16 October 1990 
PROC. ENTOMOL. SOC. WASH. 
92(4), 1990, pp. 705-715 
SYSTEMATICS OF THE WEST INDIAN MOTH GENUS 
HEURETES GROTE AND ROBINSON 
(LEPIDOPTERA: LIMACODIDAE) 
MARC E. EPSTEIN AND SCOTT E. MILLER 
(MEE) Department of Entomology, NHB 127, Smithsonian Institution, Washington, 
D.C. 20560; (SEM) Department of Entomology, Bishop Museum, Box 19000-A, Hono- 
lulu, Hawaii 96817. 
Abstract?TYie genus Heuretes Grote and Robinson, 1868, is revised. Heuretes was 
previously monotypic, comprised of ^. picticornis Grote and Robinson, 1868, and known 
only from the Island of Saint Thomas, U.S. Virgin Islands. Monoleuca albicollis Forbes, 
1930, known from Puerto Rico, is recognized as a junior synonym o?H. picticornis. New 
records expand the range ofH. picticornis to include the island of T?rtola, British Virgin 
Islands. Heuretes daidaleos, new species, and H. divisus, new species, are described from 
the Dominican Republic. Phylogenetic analysis and transitional character states support 
placement of the new species in Heuretes. Heuretes appears to be closely related to other 
Caribbean genera Alarodia Moeschler and Leucophobetron Dyar. 
Key Words:   Puerto Rico, Virgin Islands, Hispaniola, Greater Antilles, systematics, cla- 
distics, Caribbean, Zygaenoidea 
In the Lepidoptera it is not uncommon ural History (USNM), but we have also used 
for conspecific males and females to be de- material in or checked the collections of the 
scribed in different genera as different American Museum of Natural History 
species. Epstein recognized Monoleuca al- (AMNH), Bernice P. Bishop Museum 
bicollis Forbes (Limacodidae) as the junior (BPBM), British Museum (Natural History) 
S'^nonym O? Heuretes picticornis GroXt diXia (BMNH), Carnegie Museum of Natural 
Robinson while examining a female syntype History (CMNH), Natural History Museum 
of the H. picticornis. Concurrently, field- of Los Angeles County (LACM), Museum 
work by Miller in the British Virgin Islands of Comparative Zoology, Zoologisches Mu- 
yielded good series of male specimens of a seum der Humboldt Universitaet (ZMHB), 
moth that matched the holotype of Mono- Zoologische Sammlungen des Bayerischen 
leuca albicollis, known only from males. Staates, and V. O. Becker private collection 
Later, Becker collected both males and fe- (VQB). Color descriptions follow Smithe 
males at the same locality in Puerto Rico, (1975) 
verifying the synonymy. These events, plus 
the finding of two new species from the Do- Heuretes Grote and Robinson 
minican Republic prompted us to revise and 
investigate the  systematic placement of //ewr?-i^j Grote and Robinson, 1868:190.? 
Heuretes. Kirby,   1892:  551.-Dyar,   1905:  382; 
This work is based primarily on speci- 1935:  1130.?Van Eecke,  1925: 53.? 
mens in the U.S. National Museum of Nat- Fletcher and Nye, 1982: 77. 
706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Type species.?Heuretes picticomis Grote 
and Robinson, by monotypy. 
Diagnosis.?Small moths, salmon to gray 
forewing and thorax, usually light-colored 
hindwing and abdomen. Conspicuous oval 
or quadrate spots present (at least) on male 
forewing apex or middle of inner margin. 
Male anterior thorax with dorsal collar of 
buff or white scales. Foreleg dark, with coxa 
and femur scarlet orange or dark brown. 
Mid- and hindlegs light colored. Hind tibia 
with two pairs of spurs. Male antenna bi- 
pectinate to tip or near, with light-colored 
scales in contrast at apex. 
Discussion.?Presumed synapomorphies 
of Heuretes within Limacodidae include the 
scale patterns on the body and wings. These 
patterns include a distinct collar behind the 
head in males, dark-scaled forelegs in con- 
trast to light mid- and hindlegs, and light- 
colored scales at the apex of the antenna in 
contrast to dark segments on all other seg- 
ments or only near the apex. 
