black setu- 
lae on the fore femur; and, 2) the presence 
or absence of a prominent basolateral seta 
(sometimes two) on the fore coxa. Taxo- 
nomic components of male terminalia in- 
clude: degree of fusion between the hypan- 
drium and epandrium, ranging from 
separate to completely fused; degree of fu- 
sion between each cercus and corresponding 
epandrial lobe, ranging from separate to 
completely fused; structure and vestiture of 
cerci; length and shape of the phallus; length 
and shape of a pair of epandrial lobes; and, 
if developed, length and shape of a pair of 
phallic processes. 
Key to Adults of Species of 
Chelipoda Macquart of America 
North of Mexico 
1 . Crossvein dm-cu lacking, cell dm-cu open (Fig. 
1) 2 
- Crossvein dm-cu present, cell dm-cu closed (Fig. 
2) 3 
2. Phallic process extending ca. 'A length of cer- 
cus; subepandrial lobe in-turned apically with 
tip crossing mid-line and ending in 3 black 
processes (Fig. 3) C americana (Melander)'' 
- Phallic process extending slightly beyond apex 
of cercus; subepandrial lobe in-turned apically, 
but tip lacking 3 black processes apically and 
not crossing mid-line (Fig. 4) 
C. limitaria. new species" 
3. Fore femur ventrally with 2 complete rows of 
black setulae (outer row usually containing 
fewer), flanked laterally by row of ca. 4-5 light 
brown bristles (Fig. 5) 4 
- Fore femur ventrally with only 1 complete row 
' Reliable characters for identifying females were not 
discovered (see diagnosis section under both species). 
330 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
(inner) of black setulae (outer row of fewer, less 
prominent setulae may exist), flanked laterally 
by ca. 4-6 light brown bristles (Fig. 6) 6 
4. Fore coxa with 1 or 2 prominent setae baso- 
laterally (occasionally dislodged) and row of 
finer setae continuing distally (Fig. 7); male 
terminalia subequal to pre-genital segment (Fig. 
9); female cercus length and basal width sub- 
equal (Fig. 10) C. contracta Melander 
- Fore coxa lacking prominent seta basolaterally 
(Fig. 6); male terminalia distinctly longer than 
pre-genital segment; female cercus ca. 2 x lon- 
ger than basal width (Fig. 11) 5 
5. Distal tarsomere brown, remaining tarsomeres 
yellow; postgena, gena, and occiput black; fla- 
gellum dark brown; subepandrial lobe pointed, 
not strongly sclerotized apically; phallic pro- 
cess and phallus subequal in length (Fig. 12) 
C. elongata Melander 
- Tarsomeres yellow; postgena with distinct pale 
area on mid-ventral aspect that contrasts with 
darker gena and occiput (Fig. 8); flagellum yel- 
low (male) or light brown (female); subepan- 
drial lobe truncate, strongly sclerotized apical- 
ly; phallic process 'A length of phallus (Fig. 13) 
C tnincata, new species 
6. Phallus slender in dorsal view, in lateral view 
distal 'A sinuate; subepandrial lobe not appar- 
ent in lateral view, extending only ca. Vw length 
of phallus (Fig. 14) C praestans Melander*' 
- Phallus not uniformly slender in dorsal view, 
in lateral view distal 'A abruptly bent; subepan- 
drial lobe well developed, ca. % length of phal- 
lus and bent upward in lateral view (Fig. 1 5) 
C. sicaria Melander'' 
Chelipoda americana (Melander), 
New Combination 
(Figs. 1, 3) 
Phyllodromia americana Melander, 1947: 
269. 
Diagnosis.— Absence of crossvein dm-cu 
in both wings distinguishes adults of this 
species and those of C. limitaria, newly de- 
scribed below, from other species of Che- 
lipoda treated here on which dm-cu very 
'' Females are difficult to identify, with side by side 
comparison necessary: those of C. sicaria possess a 
brown antennal arista that contrasts with the pale ped- 
icel and brown thoracic setae; those of C. praestans 
possess a tan antennal arista that does not contrast with 
the pale pedicel and tan thoracic setae. 
rarely is lacking in one wing. Examination 
of terminalia is required to separate males 
of C americana and those of C. limitaria. 
The distinction between them is based on 
comparative structure of the subepandrial 
lobes and phallic processes. Males of C. 
americana possess phallic processes that are 
much shorter than the cerci and acutely in- 
turned subepandrial lobes, the tips of which 
cross and each of which ends in three black 
projections. In comparison, males of C lim- 
itaria possess phallic processes that are 
slightly longer than the cerci and lack black, 
distal projections on the subepandrial lobes, 
the tips of which do not cross. 
Description. — Length including termina- 
lia of male ca. 1.8 to 2.0 mm, of female ca. 
1.8 to 2.1 mm. General color of male yel- 
low, of female yellowish brown. Head black, 
except for yellow mouthparts and palps; 
bristles yellow. Antennal scape and pedicel 
yellow; flagellum and arista of male yellow- 
ish brown, of female brown. Thorax short, 
compact; bristles yellow. Legs yellow; distal 
tarsomere light brown. Fore femur ventrally 
with 2 rows of black setulae, each row 
flanked by row of 4-5 yellow bristles (see 
Fig. 5). Fore coxa lacking prominent ba- 
solateral seta, but row of fine setae continu- 
ing distally (see Fig. 6). Wing hyaline; cross- 
vein dm-cu lacking; cell dm open (Fig. 1). 
Abdominal terga of male greyish brown, of 
female reddish brown; sterna paler than ter- 
ga in both sexes. Male terminalia (Fig. 3) 
yellow, compact; hypandrium and epan- 
drium only partially fused medially; cercus 
and epandrial lobe completely fused; cercus 
slender distally, ending in 3 strong setulae; 
subepandrial lobe curved acutely inward 
with tip crossing mid-line in dorsal view, 
ending in 3 pointed, black processes of un- 
even size; phallic process pointed apically, 
ca. 2 X length of phallus and ca. Vi length of 
cercus. Female cercus short (see Fig. 10); 
spermatheca more or less reniform, with 
spermathecal duct arising from center of 
concave surface. 
Type material examined. — HOLOTYPE 
VOLUME 95, NUMBER 3 
331 
dm-cu 
Figs. 1,2. 1, Chelipoda americana wing. 2, Chelipoda contracta wing, dm = cell; dm-cu = crossvein. Scale 
bar = 1.0 mm. 
male, labelled "Mt. Monadnak/ 26 Jul '26 
NH/ A L Melander" (USNM). The speci- 
men is in excellent condition, at least some 
of the diagnostic features of the terminalia 
are visible without maceration, and the cor- 
rected type locality is Mt. Monadnock, New 
Hampshire. ALLOTYPE. Virginia: Great 
Falls, June (lacking head) (USNM). PARA- 
TYPES. Georgia: 1 female. Burton, May; 1 
female, Decatur Co., Jun (USNM). New 
York: 2 females, 1 male, Millwood, Jun 
(USNM). Rhode Island: 1 female. Westerly, 
Jul (USNM). Virginia: 1 specimen (lacking 
abdomen). Great Falls, Jun (USNM). 
