Vol. 8 No. 1 1997 AUSTIN et al: Hesperiidae of Rondonia - Entheus 5

TROPICAL , 8(1): 5-18 HESPERIIDAE OF RONDONIA, : ENTHEUS HUBNER, WITH DESCRIPTIONS OF NEW SPECIES (LEPIDOPTERA: HESPERIIDAE: PYRGINAE)

GEORGE T. AUSTIN', OLAF H. H. MIELKE2-4, and STEPHEN R. STEINHAUSER3

'Nevada State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas, Nevada 89107, USA 2Dept. de Zoologia, Universidade Federal do Parana, C. P. 19020, 81531-970 Curitiba, Parana, Brazil 3Allyn Museum of Entomology, 3621 Bay Shore Road, Sarasota, Florida 34234, USA

ABSTRACT.- Six species of Entheus were found in central Rondonia, Brazil. Four are described as new species: Entheus eunyas n. sp. of the E. eumelus group, Entheus aureanota n. sp. of the E. priassus group, and Entheus aureolus n. sp. and Entheus bombus n. sp. of the E. gentlus group. A neotype is designated for Papilio gentius (Cramer, [1777]). Entheus ninyas is returned to a species level taxon. The status of Entheus telemus is elaborated. KEY WORDS: Amazon, Apidae, , , Ecuador, Entheus aureanota n. sp., Entheus bombus n. sp., Entheus aureolus n. sp., Entheus eunyas n. sp.. Formicidae, Hymenoptera, Neotropical, , South America, Surinam, .

The Neotropical genus Entheus Hiibner, [1819], creates shaped antrum. Laterally, the lamella postvaginalis is a pair of taxonomic headaches from several directions. The striking sexual broad plates which are variously produced and dentate on their dimorphism and sympatry of several species makes association of caudal edge and spinose along their medial edge and mid venter. the sexes a challenge (Steinhauser, 1989). Compounding the The plates are covered by a membraneous flap which is distinctly problem are superficial similarities between species, few obvious striated medially (appearing like a feather). The lateral plates characters of the genitalia, and unrecognized taxa (Evans, 1952; were interpreted as the lamella antevaginalis by Steinhauser Steinhauser, 1989). Further, some of Evans' (1952) subspecific (1989) but these structures are clearly dorsad and caudad of the level taxa (and synonyms) may be valid species in themselves. ostium. The lamella antevaginalis is not distinguishable and is Sorting out this chaos requires extensive collections and detailed apparently fused with the antrum. The ductus bursae is bulbous studies of the various phenotypes to identify distributions, and constricted at the cervix. The corpus bursae has a short and sympatries, and variation and to correctly associate sexes and narrow neck and an expanded sac cephalad. taxa. In central Rondonia, Brazil, Entheus is represented by six The male genitalia of most Entheus are very similar. The species: one of the Entheus eumelus (Cramer) group, three of the tegumen has a pair of processes from the caudal end, the uncus Entheus priassus (Linnaeus) group, and two of the Entheus is divided, the upturned gnathos is spiculose on its lateral surfaces gentius (Cramer) group. Four of these represent undescribed caudad and joined ventrad on the caudal half by a lightly species. These species and certain synonymies are discussed, their sclerotized membrane, a process from the inner surface of the genitalia are illustrated, and four new species are described. costa of the valva extends caudad, the caudal end of the harpe is Forewing length is from base to apex of specimens from Rondon- dentate and curved inward, the penis has a long and slender ia, unless noted. Terminology for structures of the genitalia phallobase and a stout aedeagus with the caudal end sharply follows Steinhauser (1986); the penis is characterized by the pointed and opening dorsad, and the cornuti consist of several anterior phallobase and posterior aedeagus. Capitalized color prominent spikes. Godman and Salvin's (1879) figure omitted the names are after Smithe (1975, 1981). costal process of the valva and Evans (1952) showed this structure as a caudal extension of the vinculum (Steinhauser, Entheus eumelus Species-Group 1989). The female genitalia of Entheus have a narrow central portion A number of misconceptions are present in the literature of the lamella postvaginalis usually with a central notch on the concerning Entheus taxa characterized by Evans (1952) as having caudal margin. This is fused with a cephalad narrowing trough broadly orange males with no tibial tuft, continuous hyaline discal which ends at the caudal end of a distinct, sclerotized, and cup- and [sub] apical bands, and a macule between these bands in M3- 4. Contribution No. 898, Dept. de Zoologia, Universidade Federal do Parana, CuA,. The names involved are discussed below and a new Curitiba, Parana, Brazil. species is described from Rondonia. 6 AUSTIN et ai: Hesperiidae of Rondonia - Entheus TROPICAL LEPIDOPTERA

Fig. 1. Entheus males, dorsal surface. Top row: left - E. eunyas, holotype; center - E. telemus, BRAZIL: Rondonia; 62 km S Ariquemes, Fazenda Rancho Grande. 12 Jun 1993; right - E. priassus pralina, BRAZIL: Rondonia; Linha 10, 5 km S Cacaulandia, 4 May 1995. Middle row: left - E. aureanota, holotype; right - E. priassus pralina, BRAZIL: Esp. Santo; Concei9ao da Barra, 4 Sep 1969. Bottom row: left - E. gentius, neotype; center - E. bombus, holotype; right - E. aureolus, holotype Fig. 2. Entheus males, ventral surface. Same specimens as in Fig. 1. J Vol. 8 No. 1 1997 AUSTIN et al.: Hesperiidae of Rondonia - Entheus 1