Although found in other limacodid gen- 
era, the forewing radial vein pattern in Heu- 
retes, R3+R4 branched off R2, may be in- 
dependently derived. Presumed sister genera 
Alarodia Moeschler, Leucophobetron Dyar 
and Phobetron Huebner, possess wing ve- 
nation pattern in which R3+R4 branch off 
R5 or nearby?the primitive condition ac- 
cording to Brock (1971). 
Heuretes picticornis Grote and Robinson 
Figs. 1, 2, 5, 6, 9 
Heuretes picticornis Grote and Robinson, 
1868: 190. Grote, 1882: 17; 1888: 182. 
Dyar, 1905: 382; 1935: 1130. Van Eecke, 
1925: 53. 
Monoleuca albicollis Forbes, 1930: 166. 
Dyar, 1935: pi. 168, fig. g [not mentioned 
in text]. Wolcott, 1936: 505; 1951: 746. 
Martorell, 1948: 549; 1976: 33, 51, 175, 
252. NEW SYNONYMY, NEW COMBINA- 
TION. 
Diagnosis. ?Small moths with either 
salmon or dark-brown forewings. Front legs 
orange. Pale-form males have antemedian 
and postmedian bands on forewing visible, 
cream-colored hindwing; dark-form males 
with forewing bands not visible, either cream 
or dark-brown hindwing. Both male forms 
with conspicuous quadrate white patch me- 
dially along inner margin. Female without 
white patch. Male antenna broadly bipec- 
tinate, becoming unipectinate with branch- 
es abruptly shortening toward apex, scarlet- 
orange scales on shan. Uncus unusually 
short, baglike, with lateral socii; gnathos 
S-shaped laterad. Valva bifid distally; ae- 
doeagus straight basally, reaching end of 
valva, upturned distally. 
Adult male (Fig. 1).?Forewing length 6- 
7 mm (one aberrant reared male, 8 mm). 
Head: Frons mostly white. Vertex cream 
colored (54) (= color code in Smithe 1975). 
Antenna ca. half length of forewing; bipec- 
tinate, maximum width about 5 x length of 
a segment, slightly tapering to near apex, 
then abruptly short. First two short apical 
segments bipectinate, remaining nine seg- 
ments short, unipectinate, and bristly. Scales 
on shaft scarlet orange; scales on apex and 
pectinations buff (124). Labial palpus por- 
rect, extending past frons, covered with 
scarlet-orange scales. Haustellum coiled, 
longer than basal segment of labial palpus. 
Thorax: Dorsum with white scale bundle 
behind head on basal one-fifth, remainder 
matching forewing. Foreleg with coxa and 
femur scarlet orange, tibia and tarsus burnt 
orange (116) with dark bands distally. Mid- 
leg and hindleg a lighter cream color, with 
brushlike scales dorsally on tibia and tarsus. 
Midtibia with one pair of spurs and hind- 
tibia with two pairs. Forewing R3 and R4 
nearly fused, arising from R2 (Fig. 5). Dor- 
sal surface of forewing in two forms, either 
salmon or dark brown. Diffuse postmedian 
and antemedian bands of dark-brown scales 
visible only in pale morph (Fig. 1). These 
bands connect, with mostly dark-brown 
scales from wing base to tomus, below dis- 
cal cell. Conspicuous white quadrate patch 
on middle of inner margin, smaller dark 
VOLUME 92, NUMBER 4 707 
Figs. 1^. 1, Heuretes picticornis, male wing pattern (USNM) (forewing 6 mm). 2, Heurelespicticornis, female 
wing pattern (USNM) (forewing 8 mm). 3, Heuretes daidaleos, male wing pattern (USNM) (forewing 6 mm). 4, 
Heuretes divisus, male wing pattern (CMNH). 
spot where M2 and M3 arise from discal 
cell. Margins orange scarlet, preceded by 
dark brown scales on fringe and costal mar- 
gin, scattered orange scales along veins. 
Ventral forewing with inner margin cream 
colored, base of costa scarlet, otherwise 
burnt orange. Dorsal hindwing cream col- 
ored, or dark brown, outer margin fringe 
with a few dark scales. Salmon-forewing 
morph with pale hindwing, dark-forewing 
morph with either pale or matching dark 
hindwing. Ventral hindwing warm buff ( 118) 
with hint of orange, more cream color along 
inner margin and fringe. 