Other specimens examined. — CANADA. 
Ontario: 1 female, 3 km E. Carp, Jul (CNC). 
UNITED STATES. Georgia: 1 female, 
Waycross, Mar (CNC). Florida: 1 female, 
Alachua Co., Apr (USNM). Maryland: 2 
males, Bethesda, Jun (USNM). Massachu- 
setts: 1 male, Nonomesset, Jul (USNM). 
New Hampshire: 1 female, Mt. Madison, 
Dolly Copp cmpgr., Jul (CNC). New York: 
1 male, St. Lawrence Co., Jun (USNM). 
North Carolina: 1 male. Highlands, May 
(CNC). 
Distribution. — Adult males of this spe- 
cies are known from eastern New York state, 
New Hampshire, southeastern Maryland, 
and the mountains of western North Car- 
olina (Fig. 16). 
Remarks. — Some of the female paratypes 
and other females of C americana recorded 
from the southeastern United States (Fig. 
1 6) probably are females of C. Umitaria. As 
mentioned in the identification key, reliable 
characters for separating females of C 
americana and C Umitaria were not found 
(therefore the qualification pertaining to 
distribution records). However, female 
specimens from the northeastern United 
States and southeastern Canada, which 
probably are C. americana, are slightly 
smaller and possess a paler antennal arista 
than females in the type series of C. Umi- 
taria. 
Chelipoda contracta Melander 
(Figs. 2, 5, 7, 9, and 10) 
Chelipoda contracta Melander, 1947: 265. 
Diagnosis. —Adults are distinguished 
from those of other species treated here by 
the combination of two ventral rows of black 
setulae on the fore femur and at least one 
prominent seta (sometimes two) basolater- 
ally on the fore coxa. The basolateral seta 
on the fore coxa usually is prominent, but 
often is less obvious on males, and occa- 
sionally is dislodged from one or both legs. 
Males are recognized by their delicate yel- 
low bodies and small, compact terminalia. 
The relatively short, compact thorax of both 
sexes and the short cercus of females aid in 
separating adults of C contracta from those 
of C elongata and C truncata, which also 
possess two complete rows of black setulae 
ventrally on the fore femur. 
Description. — Length including termina- 
lia of male ca. 1.6-1.8 mm, of female ca. 
1.8 to 2.1 mm. General color of males yel- 
low, of females usually yellowish brown. 
332 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
'^^^P l^oelof 
idema wai 
Figs. 3, 4. 3a (lateral) and b (dorsal), Chelipoda americana male terminalia. 4a (lateral) and b (dorsal), 
Chelipoda limitaria male terminalia. ce = cercus; hyp = hypandrium; ph = phallus; php = phallic process; ep 
= epandrium; subep = subepandrial lobe. Scale bar = 0.25 mm. 
Head black, except for grey frons and post- brown; bristles yellow. Legs yellow; fore fe- 
gena; mouthparts and palps yellow; bristles mur ventrally with 2 rows of black setulae, 
yellow. Antennal scape and pedicel yellow; each flanked by row of 4-5 light brown 
flagellum and arista light brown to brown, bristles (Fig. 5); fore coxa with a prominent 
Thorax short, compact; yellow to yellowish basolateral seta (Fig. 7), weaker and less 
VOLUME 95, NUMBER 3 
333 
^^^M 
•33'? 
5 ^ 
irlema i9si 
Figs. 5-8. 5, Chelipoda contracia fore femur. 6, Chelipoda praestans fore femur. 7, Chelipoda contractu fore 
coxa. 8, Chelipoda truncata mid- ventral view of postgena. fcs = basolateral seta of fore coxa; ffst = setulae of 
fore femur; pg = postgena. Scale bars = 0.5 mm. 
prominent on male. Wing hyaline; cross- 
vein dm-cu present; cell dm closed (Fig. 2). 
Abdominal terga yellowish brown, sterna 
light brown. Male terminalia (Fig. 9) yellow; 
compact, subequal to pre-genital segment; 
cercus and phallus projecting anteriorly only 
slightly beyond the completely fused hy- 
pandrium and epandrium; cercus separate 
from epandrial lobe, nearly rectangular in 
lateral view; subepandrial lobe sword- 
shaped, ca. % length of phallus; phallus 
strongly expanded at apex in dorsal view, 
subequal in length to cercus; phallic pro- 
cesses lacking. Female cercus short; sper- 
matheca hemispherical, with spermathecal 
duct arising from center of flat surface (Fig. 
10). 
Type material examined. — HOLOTYPE 
334 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
subep , ( ph 
Figs. 9-n. 9a (lateral) and b (dorsal), Chelipoda contractu male terminalia. 10, Chelipoda contractu female 
terminalia. 11, Chelipodu elongata female terminalia. ce = cercus; ph = phallus; sp = spermatheca; subep = 
subepandrial lobe. Scale bars = 0.25 mm (Fig. 9a, b) and 0.1 mm (Figs. 10, 1 1). 
VOLUME 95, NUMBER 3 
335 
male, labelled "Petersham/ MASS 9 vi '32/ 
A L Melander" (USNM). The specimen is 
in excellent condition and at least some of 
the diagnostic features of the terminalia are 
visible without maceration. A female la- 
belled "ALLOTYPE C contracta'' with the 
same collecting data as the holotype was C. 
praestans. PARATYPES. CANADA. Brit- 
ish Columbia: 1 female, Abbotsford, Aug 
(USNM); 3 females (CU) and 1 female 
(USNM), Downie Crk., Selkirk Mts., Aug. 
Ontario: 1 male, 4 females, Waubamich, 
Jun (USNM). UNITED STATES. Con- 
necticut: 3 females, 5 males. Redding, May- 
Jun (USNM). Massachusetts: 1 female, 1 
male, Boston, Jun; 1 female, 1 male, Peter- 
sham, Jul (USNM). Maine: 2 females, Seal 
Harbor, Jul (USNM). New Hampshire: 4 
females, Breton Woods, Jul (USNM). New 
York: 4 females, 1 male, Ithaca, Jun; 1 fe- 
male, 2 males. Tuxedo. May (USNM). 
Washington: 1 1 females, 1 2 males. Index, 
Aug; 2 females, Lake Cushman, Jul; 1 1 fe- 
males, Mt. Baker, Skyline Trail, Aug; 4 
males, Mt. Constitution, Jul; 1 male, Mt. 