Entheus eumelus (Cramer) divergent; gnathos typical for genus; valva quadrate cephalad; costa with Papilw eumelus Cramer, [1777] short and blunt process; caudal end of harpe narrow, sharply bent Entheus eumelus; Evans, 1952 inward; penis short, stout, caudal end not prominently curved dorsad; 8 Entheus mina Williams & Bell. 1931 (part.; male only, female is of E. gentius) spike-like cornuti (n = 2); a male from Maues, Amazonas also has 8 The female of E. eumelus is similar to the male, and, in fact, comuti (3 are larger and 5 are of medium size). Cramer's (1777) figure is of a female. Williams and Bell (1931) FEMALE: forewing length = 17.1-17.2mm (n = 2); apex more rounded proposed Entheus mina for a putative pair from Dutch Guiana and produced than on male; dorsum dark brown; forewing with white, (Surinam). The male was said to have "a broad costal fold"; the hyaline macules as on male; base of discal cell with broad orange streak holotype was examined by Mielke and has no costal fold (no extending two-thirds distance to discal cell macule; vague orange scaling Entheus has a costal fold) as subsequently noted for a male from at mid costa anterior to discal cell macule; hindwing with large, more or Surinam (de Jong, 1983). This name is synonymous with E. eu- less rectangular white or pale yellowish patch extending from vein Rs to anal margin where broadest, this more than half the width of the wing melus. The genitalia figure by Williams and Bell (1931) for E. leaving the extreme base, costal margin, and outer margin (4mm in mina is a poor representation. The female of Williams and Bell's width) brown. (1931) "pair" of E. mina was considered Entheus gentius by Ventral forewing similar to dorsum; pale orange macules anterior to Evans (1952); this may be correct (see below under that species). and nearly the width of discal cell macule; anal margin whitish to Evans' (1952) records were for French and British Guiana. proximal edge of macule in CuA2-2A; base of discal cell pale orange; Mielke's records are for Venezuela (Bolivar: Santa Elena) and hindwing with white patch more extensive anteriorly, extending nearly Brazil (Para: Obidos, Aug). The "type" of Papilio eumelus is to the base and more distad towards apex. from Surinam as is the holotype of Entheus mina. Head black with white macules above, white beneath and behind eyes; palpi white ventrally, black on dorsal and inner surface; antennae black, Entheus ninyas H. H. Druce, rev. stat. narrowly whitish distad beneath, nudum with 18 (n = 1) or 23 (n = 1) Entheus ninyas H. H. Druce, 1912 segments; thorax black above with a few white scales in collar, scattered Entheus eumelus ninyas'. Evans, 1952 (part) pale orange and pale gray scales laterad, white beneath; legs pale brown, outer surface white, mid and hind tibiae with spurs as on male but Entheus eunyas Austin, Mielke & Steinhauser, n. sp. shorter; abdomen black above, nearly entirely overscaled with white (Fig. 1-5, 12) except caudad, white beneath. Entheus ninyas H. H. Druce, 1912 Genitalia: central lamella postvaginalis broad on straight caudal edge, Entheus eumelus ninyas: Moss, 1949 (biology, hostplant); Evans, 1952 (part) narrowing rapidly and fused with trough leading to caudal end of distinct Description.- MALE: forewing length = 17.2mm (16.7-18.0, n = 4); antrum; laterally lamella postvaginalis with broad but short plates, not forewing apex pointed, not produced, termen evenly curved; hindwing overlapping or extending caudad of caudal end of central portion of slightly angled at vein M3; dorsal forewing with bright orange (Spectrum lamella postvaginalis, caudal end broad with several short, robust teeth Orange to Chrome Orange) triangular-shaped patch basad from basal 1/3 of about equal size, covered cephalad with membranous flap, this with of anterior edge of discal cell to 3/4 distance to tornus on anal margin; feather-like striations; ductus bursae broad before narrowing to corpus remainder of forewing dark brown with pale yellow hyaline macules as bursae. follows: continuous discal band from vein Sc through discal cell to just Types.- Holotype 3 with the following labels: white, printed - BRASIL: posterior of CuA2 where narrow and triangular-shaped; parallelogram- Rondonia / 62 km S Ariquemes / linea C-20, 7 km E / B-65, Fazenda / shaped macule in M3-CuA,, about 1/2 distance from discal band to termen; continuous (but divided by dark veins), curved series of 6 Rancho Grande / 16 June 1993 / leg. G. T. Austin / (at paper lures / 1200-1230); white, printed and handprinted - Genitalia Vial / GTA - subapical-submarginal macules from R2-R3 to M2-M3, 4 subapical macules elongate, submarginal macules nearly square, terminating about 3708; red, printed - HOLOTYPE / Entheus eunyas I Austin, Mielke & Steinhauser. 1/2 distance between macule in M3-CuA, and termen; costa overscaled with orange basad: vague pale orange costal macule anterior to discal Paratypes (all BRAZIL: Rondonia, leg. G. T. Austin unless noted): same cell macule; often orange scaling posterior to end of discal band. location as holotype, 22 Oct 1992, leg. J. P. Brock, associated with Hindwing bright orange; costal margin very pale brown along basal 1/2, Eciton burchelli (Hymenoptera: Formicidae), 0825-0855h (1 3); 15 Nov darker brown distad and caudad to vein Rs; outer margin narrowly dark 1992 (1 9); linea C-20, 10 km E B-65, 3 km E Fazenda Rancho Grande, brown; fringes dark brown on both wings. lot 18, 15 Jun 1993, at paper lures, 1100-1130h (1 <5); 15 Jun 1993, at Ventral surface nearly identical to dorsum; orange paler; forewing paper lures, 1230-1300h (1