Abdomen: Cream colored. Genitalia as in 
Fig. 6. Valva bifid about three-fourth dis- 
tance from base. Gnathos simple, with gently 
undulating margins. Uncus short, expanded 
ventrally above gnathos, caudal end of un- 
cus divided. Socii forming lateral pockets 
with setae. Aedoeagus straight and about 
same length as valva; stout basally, but be- 
yond gnathos, becoming narrow, upturned 
90? distally. 
Adult female (Fig. 2).?Forewing length 
7.5-8.0 mm. 
Head: Frons salmon colored (106). Ver- 
tex cream colored. Antenna short, ca. one- 
708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Fig. 5. Heuretes picticornis, male wing venation 
(USNM 28034). 
third length of forewing, bristly, with short 
pectinations, curved at tip, cream colored 
to near apex, dark-brown scales and lighter 
scales distally, nearly buff. Palpus shaped as 
in male, covered with warm-buff and salm- 
on scales. 
Thorax: Dorsum with brown scales, with- 
out white scale bundle of male. Legs colored 
as in male. Forewings as in male dark morph, 
less scarlet underneath, with white scale 
patch absent. Discal spot faint. Only pale 
hindwings known. 
Abdomen: Genitalia as in Fig. 9. Bursa 
copulatrix short, with ductus seminalis 
broadly connected from base to middle of 
corpus bursae. Signum absent. Lobes of pa- 
pillae anales disk shaped, irregularly toothed 
on margins, and covered with setae. 
Types.?Lectotype female, here desig- 
nated (ZMHB), and paralectotype female 
(AMNH) (//. picticornis); Holotype male, 
AMNH (M. albicollis). 
Type localities.?[U.S. Virgin Islands], 
[Island of] Saint Thomas (H. picticornis); 
Puerto Rico, Coamo Springs {M. albicollis). 
Hosts.?Buchenavia capitata (Vahl) Eichl. 
(Combretaceae) (USNM); Byrsonima cori- 
?cea (Sw.) DC (Malpighiaceae) (Martorell 
1948: 549, as B. spicata (Cav.) Rich.; Wol- 
cott 1951: 746, as B. spicata; Martorell 
1976); Cedrela odorata Linn. (Meliaceae) 
(Wolcott 1951: 746, as C. mexicana Roem.; 
Martorell 1976); Montezuma speciosissima 
Sess? & Moc. (Malvaceae) (Wolcott 1951: 
746; Martorell 1976); Swietenia mahagoni 
(Linn.) Jacq. (Meliaceae) (Martorell 1948: 
549, 1976). 
Immature stages.?This description of the 
previously undescribed larva is from an 
exuvium found inside a cocoon associated 
with an adult male ("Hopk. US 33101K," 
USNM). The exuvium was softened in 10% 
KOH and stored in glycerin (descriptions of 
tubercles are vague because precise inter- 
pretation was difficult). 
Final instar: No prolegs or crochets, sub- 
dorsal row of tubercles well developed (T2- 
3, A2-9), extending to near middorsum, lat- 
eral tubercles short. Most subdorsal tuber- 
cles with long, filamentous hairs, maximum 
length ca. 1 mm, dark brown and light col- 
ored, with short lateral branches. Second or 
third subdorsal tubercle from anterior end 
with dense coat of short, filamentous hairs 
interspersed with long hairs, appearing 
darker than others. Unbranched urticating 
setae well developed on dorsal, basal por- 
tion of tubercles. Middorsal strip with small 
"skin spines." 
Cocoon: Ovate, about 5 mm long, 4 mm 
wide, 4 mm high. Typical circular emer- 
Figs. 6-8. 6, Heuretes picticornis, male genitalia: a) lateral view, right aspect (dotted outline indicates position 
of aedoeagus) (USNM 28109); b) aedoeagus (USNM 28025); c) ventral view of slide mounted specimen with 
aedoeagus removed (g = gnathos; s = socius) (USNM 28025) (scale = 0.5 mm). 7, Heuretes daidaleos, male 
genitalia: a) lateral view, left aspect (USNM 28110); b) aedoeagus (USNM 28358); c) ventral view of slide 
mounted specimen with aedoeagus removed, right valva (p = valval process) (USNM 28358) (scale = 0.5 mm). 
8, Heuretes divisus, male genitalia: a) lateral view, right aspect (CMNH); b) aedoeagus (CMNH) (scale = 0.5 
mm). 
