Rienk de JONG

National Museum of Natural History, Leiden

REVISION OF THE ORIENTAL DE NICÉVILLE (: HESPERIIDAE: PYRGINAE)

Jong, R. de, 2006. Revision of the Oriental genus Odontoptilum de Nicéville (Lepidoptera: Hesperiidae: Pyrginae). – Tijdschrift voor Entomologie 149: 145-159, figs. 1-54. [issn 0040- 7496]. Published 1 December 2006. The Oriental genus Odontoptilum, currently conceived as consisting of three and five additional subspecies, is revised based on examination of all types and a study of recent material, particularly from the . As a consequence the number of species-group taxa is raised to ten, arranged in two genera (one new, Semperium for the species Pterygospidea leptogramma Hewitson, 1868), six species and four additional subspecies. Two species, O. corria sp. n. and O. abbreviata sp. n., and one subspecies, O. pygela tawita ssp. n., are described for the first time, and one species, O. helias (Felder & Felder), is raised from subspecies rank. O. angulata sinka Evans proved to be a junior synonym of O. helias helisa Semper. Four species are restricted to Wallacea. The genera are remarkable because of the strongly asymmetrical genitalia and the occurrence of three types of secondary sexual characters, one of which is described for the first time. R. de Jong, National Museum of Natural History, P.O. Box 9517, 2300 RA Leiden, The Netherlands. E-mail: [email protected] Key words. – Hesperiidae; Odontoptilum; Semperium; secondary sexual characters; new taxa; ; distribution; Oriental region; Wallacea.

The genus Odontoptilum as currently understood in Odontoptilum and that its isolated taxonomic posi- (Evans 1949, Maruyama 1991, Eliot 1992, Bridges tion is best reflected by placement in a new genus. 1994) contains three species: O. angulata (Felder, All taxa are briefly described below to facilitate 1862), from and NW to the identification, and relevant characters are illustrated. Philippines, and the Lesser Sunda Islands, For synonymy other than proposed here, see Evans O. pygela (Hewitson, 1868) from Burma and Thai- (1949). All holotypes relating to species-group names land to Java and , and just penetrating the in Odontoptilum are in the Natural History Museum, Philippines, and O. leptogramma (Hewitson, 1868), London (bmnh), except one that is in Forschungsin- endemic to the Philippines. The species are easy to stitut und Naturmuseum Senckenberg, Frankfurt- separate on the basis of external characters. In the am-Main (smf), and the types of the newly described male genitalia O. leptogramma stands apart because of taxa, which are in the National Museum of Natural the elaborate valvae, while the valvae of O. pygela and History ‘Naturalis’ (formerly Rijksmuseum van O. angulata are of a simple and much more slender Natuurlijke History), Leiden (rmnh), and in the col- build. While O. leptogramma and O. pygela exhibit lection of C.G. Treadaway (cgt, donated to smf). minor variation, O. angulata (as conceived by Evans 1949) is highly variable, seasonally as well as geo- Odontoptilum in relation to other genera graphically, and in wing markings as well as in geni- talia. This variation, displayed by a richer material The Oriental genus Odontoptilum de Nicéville, than Evans (1949) had at his disposal, prompted re- 1890, as currently conceived, belongs to a group examination of the types and, subsequently, a revision of Afro-Oriental genera described as the Tagiades of the concept of O. angulata. Further, examination group by Evans (1949), a grouping that, on the ba- of the genitalia and secondary sexual characters led sis of molecular data, may well be monophyletic to the conclusion that O. leptogramma was misplaced (A. Warren pers. comm.). Within the Tagiades group,

145 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

1 2 3

Figs. 1-3. Venation and outline of hindwing of Semperium leptogramma (1), Odontoptilum pygela (2), and O. helias (3). The other Odontoptilum species have an outline similar to O. helias.