Vernon, Jul; 1 female. Sultan, Aug (USNM). 
Other specimens examined. —CANA- 
DA. Alberta: 1 female, Waterton Prk., Jul 
(CNC). British Columbia: 3 females, 1 male, 
Cultus Lk., Jun-Aug; 1 female Liard Hot 
Sprs., Jul; 36 females, 35 males. Terrace, 
Jun-Aug (CNC, CU, UKL, USNM). Man- 
itoba: 1 female, Brandon, Jul; 1 female, 1 
male, Forrest, Jul; 5 females, Ninette, Jul; 
8 females. Turtle Mt., Jul-Aug (CNC). New 
Brunswick: 2 females, Acadia, Jun; 1 fe- 
male, Chamcock, Jul (CNC). Newfound- 
land: 1 female. Bay of Islands, Jul; 7 males, 
8 females, St. Johns, Aug (AMNH, CNC). 
Nova Scotia: 29 females, 46 males. Cape 
Breton Highlands Nat. Prk., Jun-Aug; 14 
females, 2 males, Lockport, Jul-Aug; 1 fe- 
male, Shelbume, Aug; 5 females, Spring- 
field, Aug (CNC). Ontario: 1 male, Atiko- 
kan, Jul; 2 females. Ft. Francis, Jul; 1 male, 
3 females, Griffith, Jun-Jul; 1 female, Ke- 
nora, Aug; 1 female, Mamora, Jul; 2 fe- 
males, Maynooth, Jun; 2 females, 1 male. 
Midland, Jul-Aug; 1 male, One-Sided-Lake, 
Jun; 4 females, 4 males, Ottawa, Jun-Jul; 1 
female, S. March, Jun; 2 females, Waubam- 
ich, Jun-Jul (CAS, CNC). Quebec: 1 female, 
Abbotsford, Jun; 1 female, 1 male. Beech- 
grove, Jun; 3 females, 2 males, Brecken- 
ridge, Jun; 1 female Cap Rouge, Jul; 1 fe- 
male, Corey Hill, Jun; 1 7 females, 1 3 males. 
Old Chelsea; 2 females. Park Reserve, Jul; 
1 female, Rivere-du-Loop, Jul; 26 females, 
1 male, Wakefield, Jun-Jul (CNC, USNM). 
Saskatchewan: 1 female, 1 male, Rockglen, 
Jun; 2 females, 3 males. Scout Lk., Jun 
(CNC). UNITED STATES. Georgia: 6 fe- 
males, 5 males, Athens, Apr-May; 1 female. 
Black Rock Mt., May; 2 females, Rabun 
Co., Aug (CNC, CU, MAC). Illinois: 1 fe- 
male, Macomb, May (USNM). Maine: 2 fe- 
males, Mt. Katahdin, Jul (CNC). Maryland: 
1 female, 1 male, Bethesda, May-Jun 
(USNM). Massachusetts: 1 female, 1 male. 
Woods Hole, Aug (AMNH). Michigan: 1 
male, Cadillac, Jun; 1 female, Crawford Co., 
Jun; 2 females. Isle Royale, Jul-Aug; 2 fe- 
males, Manistee, Jul; 3 females. Midland, 
Jun-Jul; 1 male, Schoolcraft, Jun (CNC, 
USNM). Minnesota: 8 females, 2 males, 
Basswood Lk., Jul-Aug; 1 female. Eagle- 
nest, Aug; 2 females, 2 males, Itaska St. Prk., 
Jun-Jul (AMNH, UMSP). Montana: 2 fe- 
males, Fathead Lk., Aug (USNM). New 
Jersey: 1 female, Brookside, Sep (AMNH). 
New York: 1 female, Essex Co., Jul; 1 fe- 
male, Orleans Co., Aug (CAS, CNC). North 
Carolina: 1 female, Coweeta, May; 2 fe- 
males, 1 male. Highlands, May; 1 female. 
Looking Glass Pk., Jul; 1 female, Macon 
Co., Jul (CNC, UNH); 1 female, McDowell 
Co., Sep (USNM). Pennsylvania: 3 females. 
Spring Bridge, Jun (USNM). Tennessee: 4 
males. Great Smoky Mts. Nat. Prk., E. Gat- 
linburg, Jun-Jul (USNM). Virginia: 1 fe- 
male, Alexandria, Jun; 1 female. Big Mead- 
ow, Jun; 2 females, Blacksburg, May; 2 
males. Brush Mt., May; 2 males. Falls 
Church, May; 5 females, 1 male, Hawksbill, 
336 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Jun (CNC, USNM). Washington: 4 females, 
1 male, Friday Harbor, Jun-Jul (AMNH, 
CU, FSCA). Wisconsin: 1 female, Vilas Co., 
Jun (USNM). 
Distribution.— This is the most wide- 
spread Nearctic species, occurring across 
southern Canada and the northern United 
States, and south into the Appalachian 
mountains of northern Georgia (Fig. 1 7). 
Remarks. —Adults have been collected at 
sea level (Puget Sound, Washington) and 
near 1200 meters in the Appalachian 
Mountains. Specific habitats in the Appa- 
lachian Mountains include mesophytic 
hardwood, hemlock, and damp fir/spruce 
forests. Labelled specimens from Manitoba 
are from a Maple/Elm floodplain, "around 
a Populus balsamifera L. (balsam popular) 
slough," "in secondary growth vegetation 
associated with power lines," and a dead 
cat. Specimens also have been taken off 
blossoms of Vaccinium sp. in Virginia and 
in "wet, scrubby clearings" near Terrace, 
British Columbia. Several series consisting 
of males and females were collected in Mal- 
aise traps and yellow pan traps in Cape Bre- 
ton Highlands National Park, Nova Scotia, 
by J. R. Vockeroth (Biological Resources 
Division, Agriculture Canada, Ottawa). 
Chelipoda elongata (Melander) 
(Figs. 11, 12) 
Litanomyia elongata Melander, 1902: 232. 
Chelipoda elongata; Melander, 1928: 266. 
Chelipoda albiseta (Zetterstedt) sensu Me- 
lander, 1947: 265 (misident.) 
Diagnosis.— Adults of this species, sim- 
ilar to those of C. contracta, possess two 
ventral rows of black setulae on the fore 
femur, but they are distinguished by the 
elongated thorax, lack of a prominent ba- 
solateral seta on the fore coxa, and a long 
female cercus. The contrastingly darker dis- 
tal tarsomere and concolorous postgena, 
gena and occiput distinguish adults of C 
elongata from those of C. tmncata, newly 
described below. Male terminalia resemble 
those of C tmncata, but differ in possessing 
pointed subepandrial lobes and phallic pro- 
cesses that are subequal in length to the 
phallus; males of C tmncata have truncate 
subepandrial lobes and much shorter phal- 
lic processes, which are about half the length 
of the phallus. 