VOLUME 92, NUMBER 4 709 
710 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Fig. 9.   Heuretes picticornis, female genitalia (ZMHB lectotype): a) lateral view; b) ventral view (c = corpus 
bursae, d = ductus seminalis; c) papillae anales (scale = 0.5 mm). 
gence hatch at end. Whitish, mottled with 
brown, surface smooth and hard. 
Flight period.?December to August. 
Distribution.?Puerto Rico and the Vir- 
gin Islands, above 360 m. 
Material examined.?65 males and 5 fe- 
males. PUERTO RICO: Coamo Springs, 5- 
7 June 1915 (AMNH, holotype of M al- 
bicollis); El Semil, near Villabla, 1700 feet 
[520 m], lO-V-1940, W. A. Hoffman 
(USNM); El Verde Field Station, Luqillo 
Experimental Forest, 435 m, 29-XII-1970 
to 21 -I-1971, C. P. Kimball (USNM); Hotel 
Barranquitas, Barranquitas, 650 m, 22-26- 
11-1971, Kimball (USNM); Maricao, 11-14- 
VIII-1987, V. O. Becker (VOB); Maricao 
Fish Hatchery, 23-XII-1962, P. & P. Span- 
gler (USNM); Reserva Forestal Guajataca, 
360 m, 18-28-III, 14-20-IV-1971, Kimball 
(USNM); Patillas, [Sierra de Cayey, ca. 20 
km NE Campamento Real], 590 m, 5-25- 
VIII-1987, Becker (USNM, VOB); Villabla, 
ex Buchenavia capitata, Hopkins no. 
33101K, l-V-1940 (USNM); U.S. VIRGIN 
ISLANDS: [Island of] Saint Thomas, [no 
further data] (ZMHB, lectotype of H. pic- 
ticornis; AMNH, paralectotype); BRITISH 
VIRGIN ISLANDS: [Island of] T?rtola, 
Mount Sage National Park, 480 m, 7-8-VII- 
1985, S. E. &P. M. Miller, 22-24-VII-1986, 
S. E. Miller & M. G. Pogue, 13-15-VII- 
1987, S. E. Miller & V. O. Becker, ultra- 
violet light trap in "aridulate rain forest" 
(USNM, BMNH, CMNH, LACM, VOB, 
BPBM). 
Discussion.?The scarlet orange front 
coxa and femur in both male and female 
led to uncovering the synonymy between 
VOLUME 92, NUMBER 4 711 
Heuretes Heureles 
picticomis 
Leucophobetmn 
Fig. 10. Preliminary cladogram of genera related to 
Heuretes {Leucophobetron based on type species only). 
Synapomorphic characters include: 1) urticating spines 
in late instar larvae both branched and unbranched 
(known only in A. slossoniae and H. picticomis); 2) 
valvae asymmetrical (reversed in H. picticomis, see 
text); 3) white groundcolor; 4) uncus with dorsal pro- 
cess; 5) uncus with simple dorsum and curved end; 6) 
forewing R3+R4 branched ofrR2; 7) dorsal collar of 
white scales behind head; 8) antennal scales light col- 
ored at apex, in contrast to preceding scales. 
Heuretes picticomis and Monoleuca albicol- 
lis. Collection of males and females at the 
same localities in Puerto Rico by Becker 
(see above) confirmed this finding, since the 
two types are from separate, though neigh- 
boring, islands of Saint Thomas and Puerto 
Rico. 
This species is restricted to the Puerto 
Rican Bank, where it occurs between 360 
and 2560 m. Martorell (1948: 549) [as M. 
albicollis] considered it "fairly common" on 
Puerto Rico. In the United States Virgin 
Islands the species has not been recorded 
since the original description of specimens 
from Saint Thomas. This could be due to 
lack of subsequent collecting and destruc- 
tion of native forest. We record it for the 
first time from the British Virgin Islands, 
from the moist forest on the top of T?rtola 
(termed "aridulate rain forest" by D'Arcy 
1967: 392). The forest in Sage Mountain 
National Park on T?rtola appears to be the 
only suitable habitat in the British Virgin 
Islands (see also Lazell and Jarecki 1985). 
The species was not found in ultraviolet Ught 
surveys on Virgin Gorda and Guana Is- 
lands. 