Odontoptilum shares three characters with the genera species are described and figured for the first time, the Ctenoptilum de Nicéville, 1890 (Oriental), position of O. leptogramma (Hewitson) is reconsid- Wallengren, 1857 (Afro-Oriental; the congenerity ered, O. pygela (Hewitson) is briefly described (and of the Oriental and African species needs re-exami- a new subspecies is proposed), and a taxonomic revi- nation), Netrobalane Mabille, 1904, Leucochitonea sion of the O. angulata (Felder) complex is presented, Wallengren, 1857, and Abantis Hopffer, 1855 (the including the description of two new taxa. latter three genera restricted to the Afrotropics). Evans (1949) coined the name Caprona subgroup for these Delimitation of Odontoptilum genera. The characters which more or less separate them from the other genera of the Tagiades group are: The type species of Odontoptilum is O. angulata antennal apiculus blunt, subcostal vein of forewing (Felder, 1862). The genus is currently conceived as ending well before end of cell, and the male genitalia containing three species, O. angulata, O. pygela, and asymmetrical to some extent, in the valvae as well as O. leptogramma. According to Evans (1949), they in the uncus and gnathos, with the aedeagus generally share the following characters that separate them emerging from the right. Asymmetrical male genitalia from the genus Caprona: (1) hair pencil on fore coxae are not unique among Hesperiidae (see for example, of male short (long in Caprona and Leucochitonea); Burns 1964), but they are among the Tagiades group (2) tegulae of male more or less prolonged; (3) tor- (with exception of the genus Abraximorpha Elwes & nal cilia of hindwing elongate. However, the tegu- Edwards, 1897, which in other characters appears to lae of the male are also longer than in the female in be more closely related to other genera). Whether Caprona agama Moore, 1857 (Oriental region) and these characters point to a closer relationship be- in C. pillaana Wallengren, 1857 (Afrotropical re- tween, and monophyly of, the six genera remains to gion), and possibly in more species which have not be studied. There are other remarkable characters that been checked. Elongate tornal cilia on the hindwing seem to point in the same direction. One character can also be found, within the Caprona subgroup, in is the irregular outline of the wings (figs. 1-3) (but the Afrotropical genera Netrobalane and Abantis (at regular in Leucochitonea and Abantis), not unique in least in some of the species), and also in other genera the Tagiades group, but rarely found to this extent. of the Tagiades group like the Oriental genera Darpa Caprona, Leucochitonea and Odontoptilum share a Moore, 1865, and Seseria Matsumura, 1920. Hence peculiar and unique sexual character in the male, a the morphological basis for considering the three dense hair tuft on the fore coxae. Ctenoptilum has a species congeneric (i.e. more closely related to each recumbent hair pencil on the hind tibiae in the male, other than to any other species) is very weak indeed, a character widespread among Pyrginae, but absent in viz. the length of the hair pencil on the fore coxae many genera and, although of diagnostic value, ap- of the male. However, I failed to find the hair pencil parently of little help in establishing relationships. on the fore coxae of O. leptogramma. Moreover, while In this study a novel secondary sexual charac- the male genitalia of O. angulata and O. pygela are of ter is described and figured, female genitalia of the similar build, those of O. leptogramma are abundantly

146 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

different (valvae complicated, uncus with large dorsal Semperium gen. n. horn, no gnathos), negating the reason for considering Type species (and sole species included): Pterygospidea O. leptogramma more closely related to O. angulata leptogramma Hewitson, 1868. and O. pygela than to any other species. In addition, O. leptogramma proved to have a unique secondary Description. − See S. leptogramma. See also above, sexual character. The isolated taxonomic position of under Delimitation of Odontoptilum. O. leptogramma in the Caprona subgroup is best re- Etymology. − The genus is named after Georg flected by placing the species in a genus of its own Semper (1837-1909), the Nestor of Philippine lepi- with, as yet, unclear relationships. Therefore, a new dopterology. genus is described below. Semperium leptogramma (Hewitson) Secondary sexual characters in Odontoptilum Pterygospidea leptogramma Hewitson, 1868: 53. Holotype In Hesperiidae, secondary sexual characters (sscs) , in bmnh; type locality: Philippines. are characters, other than genitalic characters, that presumably play a role in sexual behaviour by scent Material examined. − Holotype , Philippines emission. They are widespread in Hesperiidae, par- (bmnh); 41, 6, Cebu, Leyte, Luzon, Mindanao, ticularly in the male, but scent distributing organs Mindoro, Samar, Sibuyan (bmnh, cgt, rmnh). in females have been described as well (Burns 1964, de Jong 1975). A common feature among females External characters (figs. 37, 40). − Length in the Tagiades group, and found in all species of of forewing, male 18.9-20.0 mm, female 20.2- Odontoptilum and the new genus Semperium, is the 20.6 mm. Upperside forewing with a fuzzy whitish- presence of thickly packed, wavy, hairlike scales at violet line from mid-dorsum to mid-costa, a narrow, the tip of the abdomen, known as ‘anal wool’. It is broken white line crossing space 1bc and space 2 (ap- reminiscent of a ssc but as described by Igarashi & parently the median spots in these spaces) and fol- Fukuda (1997) for several Tagiades species and by lowing the discoidal veins, a fuzzy violet zigzag line Bascombe et al. (1999) for O. angulata, its function is between the latter line and the outer margin, a short not in sexual behaviour: the hairs are used to conceal bluish line near the base from dorsum to cubital vein the laid eggs. In males, sscs are most frequently found and two bluish lines in basal part from radial vein on the wings, but they can be found on other body to costa; in basal half veins outlined in violet-white; parts as well, particularly on the legs. In Odontoptilum some violet scaling in spaces 1bc and 2 between the two peculiar male sscs have been described. One is two cross-lines; tiny hyaline spots in spaces 3-9. Un- a black or brown tuft of hair-like scales originating derside forewing, brown with strongly reduced mark- from the fore coxae and lying against the thorax. It ings. Upperside hindwing with central third white. also occurs in Caprona and Leucochitonea, where it Underside hindwing white colour even more exten- is much longer and more conspicuous, but it is ab- sive. Outline of hindwing slightly wavy, very shallowly sent in the new genus Semperium. The other male indented between the veins, strongest between veins ssc in Odontoptilum is a prolongation of the tegulae 4 and 6, and 7 and 8. (misnamed scapulae by Evans 1949) into a thick hair Secondary sexual characters. − On the hindtibia pencil (figs. 4, 5) fitting into a groove on the upper- close to its junction with the femur, the male with a side of the hindwing along the basal third of space 1c thin erectile hair pencil associated with a groove on (figs. 5, 6). This character is only found in some sub- the underside of the hindwing in space 1b along the species of O. angulata (Felder) as conceived by Evans thickened basal half of vein 1a, flanked by shining (1949). A third ssc, unique among Hesperiidae, white narrow scales (fig. 7). is found only in the new genus Semperium. It con- Male genitalia (figs. 8, 14-16) − Tegumen and un- sists of an apparently erectile metatibial hair pencil cus askew to the left. Uncus slightly flattened dorso- (found in many Pyrginae) associated with a groove on ventrally, with weakly serrated sides. Large middor- the underside of the hindwing (fig. 7), as described sal horn on uncus, curving to the right. Gnathos below. Usually, when a metatibial hair tuft is associ- absent. Valvae strongly asymmetrical, large, curved, ated with another structure, it is with a metathoracic more or less encapsulating the rest of the genitalia. pouch. The only exception known so far is in the ge- Right valva with more or less regular cucullus, distal nus Celaenorrhinus, where the metatibial hair tuft is edge slightly serrate; costa consisting of a regular part associated with an abdominal pouch opening through ending roundedly over cucullus and a large, spined, a slit along the side of the 2nd abdominal sternite proximal process curving over the regular part and (de Jong 1982). ending acutely; sclerotization of costa extending from proximal part to mid-valva. Aedeagus emerging