Fig. 5-11. Male genitalia of Entheus, lateral view of tegumen, gnathos, uncus, and associated structures; dorsal and ventral views of uncus, gnathos, and anterior tegumen; interior view of right valva; left lateral view of penis; and dorsal view of penis (latter without vesica and comutus). (5) E. eunyas, paratype, BRAZIL: Ron- donia; 3 km E Fazenda Rancho Grande, lot 18, 15 Jun 1993 (GTA #3707); (6) E. telemus, BRAZIL: Rondonia; Fazenda Rancho Grande, 16 Apr 1992 (GTA #2388); (7) £. priassus pralina, BRAZIL: Esp. Santo; Conceicao da Barra, 4 Sep 1969 (GTA #2387), dorsal and ventral views of uncus, etc. not shown; (8) E. aureanota, paratype, Fazenda Rancho Grande, 13 Nov 1991 (GTA #2386); (9) E. gentius, neotype (GTA #5766); (10) E. bombus, holotype (GTA #2512); (11) E. aureolus, paratype, 3 km E Fazenda Rancho Grande, lot 18, 22 Sep 1992 (GTA #2513). and M2-M3, quadrate, often square, offset distad from and usually not or slightly overlapping central lamella postvaginalis and extending contiguous with subapical macules, subapical-submarginal macular series caudad to about its caudal edge, caudal end narrow with distinct inner crossed by dark veins; no or vague white streak in costal cell anterior to tooth and several very small teeth distad, broadening gradually cephalad discal cell macule; base of discal cell with bright orange streak extending with several small teeth on inner edge caudad, covered with membran- 2/3 to 3/4 distance to discal cell macule; hindwing with relatively eous flap cephalad, this with some feather-like striations; ductus bursae narrow, more or less rectangular creamy white patch extending from vein bulbous, narrowing before elongate corpus bursae. Rs to anal margin, this about half the width of wing leaving extreme Distribution and phenology.- Evans (1952) gave the distribution of E. base, costal margin, and outer margin broadly brown. telemus as Ecuador, Peru, and Brazil. This may require modification Ventral forewing similar to dorsum; white macules anterior to and since we have seen phenotypes which superficially resemble E. telemus narrower than discal cell macule in Sc-R, and R,-R2; anal margin white but have different genitalia. In central Rondonia, this species is relatively to distal edge of posterior macule in CuA2-2A; base of discal cell common in the forest with records for all months. whitish; hindwing with white patch more extensive, extending to the Discussion.- We treat E. telemus as a full species, separate from E. base and nearly to costal margin. priassus as in the original description and in Emmel and Austin (1990). Head black with white macules above, white line beneath antennae, Among other characters, it differs from E. p. priassus and E. p. pralina white beneath and behind eyes; palpi very pale yellow, black on dorsal by the conjoined apical and discal bands on the male forewing, the and inner surfaces; antennae black, narrowly very pale yellow beneath extensive white (extending to the anal margin) on the female hindwing, distad and under club, nudum of 19 (n = 1), 20 (n = 1), or 21 (n = 3) and the male genitalia. Our recent finding of E. p. pralina near segments; thorax black above with a few white scales in collar, scattered Cacaulandia further strengthens the specific status of E. telemus. white and ochreous scales otherwise, white beneath; legs pale brown, outer surface white, mid and hind tibiae with spurs as on male but Entheus priassus pralina Evans shorter; abdomen black above, sparsely overscaled with white, white (Fig. 1-4, 7, 14) beneath. Entheus priassus pralina Evans, 1952 Genitalia: central portion of lamella postvaginalis narrow, about as Description.- MALE; forewing length = 18.0mm (n = 1); forewing apex long as broad, caudal margin with prominent central indentation, fused produced, termen evenly curved; hindwing subquadrate, slightly angled with trough which narrows cephalad, leading to ostium bursae at caudal at vein M3; dorsum nearly black; forewing with narrow, bright orange end of distinct antrum; lateral lamella postvaginalis of broad plates not (Chrome Orange) discal band from vein Sc through distal half of discal Vol. 8 No. 1 1997 AUSTIN et al: Hesperiidae of Rondonia - Entheus 11 cell nearly to outer margin in CuA2-2A, extending to but not below vein edge nearly perpendicular to anal margin in cell CuA2-2A; band mostly 2A, proximal edge not perpendicular to anal margin in cell CuA2-2A, opaque, hyaline at distal end of discal cell and anteriorly in cell CuA,- band opaque: large, triangular-shaped, much paler orange, hyaline CuA2 where adjoined to large, triangular-shaped, orange, hyaline macule macule in M3-CuA,, this extending from base of cell where joined with in M3-CuA,, this extending from base of cell nearly to outer margin on discal band to 2/3 distance to outer margin on vein CuA,, angled away vein CuA,, angled away from margin anteriorly; series of four elongate, from margin anteriorly; series of four elongate, conjoined, pale orange, conjoined, orange, hyaline, subapical macules from R2-R3 to R5-M,, hyaline, subapical macules from R2-R3 to R5-M,, separate from discal nearly touching (but clearly separate from) or touching (5 of 15 band anteriorly; these continuous and conjoined with two similar specimens) discal band anteriorly; these followed by two, conjoined, submarginal macules in cells M,-Mi and M2-M3 which terminate about roughly quadrate, otherwise similar, offset slightly distad (but continuous 1/2 distance from distal corner of macule in M,-CuA, and outer margin; with subapical macules), submarginal macules in cells M,-M2 and M2- subapical-submarginal macules crossed by