This distribution pattern makes sense 
biogeographically, because the islands of the 
Puerto Rican Bank have been extensively 
connected within recent geologic time. The 
principal islands of the Virgin Islands (ex- 
cept Saint Croix) lost their connection with 
each other and with Puerto Rico only about 
8000 to 10,000 years ago, due to eustatic 
rise in sea level (Heatwole et al. 1981). 
Heuretes daidaleos. NEW SPECIES 
Figs. 3, 7 
Diagnosis.?Small moth with speckled 
gray-brown forewing without fascia, hind- 
wing pale buff. Small buff-colored oval spot 
on forewing apex. Male antenna bipectinate 
to apex and gently tapering; light scales 
throughout except dark brown subapically. 
Uncus oblong, tapered to well sclerotized, 
clawlike tip. Valvae entire, slightly asym- 
metrical, curvilinear process on costal base 
and tegumen. Aedoeagus only about half 
length of valvae, with short obliquely curved 
process at distal end. Adult female and im- 
mature stages imknown. 
Adult male (Fig. 3).?Forewing length 
5.5-6.0 mm. 
Head: Frons buff colored (124), infuscat- 
ed with gray-brown scales. Vertex cream 
colored (54). Antenna length as in male H. 
picticomis, but pectinations relatively short 
and bipectinate to tip. Pectinations at max- 
imum ca. 3 X length of a segment, gently 
tapering apically. Shaft and pectinations with 
pale-buff scales except distal fourth with 
dark-brown scales, buff on last two. Labial 
palpus and haustellum similar to H. picti- 
comis except palpus buff scaled. 
Thorax: Dorsum directly behind head 
with narrow band of burnt orange scales 
712 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
(116), followed by wider collar of buff scales 
in basal seventh, remainder matching fore- 
wing. Foreleg as in H. picticornis, bands of 
dark scales alternating with buff rather than 
scarlet orange. Midleg, hindleg and thorax 
mostly buff ventrally. Tibial spurs as in H. 
picticornis. Forewing R veins as in H. pic- 
ticornis, but wing shape differs. Outer angle 
more rounded, extending posteriorly, with 
inner margin short. Dorsum with dark gray- 
brown scales speckled over glaucus (79-80) 
to pale-buff scales; only one color morph 
known. Apex with small, conspicuous, pale- 
buff oval spot (0.7 mm long). Fringe dark 
gray brown; costal margin yellow orange. 
Venter with dark scales in discal cell and 
along outer margin; buff on inner margin 
and apex. Hindwing with basal and fringe 
scales buff, hint of gray toward outer mar- 
gin. 
Abdomen: Buff colored. Genitalia as in 
Fig. 7. Valvae entire, digitate and acuminate 
distally, slightly asymmetrical, left valva 
larger. Narrow, curvilinear process articu- 
lated with base of costa and vinculum. Un- 
cus oblong, simple, tapering to downcurved 
and well-sclerotized tip. Gnathos relatively 
short, not reaching end of uncus. Aedoeagus 
short, ca. half length of valvae; globular at 
base, with short, curved hook pointing 
obliquely upward from left to right at distal 
end. 
Type.?Holotype male, USNM. 
Type locality. ?Dominican Republic, 
Dajabon Province, Rio Massacre, Balneario 
Don Miguel, 7 km SW Dajabon, 40 m el- 
evation. 
Flight period.?May aad July. 
Distribution.?Known only from the Do- 
minican Republic. 
Material examined.?Five males from the 
type locality (holotype and 4 paratypes) col- 
lected 26-V-1973 by D. & M. Davis (USNM, 
BMNH). 
Etymology.?Daidaleos (Gr.) means dap- 
pled or spotted, descriptive of the forewing 
in this species. The unlatinized nomen is 
treated as indeclinable in accordance with 
Article 31b in the International Code of 
Zoological Nomenclature (1985). 
Discussion.?T/ewre?e? daidaleos shares 
several male genitalic character states with 
Alarodia and Leucophobetron Dyar, includ- 
ing valval asymmetry and a costal process, 
absent in its congener H. picticornis. How- 
ever, phylogenetic evidence and transitional 
character states in another new species be- 
low support placement of H. daidaleos in 
Heuretes. The costal process on the valva 
is believed to be plesiomorphic with respect 
to Heuretes, found in Phobetron. 