147 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

Fig. 4. Odontoptilum helias helisa, male; lateral view of hairtuft on tegula.

Fig. 5. Odontoptilum helias helias, male; dorsal view of hair tufts on tegulae (yellow arrows) and closed groove on hind- wing (green arrow).

almost centrally, but, relative to the uncus, from the markings: good diagnostic characters at the species right. level, but not at the genus level. Female genitalia (fig. 28). − Weakly asymmetri- Secondary sexual characters. − The only character cal. More simply built than in Odontoptilum. Lateral found in all species, but in the male only, is the short parts of abdominal segment 7 well sclerotized and hair pencil on the fore coxae (often difficult to see as with large elongate, not elevated spiracles; tergal part it is covered by the legs and regular scales). reduced; sternal part a regular, squarish anterior plate Male genitalia. − Highly asymmetrical. In dorsal with rounded corners imperceptibly passing into ster- view uncus far to the left and aedeagus to the right, num 8; latter forming a regular, squarish posterior emerging from under a large folded gnathos. Valvae plate, smaller than the anterior plate, densely spinu- asymmetrical, simply built, elongate, generally taper- lose on the inner side; tergum 8 a strongly domed ing towards a cucullus that is extended, particularly structure bearing the anal wool and, in ventral view, in the right valva, and sometimes shaped like a swan’s rising behind the papillae anales. Ductus and corpus neck. bursae simple, no ornamentation, together relatively Female genitalia. − Irregular and asymmetrical. short, less than twice as long as distance from proxi- Abdominal segment 7 modified to form a more or mal edge of sternum 7 to tip of abdomen. less sclerotized anterior lamella (i.e. anterior to the os- Distribution (fig. 33). − The species is confined to tium) which is more or less folded, and passing imper- the Philippines. It has been found on the following ceptibly into the folds of the 8th sternum; lateral parts islands: Cebu, Leyte, Luzon, Mindanao, Mindoro, of segment 7 well sclerotized and with conspicuous, Samar, Sibuyan. elevated spiracles; tergal part of segment 7 reduced to Variation. − Very slight and not geographical. a narrow strip. Sternum 8 with very irregular, sclero- Life history. − No information available. tized folds, densely spinulose on the inner side; ter- gum 8 well developed, more or less domed, bearing the anal wool. Bursa simple, elongate, without orna- Odontoptilum de Nicéville mentation; ductus and corpus bursae together 2.5 to Odontoptilum de Nicéville, 1890: 217. Type species by origi- 3.5 times as long as distance from proximal edge of nal designation: Achlyodes sura Moore, [1866], which is sternum 7 to tip of abdomen. currently identified as a junior synonym of Pterygospidea Distribution. − Throughout the Oriental region. angulata Felder, 1862.

Odontoptilum pygela (Hewitson) External characters. − Although the species can eas- ily be separated from other Oriental Hesperiidae by Pterygospidea pygela Hewitson, 1868: 53. Holotype , in the outline of the hindwing (figs. 2, 3), there are actu- bmnh; type locality: Malacca. ally two types of wing shape (pygela and the angulata complex) and not a single one defining the genus. External characters (figs. 38-39, 41-44). − Length The same holds for characters like colour and wing of forewing, male 16.8-19.3 mm, female 18.7-

148 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

Fig. 7. Semperium leptogramma, male; groove on underside hindwing (gr), flanked by elongate shining scales; veins 1a and 1b are indicated.