narrowly bright orange veins; M3, latter smaller than former and placed 1/2 distance between anterior hindwing unmarked; fringes dark brown on both wings. distal corner of macule in M3-CuA, and termen; hindwing unmarked: Ventral surface nearly identical to dorsum; forewing narrowly pale fringes dark brown on both wings. gray-brown along anal margin. Ventral surface nearly identical to dorsum; forewing narrowly pale Head black with pale yellow-orange macules above, pale yellow- gray-brown along anal margin. orange beneath and behind eyes; palpi black, broadly pale yellow-orange Head black with pale yellow-orange macules above, pale yellow- beneath; antennae black, narrowly yellow distad beneath, nudum of 19 orange beneath eyes; palpi black, broadly pale yellow-orange beneath; segments; thorax black above with a few orange scales in collar, black antennae black, narrowly yellow distad beneath, nudum of 20 (n = 5), 21 beneath with numerous yellow-orange scales in forecoxae; legs dark (n = 5), or 22 (n = 3) segments; thorax black above with a few yellow- brown, first two pairs with yellow-orange on sides, hind legs with orange scales in collar, black beneath with pale yellow-orange forecoxae; yellow-orange on posterior 1/3 of outer face of tibia, mid tibia with one legs dark brown, first two pairs with yellow-orange at sides proximad, pair of spurs, outer somewhat shorter than inner, hind tibia short, with hind legs with red-brown on posterior half of outer face of tibia, mid one pair of spurs, inner about 2X length of outer, long brown tibial hair tibia with one pair of spurs, inner slightly longer than outer, hind tibia tuft fitting into groove on back of flattened and elongate first tarsal short, with one pair of spurs, inner longer than outer, long tibial hair tuft segment; abdomen black with vague ochreous caudad beneath. ochreous proximad, brown distad, fitting into groove on back of flattened Genitalia: tegumen long with narrow, proximate, sharply pointed pair and elongate first tarsal segment; abdomen black above, black with pale of caudal processes, their caudal end nearly reaching caudal end of yellow-orange caudad beneath. uncus; uncus relatively long, moderately broad and curved ventrad at Genitalia: very similar to most other species of the genus (e.g., middle in lateral view, arms widely divergent in ventral view; gnathos Steinhauser, 1989); tegumen relatively long with narrow, proximate, typical for genus, moderately broad in lateral view, narrower than caudal sharply pointed pair of caudal processes, their caudal end extending end of uncus in ventral view; costa of valva with very long caudally nearly to caudal end of uncus; uncus relatively long, caudally broad in directed and terminally hooked process extending as far as cephalad end lateral view, arms broadly divergent in ventral view; gnathos typical for of harpe; caudal end of harpe relatively narrow, finely dentate, curved genus, narrower than caudal end of uncus in ventral view; costa of valva inward and twisted at ventro-caudal corner; penis somewhat bulbous with very long caudally directed and terminally hooked process cephalad, narrowing somewhat caudad where sharply upturned with long extending as far as cephalad end of harpe; caudal end of harpe relatively and pointed lower lip; 12 spike-like cornuti (4-5 large, based on 4 narrow, curved mesad and twisted at ventro-caudal corner; penis bulbous specimens from various locales). cephalad, narrowing caudad where sharply upturned with long and FEMALE: unknown at Rondonia study site (see discussion below). pointed lower lip; 13-15 spike-like cornuti, 4 or 5 are much larger than Distribution and phenology.- Evans (1952) reported the others. distribution of this taxon as Brazil (Espirito Santo, Rio, San FEMALE: forewing length = 20.5mm (20.1-21.1, n = 7); apex more Salvador, Amazons). One male from central Rondonia was taken rounded and produced than on male; dorsum dark brown; forewing with on 4 May 1995. white, hyaline macules as follows: discal cell, rhomboidal, anterior edge Discussion.- A single male which we identify as this taxon is longer than posterior, distal and proximal edges excavate; M3-CuA,, parallelogram, near termen; CuA,-CuA2, rhomboidal, posterior edge known from the Cacaulandia area of central Rondonia. Its longer than anterior, midway between macules in discal cell and M3- superficial characters are very similar to specimens from else- CuA,; anterior CuA2-2A, roughly rhomboidal, anterior edge longer than where but it is smaller (a male from Espirito Santo, Brazil with posterior, much smaller than macule in CuA,-CuA2, outer edges of two FW length = 19.9mm) and the genitalia are virtually identical. No in line; posterior CuA2-2A, triangular, offset proximad from macule in females have been seen from Rondonia. These should be similar anterior portion of same cell; four subapical, elongate, posteriormost to female E. telemus but (based on material from Espirito Santo) offset slightly distad; two submarginal in cells M,-M2 and M2-M3, have the subapical/submarginal and median series of macules on square, offset distad from but contiguous with (or slightly separated the forewing continuous, more extensive orange in the forewing from) subapical macules, subapical-submarginal macular series crossed discal cell, and a smaller white area on the dorsal hindwing, this by dark veins; vague to prominent white streak in costal cell anterior to not reaching the anal margin. discal cell macule; base of discal cell with bright orange streak extending 2/3 or more distance to discal cell macule; hindwing with large, more or Entheus aureanota Austin, Mielke & Steinhauser, n. sp. less rectangular white patch extending from vein Rs to anal margin, this (Fig. 1-4, 8, 15) more than half the width of the wing leaving the extreme base, costal Description.