Although the simple, oblong shape of the 
uncus of H. daidaleos appears to be closer 
to the limacodid groundplan than that found 
in H. picticornis or Alarodia, it may repre- 
sent the loss of the dorsal process found in 
the latter group. 
Heuretes divisas. NEW SPECIES 
Figs. 4, 8 
Diagnosis.?Male genitalic characters di- 
agnostic (see below). Known from one male 
specimen. 
Adult male (Fig. 4).?Forewing length 5.5 
mm. 
Head: Frons and vertex dark brown. An- 
tenna length as in other Heuretes, bipecti- 
nate to near the tip, pectinations long as in 
H. picticornis, gently tapering apically. Shaft 
and pectinations with dark-brown scales 
throughout, buff on last two. Labial palpus 
similar to other Heuretes, the former dark 
scaled. 
Thorax: Dorsum with buff scales behind 
head (remainder undetermined). Foreleg 
with dark-brown scales. Midleg, hindleg, and 
ventral thorax mostly buff Tibial spurs as 
in other Heuretes. Forewing R3 and R4 
fused, arising from R2. Dorsum appears to 
have gray and buff scales, with salmon scales 
on anterior discal cell (badly rubbed). 
Abdomen: Buff colored. Genitalia as in 
Fig. 8. Valva deeply divided less than half 
it s length from base into two thin digitate 
arms (hence the name divisus), the one ven- 
trad slightly shorter. Process articulated with 
VOLUME 92, NUMBER 4 713 
base of costa and vinculum greatly reduced 
compared to H. daidaleos. Tegumen narrow 
with long setal brush near costa. Uncus sim- 
ple, similar to H. daidaleos though broader. 
Gnathos undulated as in H. picticornis, but 
with knoblike structure ventroproximal to 
apex. Aedoeagus similar to H. picticornis, 
though relatively shorter and broader. 
Distribution.?Known only from the Do- 
minican Republic. 
Material examined.?One male (holo- 
type) from Pedernales Province, La Abeja, 
38 km NNW Cabo Rojo, 18?09'N, 7 PSS'W, 
1250 m, 15-VII-1987, J. E. Rawlins & R. 
L. Davidson (CMNH). 
Discussion.?Maie genitalic character 
states in Heuretes divisus are plausible tran- 
sitions between character states of H. dai- 
daleos and those in rather atypical H. pic- 
ticornis. The curvilinear process at the base 
of its costa, much reduced compared to H. 
daidaleos and absent in H. picticornis, ap- 
pears to be functionally replaced by the 
coastal arm of the deeply cleft valva. The 
valva in H. picticornis, in turn, may be de- 
rived from fusion of arms, as in H. divisus, 
to near the distal end. Other character states 
in H. divisus appear intermediate between 
H. daidaleos and H. picticornis, including 
shapes of uncus and gnathos, and relative 
size of aedoeagus. Shape of the aedoeagus 
more closely resembles that ofH. picticor- 
nis. 
Relationships o? Heuretes 
Forbes (1930) suggested that Monoleuca 
albicollis [H. picticornis] "might be better 
placed in Protalima," but included it in 
Monoleuca Grote and Robinson based on 
wing pattern. Similarity in male antennal 
pectinations between the type species, 
Monoleuca semifascia Walker, and M. al- 
bicollis, may have also influenced Forbes' 
generic placement. Forbes (1930) consid- 
ered M. albicollis [H. picticornis] to be a 
"primitive" Monoleuca based on the "near- 
ly united" forewing R3 and R4 (R3 and R4 
are fused in North American Monoleuca). 
Other than Forbes' treatment (1930, as 
Monoleuca), the placement of Heuretes has 
remained uncertain. Grote and Robinson 
(1868: 190) considered Heuretes near Tor- 
tricidia Packard. Dyar (1905: 382) stated 
the "generic position [of Heuretes] is un- 
certain." 
Heuretes is not congeneric with Mono- 
leuca as the latter is presently defined, nor 
is Heuretes phylogenetically related to the 
Parasa complex of Epstein (1988) where 
Monoleuca belongs. Characters of Mono- 
leuca (sensu stricto) that distinguish it from 
Heuretes are: 1) spiny type larva similar to 
those found in Euclea (Epstein 1988), 2) 
fused forewing R3+R4 arising from R5, 
rather than R2 as in Heuretes, 3) haustellum 
absent, 4) labial palpus stout, upturned, 5) 
Signum present, and 6) one pair of hind tib- 
ial spurs. 