band from just above tornus to vein 4 and continued narrowly to vein 6, flanked by a grey area up to the irregular white line and by a grey band towards outer margin, but narrowly white again along outer margin. Underside hindwing largely white from dorsum up to radius/vein6. Outline of hindwing (fig. 2) strongly indented between veins 4 and 7 and less strongly so between veins 7 and 8, resulting in a pronounced tooth at the end of vein 7 and an even more pro- nounced one at the end of vein 4; distance from wing base to end of vein 4 considerably greater than from wing base to end of vein 7; between tornus and vein 4 slightly concave. Secondary sexual characters. − No other character Fig. 6. Odontoptilum corria, male; upperside hindwing show- than the hair tuft on the fore coxae. ing opened groove with black specialized scales at bottom. Male genitalia (figs. 9, 17-18). − Uncus long, slen- der, flattened laterally, strongly curving from the left side to the right. Right valva elongate triangular with 21.1 mm. Upperside forewing with a white, slightly strongly extended, slightly upcurving cucullus ending irregular line from mid-vein 1 through cell, sometimes with a serrated tip. Left valva more trapeziform, cuc- extending to mid-costa, and another slightly irregular ullus straight, pointing distally, serrated in distal part, white line representing the median spots in spaces 1bc shorter than cucullus of right valva. and 2 (the latter sometimes hyaline), often continued Female genitalia (fig. 29). − Ostium surrounded into a narrow hyaline median spot in space 3, nar- by very irregular folds that defy a clear description rowly white at wing base from costa to dorsum, nar- and passing imperceptibly into the each other, and row hyaline spots in spaces 8 and 9, sometimes a very from sternum 7 into sternum 8 but in ventral view, tiny one in space 6; veins concolourous with the dark no lamella directly in front of and behind the ostium. brown, little variegated ground colour; slight whitish Ductus and corpus bursae together about three times violet superscaling along dorsum and sometimes ex- as long as distance from proximal edge of sternum tending a bit onto wing. Underside forewing rather 7 to tip of abdomen. similar to upperside, markings less pronounced. Up- Distribution (fig. 32). − Sundaland, north to perside hindwing narrowly white at extreme base, a S Burma and ; it penetrates the Philippines straight pronounced line from mid-vein1b to shortly in Palawan, Sanga Sanga and Tawitawi. before mid-vein 8, and a very irregular, narrow white Variation. − Evans (1949) recognized two subspe- line distal to the straight line and sharply curving out- cies; here a third is described. side along vein 4 to halfway space 4-5 a broad white Life history. − No information available.

149 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

8 9 ae gn ae rv un

un rv

te te

lv lv

10 11

12 13

Figs. 8-13. Male genitalia of Semperium and Odontoptilum, dorsal view. – 8, S. leptogramma (Luzon, Los Baños, rmnh ins.303342). 9, O. pygela (N , East coast, Laut Tador, rmnh ins.303321). 10, O. angulata (Java, Gunung Sewu, rmnh ins.303216). 11, O. helias (N Celebes, nr Menado, rmnh ins.303258). 12, O. corria (paratype, C Luzon, Dolores, cgt). 13, O. abbreviata (paratype, ‘Felder, 460’, rmnh ins.303247). ae = aedeagus; gn = gnathos; lv = left valva; rv = right valva; te = tegumen; un = uncus.

Odontoptilum pygela pygela (Hewitson) species, except Java and the Tawitawi group in the Material examined. − Holotype , Malacca Sulu Archipelago. (bmnh); 167, 6, Burma, Thailand, Malay Penin- sula, Sumatra, Nias, Banka, Billiton, Borneo, Palawan Odontoptilum pygela javanica Fruhstorfer (bmnh, cgt, rmnh). Odontoptilum pygela javanica Fruhstrofer, 1909: 173. Holo- External characters (figs. 38, 41). − Upperside type , in bmnh; type locality: W Java. forewing spot in space 3 usually present, but often tiny. Upperside hindwing, space between the central Material examined. − Holotype , W Java (bmnh); white lines in cell and spaces 4-5 brown. Underside 42, 7, Java (bmnh, rmnh). hindwing, space 6 largely dark with two white lines. Distribution. − The distribution area of the External characters (figs. 39, 42). − Upperside