Description. — Length including termina- 
lia of male ca. 2.0-2.5 mm, of female ca. 
2.5-2.8 mm. General color yellow to yel- 
lowish or light reddish brown. Head black, 
including postgena; mouthparts and palps 
yellow to light reddish brown; bristles brown. 
Antennal scape and pedicel yellow; flagel- 
lum brownish black; arista brownish black 
on basal tenth, distally brownish to nearly 
white in some specimens. Thorax long, slen- 
der; yellow to yellowish brown; bristles 
brown. Legs yellow, except coxa and distal 
tarsomere light brown. Fore femur ventrally 
with 2 rows of black setulae, each flanked 
by row of 4-5 light brown bristles (see Fig. 
5). Fore coxa lacking prominent basolateral 
seta, but row of fine setae continuing distally 
(see Fig. 6). Wing hyaline; crossvein dm-cu 
present; cell dm closed (see Fig. 2). Abdom- 
inal terga brown, sterna light brown. Male 
terminalia (Fig. 12) yellow, projecting an- 
teriorly to abdominal segment 6; hypan- 
drium and epandrium completely fused; 
cercus fused to epandrial lobe, nearly 
straight, subequal in length to subepandrial 
lobe; subepandrial lobe slender, slightly 
curved ventrally and pointed apically; phal- 
lus and phallic process nearly straight, sub- 
equal in length, extending anteriorly beyond 
cercus and subepandrial lobe. Female cer- 
cus long; spermatheca nearly oval, with 
spermathecal duct arising from the center 
of a somewhat flattened pole (Fig. 1 1). 
Type material examined. —LECTO- 
TYPE (here designated from Melander's co- 
type series number 29182), male labelled 
"Brookings/ SD" (USNM). The lectotype 
lacks the head and most of the right front 
leg, but most of the diagnostic features of 
terminalia are visible without maceration. 
The paralectotype series includes a C. elon- 
VOLUME 95, NUMBER 3 
337 
php 
subep 
K'oelof 
idema i9Si 
Figs. 12, 13. 12a (lateral) and b (dorsal), Chclipoda elongata male terminalia. 13a (lateral) and b (dorsal), 
Chelipoda iruncata male terminalia. ce = cercus; ph = phallus; subep = subepandrial lobe. Scale bar = 0.25 mm. 
gata female with a "Mass" label, a C. prae- Other specimens examined. — CANA- 
stans female with a "Mass" label, and a DA. Manitoba: 7 females, 4 males, Ninette, 
badly damaged Chelipoda specimen with a Jul; 3 males, Turtle Mt., Jul (CNC). Ontar- 
"Mass" label. io: 1 male. Ft. Credit, Aug; 1 male, Ft. 
338 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Frances, Jul; 21 females, 6 males. Grand 
Bend, Jul; 2 females, Griffith, Jul; 1 female, 
Kearny, Jul; 3 females, 6 males, Ottawa, Jul 
(CNC, USNM). Quebec: 2 males, Wake- 
field, Jul; 4 females, 1 male, Hull, Aug 
(CNC). UNITED STATES. Florida: 1 male, 
Alachua Co., Apr; 1 female, 3 males, Elfers, 
Apr; 1 male, 1 female. Everglades Nat. Prk., 
Apr; 1 male, Gainesville, Feb; 4 females, 2 
males, Hialeah, Mar; 1 female. Homestead, 
Apr; 1 female, Jacksonville, Nov; 1 female, 
Putnam Co., May; 2 females. Royal Palm, 
Jan; 1 female, Sebring, Jun; 1 male, St. Pe- 
tersburg, Mar; 1 male. Tarpon Sprs., Apr; 
2 males, 5 females, Vero Beach, Apr 
(AMNH, CNC, UNH, USNM). Georgia: 3 
females. Black Rock Mt., May; 1 female, 
Clayton, Aug; 6 females, 6 males, Mcintosh 
Co., Sapelo I., Apr; 1 female, 1 male, Rabun 
Co., Jul (CNC, USNM). Indiana: 1 female, 
1 male, Lafayette, Jun (USNM). Maryland: 
1 male. Glen Echo, May; 1 male. Ft. Wash- 
ington, May (USNM), Massachusetts: 1 fe- 
male, Athol, Jul; 1 female, 1 male. Woods 
Hole, Aug (AMNH, USNM). Michigan: 1 
female, Ann Arbor, Sep; 1 female, Bath, 
Jun; 1 female. Branch Co., May; 1 female, 
Cheboygan Co., Aug; 2 females, Clinton Co., 
Jun; 1 female, Detroit, Jun; 1 female, Isle 
Royale, Aug; 1 female, Manistee, Jul; 3 
males. Midland Co., Jun; 1 female, Notta- 
wa, Jun; 1 female, Traverse Co., 1 female, 
Wayne Co., Jul (CNC, UKL, USNM). Min- 
nesota: 3 females, Basswood Lk., Jul; 1 male, 
Cass Co., Jul; 2 males, Itaska, Jul; 1 female. 
White Bear, Jul (UMSP). New Hampshire: 
1 male, Dixville, Jul; 1 male, 1 female. White 
Mts., Stinson Lk., Jul (UNHC, USNM). 
New Jersey: 1 female, Brookside, Sep 
(AMNH). New York: 1 male, Canajoharie, 
Jul; 1 male, 1 female, Ithaca, Jul; 1 female 
Lk. Sebogan, Aug; 1 female, Ludlowville, 
Jul; 1 male, Peekskill, Jul; 1 female, Rome, 
Jun; 1 female, Oneotona, Aug; 1 male, Sho- 
kan, Jul; 1 female Thomkins Co., Aug (CNC, 
CUIC, USNM). North Carolina: 1 male. 
Bubbling Spr. Crk., Jul; 7 females, 3 males. 
Highlands, Jun-Jul; 8 females, 2 males, 
Looking Glass Rock, Jul; 4 males, 8 fe- 
males. Nags Head, May; 1 female, Wayah 
Gap, Macon Co., Jul (CNC, USNM). Rhode 
Island: 1 female, 2 males. Westerly, Jul 
(USNM). South Carolina: 1 male, Mt. Rest, 
Oconee Co., Jul (CNC). South Dakota: 2 
males, Pierre, Jul (UMSP). Tennessee: 5 fe- 
males, 2 males, Gatlingburg, Jun-Jul 
(USNM). Virginia: 2 males, Richmond Co., 
Jul (USNM). Wisconsin: 1 female, 1 male, 
Vilas Co., Jul (UWM). 
Distribution. —This species is widely dis- 
tributed in eastern North America, occur- 
ring from extreme southern Manitoba and 
the northern Great Plains of the United 
States east to the Atlantic coast and south 
into southern Florida (Fig. 1 8). 