- MALE: forewing length = 19.8mm (17.3-21.2, n = 10); margin, and outer margin brown. forewing apex pointed, not produced, termen evenly curved; hindwing Ventral forewing similar to dorsum; white macules anterior to and quadrate, angled at vein M3; dorsum very dark brown, nearly black; nearly the width of discal cell macule in Sc-R, and R,-R2; anal margin forewing with broad, bright orange (Spectrum Orange, distinctly paler white to distal edge of posterior macule in CuA2-2A; base of discal cell where hyaline) discal band from vein Sc through distal half of discal cell whitish; hindwing with white patch extending to the base. to outer margin in CuA2-2A, not extending below vein 2A, proximal Head black with white macules above, white line beneath antennae, 12 AUSTIN et al.: Hesperiidae of Rondonia - Entheus TROPICAL LEPIDOPTERA

18 16

Fig. 12-18. Female genitalia of Entheus, ventral view. (12) E. eunyas, paratype, Fazenda Rancho Grande, 15 Nov 1992 (GTA #4398); (13) £. telemus, BRAZIL: Rondonia; B-65, 1 km N Cacaulandia, 20 Oct 1989 (GTA #2445); (14) E. priassus pralina, BRAZIL: Esp. Santo; Conceicao da Barra, 26 Sep 1968 (GTA #2390); (15) E. aureanota, paratype, BRAZIL: Rondonia; "Big Rock" trail, 3 km E Fazenda Rancho Grande, 29 Oct 1990 (GTA #3351); (16) E. gentius, SURINAM: Goliath Kreek, 24 Feb 1969 (GTA #5767); (17) E. bombus, paratype, Fazenda Rancho Grande, 19 Apr 1991 (GTA #3346); (18) E. aureolus, paratype, Fazenda Rancho Grande, 25 Sep 1992 (GTA #3353). white beneath and behind eyes; palpi white, black on dorsal and inner brown, outer surface white, mid and hind tibiae with spurs as on male surface; antennae black, narrowly whitish distad beneath becoming very but shorter; abdomen black above, nearly entirely overscaled with white, pale yellow beneath club, nudum of 20 (n = 1), 21 (n = 3), or 22 (n = white beneath. 1) segments; thorax black above with a few white scales in collar, Genitalia: central lamella postvaginalis somewhat broader than long, scattered white and ochreous scales otherwise, white beneath; legs pale caudal edge relatively straight with slight to prominent central indenta- 1997 AUSTIN et al: Hesperiidae of Rondonia - Entheus 13 tion, fused with trough which narrows cephalad, leading to ostium bursae form a smooth series with the submarginal macules which are not at caudal end of distinct antrum; lateral lamella postvaginalis of broad offset distad. Females of E. p. pralina (Fig. 3-4) have a much plates not or slightly overlapping central lamella postvaginalis and not smaller whitish area on the hindwing (this not reaching the anal extending much caudad of it, caudal end narrow, inner tooth usually margin), larger macules on the forewing (the posterior macule in twice (or more) as long as outer two or three teeth, clearly separate from CuA2-2A, however, is much smaller than on E. aureanota) with them, medial edge often with distinct teeth, cephalad portion abruptly broader than caudad, covered with membranous flap, this feather-like due the discal cell macule normally overlapping that in CuA,-CuA2, to parallel striations; ductus bursae bulbous, this narrowing somewhat and normally a broader and more intensely orange streak at the before elongate corpus bursae. base of the forewing discal cell. Types.- Holotype <5 with the following labels: white, printed BRASIL: Males of E. aureanota are also similar to E. telemus. The latter Rondonia / 62 km S Ariquemes / off B-65, vicinity / Fazenda Rancho / is as dark as E. p. pralina and also normally has a broad discal Grande, 180m / 2 November 1989 / leg. G.T. Austin; red, printed - band on specimens from Rondonia; at other locations (Manaus, HOLOTYPE / Entheus aureanota I Austin, Mielke & Steinhauser. Sao Gabriel, Belem, Maues, Januarete, "Guiana"), the discal band Paratypes (all BRAZIL: Rondonia, leg. G. T. Austin unless noted): same is narrower. The subapical and submarginal macules are con- location as holotype, 21 Jul 1994 (1 $); 4 Aug 1991, leg. G. Bongiolo joined with the discal band and macule in M3-CuA,, respectively, (1 9); 21 Nov 1991, leg. O. Mielke (1 9); 14 Nov 1991, leg. O. Mielke leaving a small, completely enclosed triangle of dark ground color (1 9); 8-19 Nov 1994, leg. O. Mielke (1 3); 22 Mar 1989 (1

Entheus gentius Species-Group flattened and elongate first tarsal segment, this with outer portion orange with black tip (1/3 its length) and very long, extending to distal end of Evans (1952) distinguished the male of E. gentius by its broad first tarsal segment, inner portion pale brown and of the same length; and largely opaque yellow markings and a presence of a rhomboi- abdomen yellow-orange beneath. dal macule in cell M3-CuAj. Females were characterized by a Genitalia: similar to E. telemus as described above; uncus thin in yellow to orange base to the ventral forewing discal cell, lateral view and nearly straight; gnathos narrow in ventral view; valva discontinuous discal and apical bands, the usual absence of a mid broad cephalad, costal process traces broadly rounded arc from origin, harpes relatively narrow; penis robust; 7 cornuti, one of which is very costal macule, a usually large lower macule in cell CuA2-2A, and long. long spines on the lateral plates of the lamella