150 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

external characters of the only taxon of which the holotype was not in the bmnh (Evans very rarely stud- ied types that were not in London) led Evans astray. uh On the basis of overall similarity in wing markings and wing shape, the taxa united by Evans (1949) un- der O. angulata clearly form a group well separated from the other two species of the genus. This is less apparent in the male genitalia, where there is a general similarity between O. angulata (at least in the nomi- au notypical form) and O. pygela. Within O. angulata sensu Evans, however, two types of ssc and three types of male genitalia occur. Since there is geograph- ic overlap in the types of male genitalia, we can only conclude that they are specifically distinct. Although the arrangement given here is not fully satisfactory, it is the best for now. Fig. 14. Semperium leptogramma, left lateral view of tegumen and uncus; au = apex of uncus, uh = uncus horn. The various forms can be arranged in two groups on the basis of presence or absence of a peculiar ssc: tegulae in the male prolonged into a hair pencil forewing, no spot in space 3. Upperside hindwing, (figs. 4, 5) that fits into a groove on the upperside of space between the central white lines in cell and spac- the hindwing in the basal half of space 1c (figs. 5, 6); es 4-5 white. Underside hindwing as in ssp. pygela. the groove is filled with specialized scales. This ssc Distribution. − Restricted to Java. occurs in all forms flying in Wallacea (Sulawesi and the Philippines), and is absent in the forms occurring in continental Asia and Sundaland, and the Lesser Odontoptilum pygela tawita ssp. n. Sunda Islands as far as Flores. Material examined. − Holotype, , philip- pines, Tawitawi group, Sanga Sanga Is., Boloboc, Odontoptilum angulata (Felder) 17.vi.1992. Paratypes: 1, 2, same locality, but 8.ii.1989, 22.iv.1989, and 9.vii.1990; 1, Tawitawi Pterygospidea angulata Felder, 1862: 488. Holotype  in Is., Tarawakan, 26.vi.1992; 2, 4, Tawitawi Is., bmnh; type locality: Hong Kong. 19-22.ii.2006. All types in cgt, except one  and one  paratype in rmnh. External characters (figs. 45, 48). − Length of forewing, male 16.9-20.8 mm, female 20.2- External characters (figs. 43-44). − As ssp. pygela, 22.3 mm. Upperside forewing strongly variegated but on underside hindwing white colour more exten- with paler and darker brown colours, hyaline spots sive, reaching vein 7, largely obscuring dark spots in in spaces 2, 3 (may be absent), 7 and 8, rarely a tiny space 6; dark basal spot in space 7 smaller than in dot in space 6. Underside forewing paler than upper- ssp. pygela. side. Upperside hindwing pale brown, darker brown Distribution. − Only known from the Tawitawi in basal third followed by an almost straight pale line group of islands in the southwestern part of the Sulu from vein 1b (well beyond its middle) to costa; a very Archipelago. irregular thin and inconspicuous pale line more dis- tally; dark brown at apex; pale brown to white line from just above tornus to near end of vein 4; veins The Odontoptilum angulata complex 1b to 4 paler outlined from irregular line to termen; External characters. − Upperside forewing without pale brown to white line along termen; fringes pale white cross-lines. Outline of hindwing (fig. 3) indent- brown to white. Underside hindwing mainly white, ed between veins 7 and 8, slightly concave in space darkened to various degrees mainly at apex and along 4-5, almost straight from vein 4 to tornus; distance termen from tornus to vein 4, conspicuous round from wing base to end of vein 7 greater than from black spot at base of space 7. wing base to end of vein 4. Secondary sexual characters. − No other character than the hair tuft on the fore coxae. While O. leptogramma and O. pygela exhibit minor Male genitalia. − Highly asymmetrical; in dorsal variation, Evans (1949) distinguished five subspe- view uncus curving from the left side to about the cies for O. angulata. Incorrect interpretation of the middorsal line, expanding to a rounded head. Right

151 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

15 16

17 18

19 20

21 22

23 24

25 26

Figs. 15-26. Inside of left (odd numbers) and right (even numbers) valvae of Semperium and Odontoptilum. 15, 16, S. leptogramma (Luzon, Los Baños, rmnh ins.303342). 17, 18, O. pygela (N Sumatra, East coast, Laut Tador, rmnh ins.303321). 19, 20, O. angulata (Java, Gunung Sewu, rmnh ins.303216). 21, 22, O. helias (N Celebes, nr Menado, rmnh ins.303258). 23, 24, O. corria (paratype, C Luzon, Dolores, cgt). 25, 26, O. abbreviata (paratype, ‘Felder, 460’, rmnh ins.303247).

152 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

and left valvae subequal in length, but unequal in , and throughout Sundaland to Bali. shape. Right valva subtriangular, cucullus elongate, Life history. − Early stages have been described more or less curving up, spined distally. Left valva, and illustrated by Bascombe et al. (1999), Igarashi see subspecies. & Fukuda (1997), and Piepers & Snellen (1910). Female genitalia. − See under subspecies. Hostplants include members of four families, viz., Distribution (fig. 31). − Continental Asia, from Bombacaceae (Ceiba, Eriodendron), Malvaceae Sri Lanka, and NW Himalayas to Hong Kong (Hibiscus, Urena), Sapindaceae (Allophylus) and and W Malaysia, and through Sundaland to the Tiliaceae (Grewia, Microcos). Lesser Sunda Islands as far east as Flores. Variation. − In addition to seasonal variation (see Odontoptilum angulata hyperides (Doherty) Evans 1949) two subspecies can be distinguished. Life history. − See under the nominotypical form. Abaratha hyperides Doherty, 1891:195. Holotype  in Identification. − The only other species of the ge- bmnh; type locality: Sumbawa. nus with which O. angulata flies is O. pygela. The dif- ferences in wing pattern and outline of hindwing are Material examined. − Holotype , ‘Sambawa’ so pronounced that confusion is highly unlikely. (bmnh); 2, Lombok, 3 Sumbawa (all bmnh).