Remarks.— Owing to the nearly white 
arista on many specimens, C elongata 
probably was the species considered to be 
Chelipoda albiseta, a Palearctic species in- 
cluded in earlier state lists of insects. Me- 
lander (1947) expressed doubt that females, 
with a "strikenly white arista" were C. al- 
biseta, suggesting instead that "when the 
male with a white arista is discovered a new 
name probably will be required." My ex- 
amination revealed that variation exists in 
arista coloration of males of C. elongata, 
and perception of whiteness or darkness of 
a given arista is influenced by the incidence 
of light as specimens are rotated under the 
microscope. 
Specimens of C elongata have been col- 
lected from Betula glandulosa Michx. (tun- 
dra dwarf birch) in Manitoba, during sweep- 
ing of foliage ofQuercus laevis Walt. (Turkey 
oak) in Florida, and in light traps. 
Chelipoda limitaria MacDonald, 
New Species 
(Fig. 4) 
Diagnosis. — Absence of crossvein dm-cu 
in both wings distinguishes adults of this 
species and C americana from those of oth- 
er Chelipoda treated here. The distinction ' 
between C limitaria and C. americana is 
based on comparative structure of male 
VOLUME 95, NUMBER 3 
339 
genitalia, as presented in the diagnosis sec- 
tion pertaining to C. americana, with males 
of C limitaria possessing phallic processes 
that are longer than the cerci and lacking 
black, distal projections on the subepandrial 
lobes, the tips of which do not cross. 
Description. — MALE: Body length in- 
cluding terminalia ca. 1.8 mm. General col- 
or yellow to yellowish brown. Head: black, 
except for yellow mouthparts and palps; 
bristles yellow. Antennal scape and pedicel 
yellow; flagellum and arista brown. Thorax: 
short, compact; bristles yellow; scutum yel- 
lowish brown, darker brown along noto- 
pleural suture and at apex; scutellum and 
postnotum brown. Legs: yellow, except dis- 
tal tarsomere light brown. Fore femur ven- 
trally with 2 rows of black setulae, each 
flanked by row of 4-5 yellow bristles (see 
Fig. 5). Fore coxa lacking prominent ba- 
solateral seta, but with row of setae con- 
tinuing distally (see Fig. 6). Wing: hyaline; 
crossvein dm-cu absent; cell dm open (see 
Fig. 1). Abdomen: terga greyish brown; ster- 
na paler brown. Terminalia (Fig. 4) yellow, 
compact; hypandrium and epandrium al- 
most completely separate; cercus fused to 
epandrial lobe, subequal in length to subep- 
andrial lobe, slender and ending in 3 strong 
setulae; subepandrial lobe horseshoe-shaped 
in dorsal view; phallus ca. % length of phal- 
lic process, not extending past epandrial lobe 
in lateral view; phallic process slender, 
pointed apically, and extending slightly past 
cercus and subepandrial lobe. FEMALE: 
length including terminalia ca. 2. 1-2.4 mm; 
general body color darker than male, thorax 
yellowish brown, darker brown along no- 
topleural suture, with dark brown median 
stripe on scutum; abdominal terga 1-6 light 
brown, terga 7-8 yellowish; cercus short (see 
Fig. 1 0); spermatheca more or less reniform, 
with spermathecal duct arising from center 
of concave surface. 
Type material. -HOLOTYPE male, la- 
belled "GA: Mcintosh Co./Sapelo Island/ 
28. IV.-9. V, 1987: MT/Live Oak Forest/ 
BRC HYM. TEAM" (CNC, holotype no. 
21333). The specimen is in excellent con- 
dition and at least some of the diagnostic 
features of terminalia are visible without 
maceration. ALLOTYPE labelled "GA: 
Mcintosh Co./ Sapelo Island/28. IV, 1987: 
MT/Live Oak Forest/BRC. HYM. TEAM," 
deposited in CNC. PARATYPES. Georgia: 
43 females, same collection site as holotype 
(CNC). North Carolina: 1 male, Fort Bragg, 
May-Jun (CAS). Texas: 1 male (macerated 
terminalia in glycerin microvial), 12 fe- 
males, Montgomery Co., Jones St. For., 8 
mi S. Conroe, Apr (CNC). 
Distribution. — Males of this species are 
known only from the coastal plain of south- 
eastern Texas and Georgia, and the inland 
coastal plain of North Carolina (Fig. 16). 
Etymology. —The specific epithet is in 
reference to the apparent limited distribu- 
tion of this species. 
Remarks. —The existence of this new spe- 
cies was brought to my attention by Ralph 
Idema and Brad Sinclair (Biological Re- 
sources Division, Ottawa) after their ex- 
amination of macerated terminaha of spec- 
imens labelled ''americana'" that had been 
dissected in preparation of the figures for 
this paper. The holotype and nearly all the 
paratypes in the type series were taken in 
Malaise traps set up in a live oak forest by 
members of the "Hymenoptera team" at the 
Biological Resources Centre, Agriculture 
Canada. 
Chelipoda praestans Melander 
(Figs. 6, 14) 
Chelipoda praestans Melander, 1947: 267. 
Diagnosis.— Adults resemble those of C 
contracta in size and coloration, but possess 
only one ventral row of black setulae on the 
fore femur and the female cerci are long. 
Males superficially resemble those of C. si- 
caria, particularly when the anterior por- 
tions of the terminalia of the latter project 
ventrally into the pregenital abdominal seg- 
ments and are not clearly visible. Macerated 
terminalia of these two species are distinc- 
340 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
l<.oelof 
idema is9i 
Figs 14 15 14a (lateral) and b (dorsal), Chelipoda praestans male terminalia. 1 5a (lateral) and b (dorsal), 
Chelipodasicaria male termmal.a. ce = cercus; ph = phallus; subep = subepandnal lobe. Scale bar - 0.25 mm. 
VOLUME 95, NUMBER 3 
341 
tive, however. Those of C praestans possess 
a uniformly slender phallus in dorsal view 
and very small subepandrial lobes, whereas 
those of C sicaria possess a phallus that is 
distinctly expanded apically and prominent 
subepandrial lobes that are at least half the 
length of the phallus. In areas of sympatry, 
identification of females of these two species 
is difficult and must be based on close com- 
parison, with the much paler antennal arista 
and paler thoracic setae being the most re- 
liable characters for distinguishing females 
of C. praestans from those of C. sicaria. 
Description. — Length including termina- 
lia of male ca. 2.0-2.4 mm, of female ca. 