Heuretes shares presumed apomorphic 
larval character states with Alarodia (only 
A. slossoniae known), Phobetron, and Iso- 
chaetes Dyar (Dyar 1899: 244); the "tropic 
hairy eucleids" of Dyar (1899). This group 
is defined by tubercles that are strongly de- 
veloped subdorsally and weakly developed 
laterally, and branched, filamentous hair- 
like setae. Larval Phobetron differ from 
Heuretes and Alarodia by having tubercles: 
1) with short and stout hairs only, 2) without 
unbranched urticating setae, and 3) that are 
deciduous, later incorporated into the co- 
coon. Immature characters shared between 
Heuretes and Alarodia include: 1) larval tu- 
bercles with long filamentous hairs, 2) short, 
unbranched urticating setae, and 3) white 
cocoons. Socii found on the uncus in Alar- 
odia slossoniae (Packard) and H. picticornis, 
absent in aU other taxa in both genera, are 
independently derived according to the 
phylogenetic analysis below. 
Leucophobetron argentiflua (Geyer) from 
Cuba, the type species of Leucophobetron, 
is probably congeneric with Alarodia (Ep- 
stein in prep.). Leucophobetron includes one 
other species from Colombia, Leucophobe- 
tron punctata (Druce). Though we have not 
714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
examined it, L. punctata is of uncertain 
family status (Dyar unpublished, Nat. Agri. 
Libr.; Becker and Epstein in press). There 
are a number of taxa of this general descrip- 
tion in several moth families in South 
American. Whereas, no other white species 
of Limacodidae are reported in the neo- 
tropics outside of the Caribbean basin. 
A detailed phylogeny of the genera related 
to Heuretes and resolution of generic limits 
of Alarodia and Leucophobetron are beyond 
the scope of this paper. However, we present 
the results of a preliminary phylogenetic 
analysis that we undertook to determine the 
correct generic placement of H. daidaleos 
{H. divisus was not included because it was 
discovered after completion). The analysis 
was performed using McClade, with Pho- 
betron as the outgroup. Alarodia, Leuco- 
phobetron (based on the type species only), 
and two species of Heuretes were the other 
terminal taxa. Eight characters with a total 
of 21 states were used in the analysis, and 
multistate characters were ordered with 
morphoclines. 
Heuretes daidaleos was found to be the 
sister species ofH. picticornis in two equally 
parsimonious cladograms (including Fig. 10, 
consistency index of .93). Figure 10 and a 
cladogram with Alarodia as sister group to 
Leucophobetron + Heuretes are supported 
biogeographically, since the Puerto Rican 
Bank {H. picticornis) is closer to Hispaniola 
(//. daidaleos) than it is to Cuba (Leucopho- 
betron and Alarodia), or Jamaica (Alarodia). 
Alarodia (Dyar 1897, 1935) and now 
Heuretes are the only genera in Limacodi- 
dae (ca. 300 neotropical species), with more 
than one described species in the Greater 
Antilles (Becker and Epstein in press). Two 
undescribed species, one of uncertain place- 
ment (not in Alarodia or Heuretes) from 
Cuba and the Bahamas (USNM, AMNH, 
CMNH), the other in the Perola complex 
from the Dominican Republic (CMNH), in- 
dicate that the limacodid fauna may in- 
crease with future examination. Limacodid 
genera Perola Walker and Semyra Walker 
have one described species each in the Less- 
er Antilles. There is one undescribed species 
in the Natada complex in this region. 
Poor dispersal ability of Limacodidae 
(Wood 1984), and related families Dalcer- 
idae (Miller in press) and Megalopygidae, 
may explain their low species richness in 
the West Indies. Dalceridae (85 spp.) and 
Megalopygidae (>200 spp.), mostly neo- 
tropical famiUes, have only one West Indian 
species each. 
ACKNOWLEDGMENTS 
Miller's fieldwork, as well as preparation 
of the illustrations, was supported by The 
Conservation Agency, with a grant from the 
The Falconwood Corporation. L. Laszlo 
Meszoly and Epstein illustrated the geni- 
talia. We thank the curators of the collec- 
tions consulted for use of specimens. R. W. 
Hodges, D. R. Davis, N. L. Evenhuis, and 
S. J. Weiler reviewed the manuscript. 
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