External characters. − Length of forewing, male Odontoptilum angulata angulata (Felder) 19.0-20.7 mm; Evans (1949) indicates that this form Material examined. − 196, 54, Sri Lanka, is smaller than ssp. angulata, but this does not agree India, Nepal, Bhutan, Sikkim, Burma, Thailand, with my measurements. Wings more rounded. Upper- Indo-, Hainan, Hong Kong, W Malaysia, side hyaline spots on forewing reduced; darker, more Sumatra, Borneo, Java (bmnh, rmnh). uniformly coloured, but pale brown along termen of hindwing replaced by white (and fringes white) leav- External characters. − Length of forewing, male ing a conspicuous row of white-edged brown spots. 16.9-20.8 mm, female 20.2-22.3 mm. Upperside Male genitalia. − Right valva slender, costa hardly strongly variegated with various shades of brown; developed, cucullus hardly curving up, but slightly outer third of forewing warm brown; spot in space curving to the outside, slightly broadening towards 2 well-developed, more or less crescentic, spot in spined tip. Left valva without the dorsal hump space 3 small or absent, spots in spaces 7 and 8 well- of the nominate subspecies, costa imperceptibly pass- developed. Upperside hindwing paler or darker brown ing into the almost straight cucullus, which is club- along termen, not whitened, fringes pale brown. shaped, flattened, slightly curved and serrate in distal Male genitalia (figs. 10, 19-20). − Right valva dor- part. sally humped just beyond middle, cucullus long, slen- Female genitalia. − No females known. der, curving up, and down again, as a swan’s neck, Distribution. − Lombok, Sumbawa and Flores. spined distally. Left valva subrectangular, costa sharp- Discussion. − In the original description, Do- ly bending down to triangular cucullus with a ventral herty mentioned that ‘Another species, more like line curving up to the almost horizontal dorsal line, A. angulatus, was found in Sumba, but no specimens and then cucullus extending into a strongly serrated, survived’. This could underline the closer relation- pointed and down-curving projection. ship of ssp. hyperides with ssp. angulata than with Female genitalia (fig. 30). − Asymmetrical; ostium any other taxon in the complex. It is in line with the displaced to the left. Abdominal segment 7 laterally absence of the ssc consisting of the hair pencil at sclerotized with prominent, raised spiracles, sternally the tegulae associated with the groove on the hind- lightly sclerotized to membraneous; laterally the scle- wing. rotization passing imperceptibly into the folded and highly irregular posterior lamella (or rather plate) of Odontoptilum helias (Felder & Felder) the 8th segment, which is densely spinulose along the edge and on the innerside; central part of posterior Pterygospidea helias Felder & Felder, [1867]: 529. Holotype plate slantingly squarish, distal edge slightly concave, , in bmnh; type locality: Celebes. not indented, about half the width of sternum 7. Bursa copulatrix hardly more than a broadening of External characters (figs. 46-47, 49-50). − Up- the ductus bursae; ductus and corpus bursae together perside hindwing, pale line bordering basal dark area about 2.5 times as long as distance from proximal reaching vein 1b at or just before the middle of the edge of sternum 7 to tip of abdomen. vein. Distribution. − Continental Asia, from Sri Lanka, Secondary sexual characters. − In addition to the India and NW Himalayas to Hong Kong and W hair tuft on the fore coxae there are specialized grey

153 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

27 29

28 30

Figs. 27-30. Female genitalia of Semperium and Odontoptilum, ventral view. 27, O. helias (N Sulawesi, Menado, rmnh ins.303256). 28, S. leptogramma (Samar, rmnh ins.303344). 29, O. pygela (NE Sumatra, Tanjung Merawa, rmnh ins.303276). 30, O. angulata (Java, Depok, rmnh ins.303229).

scales at the bottom of a groove on the upperside squarish apex. hindwing. Female genitalia (fig. 27). − Asymmetrical; ostium Male genitalia (figs. 11, 21-22). − Uncus not as displaced to the left. Similar to O. angulata; sternum strongly asymmetrical as in O. angulata, displaced to 7 more strongly sclerotized, central part of posterior the left, curving down, but not sideways, broadening plate larger, squarish, about as wide as sternum 7, dis- to a rounded apex. Right and left valva very unequal tally indented, lateral folds of plate smaller than in in size and shape. Right valva slender, subtriangular, O. angulata. Ductus and corpus bursae together costa rather flat, cucullus narrowly extending to a about 4 times as long as distance from proximal edge down-curving, serrate, blunt apex. Left valva about of sternum 7 to tip of abdomen ⅔ length of right valva, subrectangular, costa bulg- Distribution (fig. 34). − Sulawesi and Philippines. ing dorsally in proximal half, cucullus in continuation Variation. − Two subspecies can be distinguished, of costa, straight, with strongly serrate, blunt or differing in size, wing markings and genitalia.

154 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

31 32

Figs. 31-32. Distribution areas of Odontoptilum angulata (31) and O. pygela (32).