1.8-2.4 mm. General body color yellow to 
yellowish grey. Head black, except for yel- 
lowish grey gena and postgena; mouthparts 
and palps yellow; bristles light brown. An- 
tennal scape and pedicel yellow; flagellum 
yellowish brown; arista light brown. Thorax 
short, compact; yellowish grey, paler ven- 
trally; bristles light brown. Legs yellow. Fore 
femur ventrally with an inner row of 16-20 
black setulae and incomplete outer row of 
ca. 6-10 weaker black setulae, each such 
row flanked by row of 5-6 light brown bris- 
tles (Fig. 6). Fore coxa with 1 or 2 prominent 
basolateral setae and row of weaker setae 
continuing distally (see Fig. 7). Wing hya- 
line; crossvein dm-cu present; cell dm closed 
(see Fig. 2). Abdominal terga yellowish grey, 
sterna yellow. Male terminalia (Fig. 14) yel- 
low, projecting anteriorly nearly to abdom- 
inal segment 5; hypandrium and epandrium 
completely fused; cercus and epandrial lobe 
separate; cercus slender, curved ventrally 
distally, subequal in length to phallus; sub- 
epandrial lobe ca. '/,o length of phallus; 
phallus slender, strongly curved ventrally 
over distal 'A in lateral view; phallic pro- 
cesses lacking. Female cercus long (see Fig. 
1 1); spermatheca hemispherical, with sper- 
mathecal duct arising from center of flat sur- 
face (see Fig. 1 0). 
Type material examined. — HOLOTYPE 
male, labelled "Redding CT/ 10 June '29/ 
A L Melander" (USNM). The specimen is 
in excellent condition and most of the di- 
agnostic features of the terminalia are vis- 
ible without maceration. Two females on 
the pin with the holotype and labelled "C 
praestans ALLOTYPES" were females of 
C contract a. PARATYPES. CANADA. 
Ontario: 1 male (lacking head), Waubam- 
ick, Jul (USNM). UNITED STATES. Con- 
necticut: 7 females, 6 males (1 pin also in- 
cludes a C contracta male and another pin 
also includes 2 C contracta females), Redd- 
ing, May-Jun (USNM). Massachusetts: 9 
females, 1 male (one female labelled "al- 
lotype Chelipoda contracta''). Petersham, 
Jul (USNM). Maine: I female. Seal Harbor, 
Jul (USNM). New Hampshire: 1 female, Mt. 
Monadnock, Jul; 2 females, Mt. Washing- 
ton, Jul; 1 female, Pinkham Notch, Jul 
(USNM). New York: 1 female, 2 males. Bear 
Mt., Jul (USNM). Pennsylvania: 1 female, 
Chester Co. (USNM). 
Other specimens examined.— CANADA. 
Newfoundland: 2 females, 1 male, St. Johns, 
Jul (CNC). Nova Scotia: 2 females, 6 males, 
Cape Breton Highlands Nat. Prk., Jul; 5 fe- 
males, 4 males. Cranberry I., Jul; 1 female, 
1 1 males, Lockeport, Jul-Aug; 2 females, 6 
males. Lone Shieling, Jun-Jul; 1 female, 1 
male, Springfield, Jun-Aug; 2 males, 12 fe- 
males, St. Anne De Ruisseau, Jul (CNC, 
USNM). Ontario: 3 females, 1 male, Iro- 
quois Falls, Jun (CNC). Quebec: 1 female, 
Beechgrove, Jun; 1 male. Old Chelsea, Jun 
(CNC). UNITED STATES. Georgia: 2 
males, Rabun Bald, Jul-Aug (CNC). Maine: 
12 females, 14 males. Seal Harbor, Mt. Des- 
ert I., Jul (CNC). New Hampshire: 4 males, 
2 females, Wonalancet, Jul (UNH). New 
York: 6 females, 8 males. Lake Placid, Jul 
(CNC). North Carolina: 1 female, Devils 
Court House, Aug; 36 females, 13 males. 
Highlands, May-Aug; 1 female, Lake Tax- 
away, Jul; 1 female, 1 male, Yancey Co., 
Mt. Mitchell, Jun; 2 females, 1 male, Wayah 
Gap, Jul (CNC, UKL). Pennsylvania: 1 fe- 
male. Spring Bridge, Jun; female/male in 
342 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
C2 
h 
y-^'i 
o-.m 
W' 
/"""nti. 
'1J- 
.--^-..A 
O Chelipoda americana males 
A Chelipoda limitaria males 
C. americana or limitaria females 
v 
( \ V 
\-0' 
€^ 
Fig. 16. Distribution oi Chelipoda americana and Chelipoda limitaria. 
O 
i::^ 
copula, Luzerne Co., Bicketts Glen Prk., Jul ia: 1 female, Giles Co., Mt. Lake Biol. Sta., 
(UKL). Tennessee: 1 female, Clingman's Jun; 1 female, 3 males, Hawksville, Jun 
Dome (CNC); 1 male, Great Smoky Mts. (CNC). 
Nat. Prk., Beech Gap, Jul (USNM). Virgin- Distribution.— This species occurs from 
VOLUME 95, NUMBER 3 
343 
a 
o\ 
d-Q- 
y --< 
'") 
b 
a- 
O^'O 
^^" 
v^. 
V-- 
l \ 
•- 
\. 
X 
r"-x..v; 
■.A\ 
--AX' 
^^^- 
Fig. 1 7. Distribution of Chelipoda contracta. 
344 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Fig. 1 8. Distribution of Chelipoda elongata and Chelipoda truncata. 
VOLUME 95, NUMBER 3 
345 
Fig. 19. Distribution of Chelipoda praestans and Chelipoda sicaria. 
346 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
southeastern and extreme eastern Canada, 
south along the Appalachian Mountains into 
northern Georgia (Fig. 19). 
Remarks. —Specimens of this species have 
been taken from flowers of Castenea pumila 
(L.) Mill, (eastern chinquapin) in Great 
Smoky Mountains National Park and from 
aphid-infested Veratrum sp. near Lake 
Placid, New York, but no prey records exist. 
Chelipoda sicaria Melander 
(Fig. 15) 
Chelipoda sicaria Melander, 1947: 268. 
Diagnosis. —Males and females of C si- 
caria closely resemble those of C praestans. 
Adults of both species possess only one 
complete row of black setulae ventrally on 
the fore femur and the female cerci are rel- 
atively long. Identification of males is based 
on terminalia differences, but females are 
difficult to place; discussion of the charac- 
teristics employed is presented in the di- 
agnosis section pertaining to C praestans. 
Description. — Length including termina- 
lia of male ca. 2.0-2.4 mm, of female ca. 
2.4-2.8 mm. General body color yellow to 
dark reddish brown. Head brownish black 
to reddish black; mouthparts and palps light 
reddish black; bristles dark brown. Anten- 
nal scape and pedicel yellow; flagellum yel- 
lowish brown; arista dark brown. Thorax 
short, compact; yellow to dark reddish 
brown; bristles dark brown. Legs yellow to 
light brownish black. Fore femur ventrally 
with an inner row of 16-20 black setulae 
and an incomplete outer row of ca. 6-10 
weaker black setulae, each such row ffanked 
by row of 5-6 light brown bristles (see Fig. 