postvaginalis. FEMALE: forewing length = 19.6mm (n = 1); apex more produced than Evans (1952) noted that males were "very variable in respect on male; dorsum dark brown; forewing with white, hyaline macules as of size, tone of orange coloring and width border H" and "the follows: discal cell, rectangular, distal edge concave; M3-CuA,, form of the clasp very variable." He also stated that the uncus parallelogram, nearer termen than to discal cell macule; CuA,-CuA2, was asymmetrical (his figure does not indicate this; no Entheus rhomboidal, posterior edge longer than anterior, nearly contiguous with we have seen has an asymmetric uncus). De Jong (1983) noted discal cell macule, far from macule in M3-CuA,; anterior CuA2-2A, more variation among females in the placement of forewing macules or less square, anterior edge slightly longer than posterior, somewhat and configuration of the genitalia. It is unknown if they were smaller than macule in CuA,-CuA2, edges of two in line; posterior CuA,- looking at more than one species. Among our material from 2A, nearly opaque, more or less quadrate, smaller than and offset Rondonia, there are two distinct species and we questioned which, proximad from macule in anterior portion of same cell; four subapical, if either, represented Papilio gentius Cramer described from elongate, posteriormost slightly longer than others; two submarginal in cells M,-M2 and M2-M3, square, offset slightly distad from but contigu- Surinam. Since a type does not exist for this taxon, it was ous with subapical macules forming nearly continuous series, subapical- necessary to first determine its identity. Material from Surinam submarginal macular series crossed by dark veins; short (or none on was requested from R. de Jong at the Nationaal Natuurhistorisch another specimen), pale orange, mid costal macules anterior to discal cell Museum in Leiden. He sent four specimens, a male of which macule; base of discal cell with moderately broad yellow-orange streak appears to represent Cramer's (1777) concept. This (plus speci- extending 2/3 distance to discal cell macule; hindwing with large white mens examined by Mielke) is described and discussed before patch extending from vein Rs to anal margin where broadest, this more treatment of the Rondonia species. than half the width of the wing leaving the extreme base, costal margin, and outer margin brown, brown outer margin vaguely edged proximad Entheus gentius (Cramer) by pale yellow-orange. (Fig. 1-4, 9, 16) Ventral forewing similar to dorsum; white macules anterior to and Papilio gentius Cramer, [1777] same width as discal cell macule; anal margin whitish to under distal Entheus gentius; Moss, 1949 (hostplant); Williams and Bell, 1931 (misident. see edge of posterior macule in CuA2-2A; base of discal cell pale yellow- below); Evans, 1952 (part?) orange extending to costa; hindwing with white patch more extensive, Entheus mina Williams and Bell, 1931 (part, female only); Evans, 1952 extending to base and nearly to costal margin. Description.- MALE: forewing length = 18.5mm (neotype); forewing Head black with white macules above, white line beneath antennae, apex pointed, not produced, termen and costa slightly convex; hindwing white beneath and behind eyes; palpi white, black on dorsal and inner subquadrate, slightly angled at vein M3; dorsum marked with black and surfaces; antennae black proximad, whitish beneath distad and beneath opaque bright yellow-orange (Orange Yellow); forewing with yellow- club, nudum of 26 segments (n = 1); thorax brown above with a few- orange basal third; broad yellow-orange discal band from vein Sc white scales in collar and scattered yellow scales, white beneath; legs through distal end of discal cell approaching outer margin in CuA2-2A, pale brown, outer surface white, tibial spurs shorter than on male; edges nearly parallel, distal edge somewhat irregular anteriorly, portion abdomen white, slight yellow tinge on sides. in discal cell hyaline (this latter appears to be a unique character for E. Genitalia: central lamella postvaginalis more or less rectangular, gentius); yellow-orange parallelogram-shaped hyaline macule in M3- broader than long with prominent central indentation on caudal edge, CuA,, equidistant between termen and discal band; slightly curved series fused cephalad with long trough leading to ostium bursae at caudal end of four elongate, conjoined, yellow-orange subapical hyaline macules of distinct antrum; lateral lamella postvaginalis with very broad plates from vein R2 to M,, clearly separate from discal band anteriorly; these nearly touching centrally, not overlapping caudal end of central lamella continuous with similar hyaline submarginal macules in cells M,-M2 and postvaginalis, caudal end with numerous teeth, these moderately long and M2-M3, latter smaller than former and placed halfway between anterior extending to or beyond caudal end of central lamella postvaginalis distal corner of macule in M3-CuA, and termen, entire series of macules laterad and shorter centrally, covered cephalad by membranous flap with decreasing in size posteriad; hindwing yellow-orange with very broad prominent striations; ductus bursae bulbous; corpus bursae not seen. (nearly 1/3 wing width) black outer margin; between 1A and anal margin Type.