(bmnh); 5, Cebu (3 cgt, 2 rmnh); 1, Duma- Odontoptilum helias helias (Felder & Felder) ran (cgt); 7 1, Luzon (5 cgt, 2 1 rmnh); Material examined. − Holotype , ‘Celebes’ 1, Marinduque (cgt); 1 1, Mindanao (cgt); (bmnh), 42, 2, Sulawesi (N, C and S) (bmnh, 7, Mindoro (cgt). rmnh). External characters (figs. 47, 50). − Considerably External characters (figs. 46, 49). − Length of smaller than ssp. helias, length of forewing, male forewing, male 21.4-23.9 mm, female 24.9 mm. 16.2-20.4 mm, female 21.2 mm. Upperside darker, Upperside strongly variegated, dark areas on forew- less strongly variegated, no pale blue or violet su- ing contrasting with the paler brown colours; hyaline perscaling, white colour along termen on upperside spots in spaces 2, 7 and 8 well developed, in space 3 hindwing less extensive, a more or less continuous small but usually present, sometimes one or two tiny grey band between white terminal line and the white hyaline spots in cell. Upperside hindwing strongly band from tornus to end of vein 4. Hyaline spots on whitened along termen, grey band between white forewing: in spaces 2, 7 and 8 always present, usually terminal line and white band from tornus to end of also in space 3, and one or two tiny spots in cell. vein 4 more or less divided into spots and becoming Male genitalia. − Cucullus of left valva longer, up obsolete from tornus to vein 4. In male, central part to 4 times as long as high, apex more rounded; ventral of hindwing with pale violet superscaling. edge curving upwards more abruptly. Male genitalia (fig. 21). − Cucullus of left valva Female genitalia. − See species description. The fe- relatively short, about twice as long as high. males listed above have been assigned to the present Female genitalia. − See species description. taxon by similarity to the males in external characters. Distribution. − Sulawesi. Since we are not yet sure about the correct identity of Identification. − The largest form in the genus, it the females listed under O. corria and O. abbreviata, cannot be confused with other Odontoptilum species we are not yet sure to what extent these taxa differ in since it is the only species in Sulawesi. female genitalia. Distribution. − Philippines: Basilan, Cebu, Luzon, Marinduque, Mindanao, Mindoro, Negros. Odontoptilum helias helisa Semper Identification. − Flies with O. corria in Luzon Odontoptilum helias helisa Semper, 1892: 311. Holotype , and Negros, and with O. abbreviata in Basilan and in smf; type locality: Mindanao. Mindanao. Externally it can be distinguished from Odontoptilum angulata subangulata Fruhstorfer, 1909: 172. Holotype , in bmnh; type locality: Basilan. both by the absence of pale violet superscaling on the Odontoptilum angulata sinka Evans, 1949: 159. Holotype upperside, the usual presence of a tiny hyaline spot in , in bmnh; type locality: Los Baños [Luzon]. the forewing cell of the male, and by the more con- tinuous and greyer band along termen on upperside hindwing; in the other species the grey band tends to Material examined. − Lectotype helisa, , Luzon be subdivided into spots, strongest in O. abbreviata (smf), holotype sinka, , Los Baños (bmnh), holo- where it is, moreover, narrower than in the other two type subangulata, , Basilan (bmnh); 1 , Basilan species.

155 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

33 34 35 36

Figs 33-36. Distribution areas of Semperium and Odontoptilum. 33, S. leptogramma. 34, O. helias. 35, O. corria. 36, O. abbreviata.

Discussion. − In the original description Semper Odontoptilum corria sp. n. said he had 18 specimens from Luzon, Samar, Bohol and Mindanao. He did not specify a holotype. Material examined. − Holotype , Philippines, Evans (1949), without having seen a specimen from Negros occ., Murcia, Canlandog, 800-900 m, Semper’s series, selected a male from Mindanao as 26.v.1995, leg. A. Buenafa (rmnh). Paratypes: 20, holotype, and he described the subspecies as: ‘In all Philippines, as follows: Luzon, 11 (10 cgt, helisa  the entire uph, except for the base and apex, 1 rmnh); Panay, 1 (cgt); Negros, 7 (6 cgt, 1 is covered with pale blue scaling, which extends, more rmnh); Polillo, 1 (bmnh). sparsely, to centre of upf.’ However, Semper did not Further material. − Negros, 1 (cgt); Panay, 1 mention the pale blue scaling at all. Moreover, in smf (rmnh); excluded from the type series. The correct a single specimen of ssp. helisa is kept, with a type assignment to this species must remain uncertain as label and originating from Luzon. It is in good con- long as also the correct assignment of the only avail- dition and does not have the pale blue scaling. We able possible female of O. abbreviata is uncertain. do not know what happened to the remainder of the More material is needed to make sure that possible series, and whether or not the other specimens also differences found in the genitalia are specific and not bore type labels. As it is possible that more than one due to individual variation. species was included in Semper’s series, the specimen from Luzon is designated lectotype here, even though External characters (figs. 51, 54). − Length of forew- the chance of any further specimens of the original ing, male 19.7-21.5 mm. Upperside forewing rather series turning up is extremely remote. uniformly dark brown with, at least in the male (no As a consequence of Evans’ incorrect concept of fresh females available), centrally a broad band of pale ssp. helisa, he described ssp. sinka, which on exami- violet superscaling from dorsum to the apical spots; nation proved to be indistinguishable from the true hyaline spots in spaces 2, 7 and 8, and usually also in ssp. helisa. space 3, present, no spots in cell, except in one female.

Figs. 37-54. Upper and undersides of Semperium and Odontoptilum. 37, 40, S. leptogramma, female (S Luzon, Mt Banahaw; cgt). 38, 41, O. pygela pygela, male (W Malaysia, Cameron Highlands; rmnh). 39, 42, O. pygela javanica, male (W Java, Palabuhan Ratu; rmnh). 43, 44, O. pygela tawita, male (holotype; Tawitawi group, Sanga Sanga Is.). 45, 48, O. angulata an- gulata, male (India; rmnh). 46, 49, O. helias helias, male (Sulawesi, Tondano; rmnh). 47, 50, O. helias helisa, male (lectotype; Luzon; smf). 51, 54, O. corria, male (holotype; Negros, Murcia; rmnh). 52, 53, O. abbreviata, male (holotype; Mindanao, S Cotabato Prov.; rmnh).