6). Fore coxa with 1 or 2 prominent baso- 
lateral setae and row of weaker setae con- 
tinuing distally (see Fig. 7). Wing hyaline; 
crossvein dm-cu present; cell dm closed (see 
Fig. 2). Abdominal terga brownish to 
brownish black, sterna light brown to light 
brownish black. Male terminalia (Fig. 15) 
yellowish brown to brownish black, pro- 
jecting anteriorly to abdominal segment 7 
or projecting ventrally into pre-genital ab- 
dominal segments; hypandrium and epan- 
drium completely fused; cercus and epan- 
drial lobe separate; cercus curved ventrally 
at apex, subequal in length to phallus; point- 
ed subepandrial lobe upturned in lateral 
view, ca. Vi length cercus; phallus sinuate in 
lateral view, prominently expanded later- 
ally at distal V3 in dorsal view; phallic pro- 
cesses lacking. Females paler than concur- 
rently collected males; cercus slightly longer 
than width at base; spermatheca hemispher- 
ical, with spermathecal duct arising from 
center of ffat surface (see Fig. 10). 
Type material examined.— HOLOTYPE 
male, labelled "Gt. Smokie NP/Newfnd 
Ridge/ 1 1 July '41/A L Melander" (USNM). 
The specimen is in excellent condition and 
most of the diagnostic features of the ter- 
minalia are visible without maceration; the 
elaborated type locality is Great Smoky 
Mountains National Park, Newfound Ridge, 
along the border of Tennessee and North 
Carolina. ALLOTYPE, same data as ho- 
lotype (USNM). PARATYPES. North Car- 
olina: 1 male (lacking head). Great Smoky 
Mts. Nat. Prk., Andrews Bald, Jul; 1 female, 
1 male (lacking head). Great Smoky Mts. 
Nat. Prk., Newfound Gap, July (USNM). 
Other specimens examined. — UNITED 
STATES. North Carolina: 14 males, 17 fe- 
males, Great Smoky Mts. Nat. Prk., Balsam 
Mt., Jun (PERC); 6 males, 9 females, Great 
Smoky Mts. Nat. Prk., Clingman's Dome 
(CNC). North Carolina-Tennessee border: 
6 males, 8 females. Great Smoky Mts. Nat. 
Prk., 1 mi S. Newfound Gap, Jun (USNM); 
3 males. Great Smoky Mts. Nat. Prk., New- 
found Ridge, Jul (USNM). Tennessee: 12 
males, 9 females, Great Smoky Mts. Nat. 
Prk., Gatlinburg, Jun-Jul (USNM). 
Distribution. —This species is known only 
from the higher elevations of Great Smoky 
Mountains National Park (Fig. 1 9). 
Remarks.— Adults of this species vary 
greatly in coloration, from nearly concol- 
orous yellow (most specimens in collec- 
tions) to nearly concolorous dark reddish 
VOLUME 95, NUMBER 3 
347 
brown, with males darker in coloration than 
concurrently collected females. Examina- 
tion of macerated males representing the 
range of coloration revealed identical ter- 
minalia. Included in the examination were 
two series collected 33 years apart from near 
Clingman's Dome, Great Smoky Moun- 
tains National Park: males in one series col- 
lected June 18, 1957 were dark reddish- 
brown; males in another series collected June 
15, 1990 were yellow to yellowish brown. 
Specimens in the June 15, 1990 series, 
mentioned above, were swept off foliage in 
late afternoon, in dense, fern undergrowth 
along a small stream under dense hardwood 
canopy. Another series, taken June 1 3, 1 990, 
below Balsam Mt. campground were swept 
off emergent vegetation in a seepage area 
along Flat Creek under a hardwood canopy, 
again in late afternoon. 
Chelipoda tmncata MacDonald, 
New Species 
(Figs. 8, 13) 
Diagnosis. —Adults closely resemble those 
of C. elongata, but differ in having entirely 
yellow tarsi and in possessing a distinct pale 
area (usually golden yellow) on the median 
margins of the postgenae. Male terminalia 
of these two species are similar superficially, 
and may require maceration in order to re- 
solve them. Terminalia of males of C. tnm- 
cata include strongly sclerotized, blunt sub- 
epandrial lobes and phallic processes that 
are about two-thirds the length of the phal- 
lus, whereas those of male C. elongata in- 
clude pointed subepandrial lobes and phal- 
lic processes that are subequal to the length 
of the phallus. 
Description. — MALE: length including 
terminalia ca. 2.0-2.4 mm. General body 
color yellow. Head: nearly black, except yel- 
low area on frons and yellow area approach- 
ing a stripe on mid ventral aspect of post- 
gena (Fig. 6); gena yellow grading into black, 
with fringe of white hair; vertex and occiput 
black; mouthparts and palps yellow; bristles 
yellow. Antennal scape and pedicel yellow; 
flagellum yellowish brown, slightly darker 
apically; arista light brown. Thorax: long, 
slender (notopleural suture ca. Vi length of 
scutum and scutellum); yellowish brown 
dorsally, paler ventrally; bristles yellow. 
Legs: yellow; fore femur ventrally with 2 
rows of black setulae, each flanked by row 
of 6-8 brown bristles (see Fig. 5); fore coxa 
lacking prominent basolateral seta, but with 
row of setae continuing distally (see Fig. 6). 
Wing hyaline; crossvein dm-cu present; cell 
dm closed (see Fig. 2). Abdomen: yellowish 
terga, sterna lighter yellow. Male terminalia 
(Fig. 1 3) yellowish, projecting anteriorly to 
abdominal segment 7; hypandrium and 
epandrium completely fused; cercus and 
epandrial lobe separate; cercus thicker api- 
cally, shorter than epandrial lobe; epandrial 
lobe slender, truncate and sclerotized api- 
cally, longer than cercus and shorter than 
phallus; phallus and phallic process slender, 
smoothly curved ventrally in lateral view, 
phallic process ca. % length of phallus. FE- 
MALE: length including terminalia ca. 2.2- 
2.6 mm; flagellum brown, arista light brown; 
cercus long (see Fig. 1 1 ); spermatheca nearly 
hemispherical, with spermathecal duct aris- 
ing from center of flat surface. 
Type material. -HOLOTYPE male, la- 
belled "Warwomen Cr. GA. /Rabun Co. 