- As mentioned the type of this species is lost. A male (without and partially distad between 2A and 3A more narrowly black, grayer antennae) at the Nationaal Natuurhistorisch Museum, Leiden, is here proximad; fringes black on both wings. designated as the neotype of Papilio gentius Cramer, [1777]. It is labeled Ventral surface nearly identical to dorsum; yellower; black margin of as follows: white, handprinted (difficult to decipher) - E. H. Jonkers / hindwing much broader at tornus and along anal margin to wing base. bosgebied / Rama aan / Sur. rivier / / 18-3-1974; white, printed Head yellow-orange, white to pale yellow beneath and behind eyes; and handprinted - Genitalia Vial / GTA-5766; red, printed - NEOTYPE palpi black above and on inner surface, otherwise pale yellow; antennae / Papilio gentius I Cramer, [1777] / designated by / Austin, Mielke & missing on neotype, black with club and adjacent shaft pale yellow on Steinhauser /1995. The associated female (described above) is also from other specimens, nudum of 24 (n = 4) or 25 (n = 1) segments; thorax Surinam (Goliath Kreek, February). and abdomen bright yellow-orange above, thorax yellow-orange beneath Distribution and phenology.- Evans (1952) indicated a broad including forecoxae; legs yellow-orange, mid tibia missing on neotype (yellow on other specimens), hind tibiae with one pair of spurs, inner distribution for E. gentius in the northern two-thirds of South more than 2X length of outer and hair tuft fitting into groove on back of America. Further evaluation of the variation mentioned by Evans Vol. 8 No. 1 1997 AUSTIN et al.\e of Rondonia - Entheus 15

(1952) and de Jong (1983) and the now known existence of at segments; thorax bright yellow above, collar yellow, yellow beneath with least two additional species in this group will require refinement pale yellow forecoxae; legs yellow, mid tibia with one pair of spurs, of this. At present, E. gentius is known from Surinam (the type outer shorter than inner, hind tibia with one pair of similar short spurs locality) and Mielke has seen additional material from northern and hair tuft fitting into groove on back of flattened and elongate first portions of Brazil (Amazonas: Vila Nova, Manaus; Para: Obidos) tarsal segment, this with inner portion pale yellow, extending about 7/8 length of first tarsal segment, outer portion yellow-orange and relatively and Colombia (Amazonas: Leticia). These have collection dates long, extending distad 1/2 to 2/3 length of first tarsal segment; abdomen from April to September. yellow above, pale yellow beneath. Discussion.- Of the two broadly yellow species of Entheus seen Genitalia'. similar to E. gentius', differ as follows: caudal processes of from Surinam (E. gentius and the second new species described tegumen slightly more robust; uncus more robust, broader and more below), the present phenotype matches most closely the somewhat angular in lateral view, arms shorter in ventral view; gnathos with sides stylized representation of Papilio gentius in Cramer (1777). This parallel in ventral view; costa of valva with inner arm angled more figure (Plate CLXX, Fig. C) shows a more yellow (rather than abruptly ventrad and straighter (this broadly curved on E. gentius), yellow-orange) with a very broad black margin on the thinner caudad; caudal end of harpe broader; penis somewhat more hindwing and an entirely yellow head and body (see also robust; 5 (n = 2), 6 (n = 2) or 8 (n = 1) spike-like cornuti, 2-3 of these discussions under following two species). Williams and Bell larger than others. FEMALE: forewing length = 20.2mm (paratype); apex more rounded (1931) and Evans (1952) illustrated male genitalia determined as and produced than on male; dorsum dark brown; forewing with white, E. gentius. The latter are too schematic to determine species. The hyaline macules as follows: discal cell, nearly square, anterior edge very Williams and Bell (1931) figure is somewhat better and appears slightly longer than posterior; M3-CuA,, parallelogram, near termen; not to be of E. gentius based upon the short processes of the CuA,-CuA2, rhomboidal, posterior edge much longer than anterior, tegumen and the thin penis (see below). The female described midway between macules in discal cell and M3-CuA,; anterior CuA2-2A, above from Surinam has genitalia different from both other roughly square, anterior edge slightly longer than posterior, somewhat known species of the E. gentius group and thus is associated with smaller than macule in CuA,-CuA2, outer edges of two in line; posterior E. gentius by default. The female illustrated by Williams and Bell CuA2-2A, parallelogram, smaller than and offset proximad from macule (1931) as E. mina is virtually identical superficially with the in anterior portion of same cell; four subapical, elongate, posteriormost female assigned to E. gentius here and Evans (1952) was correct offset slightly distad; two submarginal macules in cells M,-M, and M2- M3, square, offset distad from but contiguous with subapical macules, in synonymizing that specimen with E. gentius. A second female subapical-submarginal macular series crossed by dark veins (not seen from Surinam is of similar size (19.5mm) and is marked continuous as on E. gentius); short streak in Sc-R, and longer streak in similarly but with the macules in CuA,-CuA2 and anterior CuA2- costal cell anterior to discal cell macule pale orange; base of discal cell 2A offset distad by their width from the discal cell macule. This with broad bright orange streak extending 3/4 distance to discal cell also appears to be E. gentius but the genitalia could not be macule; hindwing with large whitish (may be slight yellow tinge) patch examined since the abdomen was missing. extending from vein Rs to anal margin where broadest, this more than half the width of the wing leaving the extreme base, costal margin, and Entheus bombus Austin, Mielke & Steinhauser, n. sp. outer margin brown. (Fig. 1-4, 10, 17) Ventral forewing similar to dorsum; white macules anterior to and Description.