1500731. VIL 1957/J. G. Chillcott" (CNC, 
holotype no. 21334). The specimen is in 
good condition and most of the diagnostic 
features of the terminalia are visible without 
maceration. ALLOTYPE, labelled "Rabun 
Bald, GA./Rabun Co. 3000714-VII-1957/ 
J. G. Chillcott" (CNC). PARATYPES. 
CANADA. Ontario: 1 male, Griffith, Jul 
(CNC). Quebec: 1 male, Hull, Aug(USNM). 
UNITED STATES. Georgia: 1 male (mac- 
erated terminalia in glycerin microvial at- 
tached to pin), Rabun Co., Jul (CNC). Ken- 
tucky: 2 males, Kentucky Ridge St. For., 
Jun (USNM). Minnesota: 4 females, 1 male, 
Basswood Lk., Jul-Aug (UMSP). North 
Carolina: 1 female. Looking Glass Rock, 
Pisgah Nat. For., Jul; 2 females, Macon Co., 
Jul-Aug (CNC). Virginia: 1 male, Alexan- 
348 
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
dria Co., Jun (USNM). Wisconsin: 1 female, 
Waupaca Co., Aug (UWM). 
Distribution. —This species is largely 
sympatric with C elongata, but it is known 
only as far south as the mountains of north- 
em Georgia (Fig. 1 8). 
Etymology.— The specific epithet is in 
reference to the structure of the subepan- 
drial lobes, each of which is truncate and 
heavily sclerotized at its apex. 
Concluding Remarks 
Adults of species of Chelipoda are thought 
to be predacious because of their raptorial 
fore legs, but no records of prey capture ex- 
ist. Collecting data on labels reveal that nu- 
merous specimens have been taken off flow- 
ers or foliage of plants infested with aphids. 
Other specimens have been swept off' low 
vegetation, usually in forested areas, and 
some labels make reference to swampy ar- 
eas and sloughs. In addition, specimens have 
been taken in Malaise traps, in yellow pan 
traps, and at lights. During the present study, 
adults of C. sicaria were swept and aspirated 
from understory foliage at higher elevations 
in Great Smoky Mountains National Park, 
where they appeared to be active only dur- 
ing mid- to late afternoon. 
Larvae and pupae of only one species, the 
Palearctic Phyllodromia melanocephala 
Fab., are described, having been collected 
in moist humus under beech trees in Europe 
(Trehen 1969). No immature stages of Ne- 
arctic species of Chelipoda have been de- 
scribed, but Harper (1980) presented indi- 
rect evidence that the larvae of three Nearctic 
species may be aquatic. During his study of 
insects in a Laurentian stream system in 
Quebec, Canada, Harper collected a few 
small series of C. contracta, C. elongata and 
C. praestans in emergence traps placed over 
streamlets, including one that was inter- 
mittent. 
The findings by Harper (1980) and the 
discovery of relatively large numbers of 
adults of C. sicaria in a specific habitat in 
Great Smoky Mountains National Park, re- 
ported above in the remarks section per- 
taining to this species, could encourage fu- 
ture biological studies of this group which 
has received little attention. Awareness of 
an afternoon period of adult activity in a 
specific habitat, namely understory foliage, 
could facilitate study of both reproductive 
and predatory behavior. Knowledge of a 
probable larval developmental site for Ne- 
arctic species of Chelipoda, namely stream- 
lets (including intermittent ones), will allow 
focused efforts to collect the immature 
stages, but special techniques will be re- 
quired for specimens of such small size in 
order to pinpoint a specific microhabitat, if 
one exists. 
Relatively few phylogenetically impor- 
tant characters have been revealed in the 
Hemerodromiinae, but components of male 
terminalia and shape of the female sper- 
matheca appear to be of value. For example, 
preliminary evaluation suggests that Che- 
lipoda americana and C. limitaria comprise 
a monophyletic group distinct from other 
Chelipoda treated here. This contention is 
not founded upon wing venation. Instead, 
it stems from their shared possession of a 
reniform spermatheca, lack of complete fu- 
sion of the hypandrium and epandrial lobes, 
and fusion of each cercus to a corresponding 
epandrial lobe. Importantly, adults of Phyl- 
lodromia melanocephala, the type species 
of Phyllodromia Zetterstedt established pri- 
marily on the basis of the lack of crossvein 
bm-cu, differ in other respects from those 
of C americana and C. limitaria. Females 
of P. melanocephala lack the reniform sper- 
matheca characteristic of C americana and 
C. limitaria, and males of P. melanocephala 
possess a hypandrium and epandrium that 
are completely fused. The latter feature is 
characteristic of males of other Nearctic 
Chelipoda treated here and of males of other 
Palearctic Chelipoda. Consideration of these 
characters, together with awareness of vari- 
ation in wing venation of other hemerod- 
romine species, supports the contention that 
the present concept of '' Phyllodromia''^ based 
VOLUME 95, NUMBER 3 
349 
on wing venation is most likely invalid. Such 
awareness also leads to the conclusion that 
the present taxonomic status of other genera 
included in the "Chelipodini" (for example, 
Afwdromia Smith, Chelipodoziis Collin, 
Doliodromia Collin, Monodromia Collin, 
and Ptilophyllodromia Bezzi) might be 
questioned also, since they too are based 
largely on minor differences in venation. 
Drawing upon the interpretations of ho- 
mologies of male terminalia outlined in 
Cumming and Sinclair (1990), taxa pres- 
ently allied with Chelipoda and Phyllodwm- 
ia appear to constitute the sister group of 
the remainder of the subfamily Hemerod- 
romiinae, but the groundplan of the sub- 
family has yet to be established. 
Acknowledgments 
I thank the following individuals for the 
loan of material: P. H. Amaud, Jr. (CAS); 
R. W. Brooks (UKL); D. S. Chandler 
(UNH); P. J. Clausen (UMSP); W. J. Han- 
son (USU); L. Knutson (USNM); S. Krauth 
(UWM): J. K. Liebherr (CU); B. Lindeberg 
(HZM): R. T. Schuh (AMNH); H. J. Teskey 
and J. M. Cumming (CNC); and H. V. 
Weems (FSCA). 
Sincere appreciation is extended to dip- 
terists at the Biological Resources Division, 
Agriculture Canada, Ottawa who initially 
facilitated this study, and especially to J. M. 
Cumming and B. J. Sinclair who not only 
shared their interpretation of the homology 
of male terminalia but assisted me in un- 
derstanding the genitalic components of 
males of Chelipoda. Special thanks go to R. 
Idema and B. J. Sinclair for bringing to my 
attention the undescribed species from Tex- 
as. In addition, R. Idema not only prepared 
the excellent illustrations, but provided col- 
or-coded figures of individual components 
of male terminalia. I also thank R. Wagner 
(Max Planck Instituts fiir Limnologie, 
Schlitz, Germany) for sharing copies of his 
illustrations of Palearctic Chelipoda. This 
paper is Purdue Agricultural Experiment 
Station journal no. 11, 511. 
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