- MALE: forewing length = 18.0mm (17.2-18.9, n = 5); nearly the width of discal cell macule; anal margin white to under distal forewing apex pointed, not produced, costa relatively straight, termen edge of posterior macule in CuA2-2A where pale orange; base of discal evenly curved; hindwing quadrate, angled at vein M3; dorsum marked cell whitish anteriorly, pale orange posteriorly; hindwing with white with black and opaque bright yellow with slight orange cast (near patch extending to the base and nearly to costal margin, distal edge pale Orange Yellow); forewing with yellow basal third; broad yellow discal orange towards anal margin. band from vein Sc through distal end of discal cell approaching outer Head black with white macules above, white line beneath antennae, margin in upper half of CuA2-2A, edges nearly parallel, distal edge white beneath and behind eyes; palpi white, black on dorsal and inner somewhat irregular anteriorly; yellow parallelogram-shaped macule in surfaces; antennae black proximad (distal portions missing); thorax gray- M3-CuA,, midway between discal band and termen; curved series of four green above with a few white scales in collar, white beneath; legs pale elongate, conjoined, yellow subapical hyaline macules from R2-R3 to R3- brown, outer surface white, tibial spurs as on male; abdomen black M,, clearly separate from discal band anteriorly; these continuous with above with much white overscaling, white laterally and beneath. similar, but opaque, submarginal macules in cells M,-M2 and M2-M3, Genitalia: caudal edge of central lamella postvaginalis slightly curved latter smaller than former and placed halfway between anterior distal with relatively broad "v"-shaped central indentation, fused cephalad with corner of macule in M3-CuA, and termen; hindwing yellow with broad long trough which narrows cephalad leading to ostium bursae at caudal (nearly 1/4 wing width) black outer margin and narrower black anal end of distinct antrum; lateral lamella postvaginalis with broad plates margin from half way between 2A and 3A to anal margin; extreme base slightly overlapping central lamella postvaginalis, caudal end with teeth of wing with black smudge on some specimens, this extending narrowly of moderate and about equal length (two on right, three on left), these about 1/4 distance to apex along costa; fringes black on both wings. barely extending caudad of central lamella postvaginalis, medial edge Ventral surface nearly identical to dorsum; forewing narrowly paler with several prominent teeth, flap over anterior portion with well yellow along anal margin proximad; hindwing with no basal black; anal developed feather-like striations; ductus bursae bulbous, this narrowing margin normally with considerable yellow mixed with black at 3A and somewhat before bulbous corpus bursae. especially marginad or entirely black. Types.- Holotype 3 with the following labels: white, printed: BRASIL: Head bright yellow with scattered black scales, white beneath eyes; Rondonia / 65 km S Ariquemes / linea C-20, 10 km E / B-65, 3 km E palpi with protruding segment black, otherwise pale yellow distad Fazenda / Rancho Grande, lot 18 / 23 September 1992 / leg. G. T. grading to white proximad, upper and inner surfaces black; antennae Austin; white, printed and handprinted - Genitalia Vial / GTA - 2512; black above, yellow beneath, nudum of 25 (n = 2) or 26 (n = 2) white, printed and handprinted - Genitalia Vial / SRS - 4373 / File No.; 16 AUSTIN et al: Hesperiidae of Rondonia - Entheus TROPICAL LEPIDOPTERA red, printed - HOLOTYPE / Entheus bombus I Austin, Mielke & Description.- MALE: forewing length = 19.3mm (18.6-20.2, n = 10); Steinhauser. forewing apex pointed, not produced, termen evenly curved; hindwing Paratypes: same location and collector as holotype, 18 Jul 1994 (1 3); subquadrate, slightly angled at vein M3; dorsum marked with black and 24 Sep 1992 (1

We thank Drs. L. D. and J. Y. Miller (Allyn Museum of Entomology, ^TlM^T™ HeSpCliidae' *""' A//>'" MuSmm (S^' Sarasota, Florida) for their assistance in this study and for reviewing the ' .' manuscnpt. J. D. Turner (Huntsville, Alabama) made helpful comments 1989' Taxonomic notes and descriptions of new taxa in the Neo- on an early draft. R. de Jong, at the Nationaal Natuurrnstorisch Museum, ^'aTota 127°! 7^' ^ ^^ Leiden, Netherlands, graciously loaned material from Surinam. T. C. w.,,. Emmel (Univ. of Florida, Gainesville, Florida) continues to provide /^f^lT '..^ *" *" „ "*" encouragement during our studies. G. Lamas and an anonymous rev,ewer 193L Hesperiidae of the Forbes expedition to Dutch and British provided thoughtful comments. Acknowledgments are due to G. ^mana (Lepidoptera). Trans. Amer. Ent. Soc. (Philadelphia), Bongiolo, J. Brock, O. Gomes, and S. Kohler for assistance in the field ,.,,„,.,..,,. ... ,, . , .. ,., -n, 1934. Studies in American Hespenoidea, Paper IV (Lepidoptera). On and to the Harald Schmitz family for providing excellent hospitality. The Conselho Nacional de Desenvolvimento Cientifico e Tecnologico kindly *e W™™ <"£ *e male. gfn *a" fme sPecies' Trms' issued the authorization permits from the Ministerio da Ciencia e Amer' EnL Soc' (Philadelphia), 60:265-280. Tecnologia for our studies in Rondonia in collaboration with EMBRAPA/CPAC.