156 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

37 38 39

40 41 42

43 44 45

46 47 48

49 50 51

52 53 54

157 Downloaded from Brill.com09/27/2021 09:27:07AM via free access T  E,  149, 2006

Underside forewing uniformly brown with faint pale similar to O. abbreviata, but the hyaline spots on the violet superscaling. Upperside hindwing more vari- forewing are generally slightly better developed and egated than forewing, darker near base and apex, rest the terminal band of grey spots on the upperside of of wing with pale violet superscaling, along termen the hindwing is a bit broader; the two species are a grey band, more or less divided into spots by pale widely separated geographically. veins, in spaces 2 and 3 these spots about as high as wide; pale line bordering dark basal area reaching vein Odontoptilum abbreviata sp. n. 1b before the middle of the vein (i.e., slightly closer to the wing base). Underside hindwing largely white, Material examined. − Holotype , Philippines, brown along costa and at apex, in space 7, in addition Mindanao, South Cotaboto Prov., Salakot T’Boli, to the dark apex a dark brown rounded spot near the 2.vi.1985, leg. J. de los Reyes (rmnh). Paratypes: base and a squarish brown spot beyond the middle, 7, all Philippines, as follows: 2, Mindanao, South along termen a dark spot in space 1c and vague grey- Cotabato Prov., Koronadal, leg. R. Müller (1 rmnh, ish spots in other spaces. 1 cgt); 1, idem but Parker Mts (rmnh); 1, Secondary sexual characters. − In addition to the Mindanao, Agusan del Norte, 25.xi.1980 (rmnh); hair tuft on the fore coxae there is a groove on the up- 1, Mindanao, Mt Malindang, 10.vi.1987 (cgt); perside of the hindwing in space 1c with specialized 1, Basilan, Malcong, 20.xi.1932 (rmnh); 1, black scales at the bottom. Basilan (bmnh); 1 , ‘Felder, 460’ (rmnh). Male genitalia (figs. 12, 23, 24). − Strongly asym- Further material. − Mindanao, 1 (rmnh); ex- metrical in dorsal view, uncus displaced to the left, cluded from the type series for reasons mentioned slightly curving back to about the middorsal line, under O. corria. slender, with slightly expanded rounded tip and with a flattened lobe before the tip. Right and left valva External characters (figs. 52-53). − Length of subequal in length, but different in shape. Right valva forewing, male 19.2-21.2 mm, female unknown. reminiscent of O. angulata, but free part of cucullus Upperside forewing rather similar to O. corria, but longer than rest of valva, less strongly curved at apex; hyaline spots smaller, spot in space 2 a narrow line, in costa without bump, but with slight curvature. Left space 3 present in only 1 specimen; pale violet super- valva, cucullus in continuation of costa; under the scaling less pronounced. Underside forewing almost elegant down-curving free part of the cucullus (with plain brown. Upperside hindwing rather strong pale strongly toothed apex) the valva bulging out promi- violet superscaling, grey terminal band divided into nently. spots wider than high; pale line bordering dark basal Female genitalia. − See above, under Further mate- area reaching vein 1b before the middle of the vein rial. (i.e., slightly closer to the wing base). Underside hind- Distribution (fig. 35). − Philippines: Luzon, wing rather like O. corria. Negros (?), Panay, Polillo. Secondary sexual characters. − In addition to the Etymology. − The name refers to my wife, Corrie, hair tuft on the fore coxae there is a groove on the who has always been an enthusiastic field companion upperside hindwing in space 1c with specialized black and has always supported my endeavours wholeheart- scales at the bottom. edly. Male genitalia (figs. 13, 25-26). − Strongly asym- Discussion. − O. corria and O. abbreviata share sev- metrical in dorsal view, tegumen and uncus displaced eral external characters and, since they are allopatric, to the left, uncus broad, curving to the right side, they could well be considered to form a single spe- ending in a broadened apex. Right and left valvae cies. However, in the male genitalia they represent the unequal in shape and size. Right valva longer than extremes in the development of the valvae, O. corria left valva, elongate-triangular, distally narrowing to a having two long, slender and curving valvae, reminis- rather straight, strongly toothed, acutely ending cu- cent of O. angulata, while O. abbreviata has both val- cullus; costa bulging upwards at ⅓ from base. Left vae short and straight. This is the more remarkable, valva, proximal half subrectangular, but then costa since O. helias helisa, which overlaps both other spe- strongly bending down and dorsal line smoothly con- cies in distribution, occupies an intermediate position tinued into the relatively short cucullus (narrow part in the male genitalia. Based on present-day knowl- about 2.5 times as long as high) with large teeth on edge it cannot be decided which two of the three taxa the ventral, distal and inner sides. are more closely related. Hence a species recognition Female genitalia. − See above, under Further for each is advisable for the moment. material. Identification. − The species almost completely Distribution (fig. 36). − Philippines: Basilan, overlaps with O. helias helisa; for differences, see un- Mindanao. der latter. Apart from the male genitalia O. corria is Etymology. − The name refers to the shortened

158 Downloaded from Brill.com09/27/2021 09:27:07AM via free access de Jong: Revision of Odontoptilum

valvae compared to O. angulata. Felder, C., 1862. Verzeichniss der von den Naturforschern Identification. − For differences from O. helias der k.k. Fregatte ‘Novara’ gesammelten Macrolepidop- helisa, with which it overlaps, see the latter species. teren. − Verhandlungen zoologisch-botanischer Gesell- schaft Wien 12: 473-496. For differences from O. corria, see there. Felder, C. & Felder, R., [1867]. Reise der österreichi- schen Fregatte Novarra um die Erde (3). − Kaiserlich- Acknowledgements Königlichen Hof- und Staatsdrückerei, Wien: 379-536. Fruhstorfer, H., 1909. Neue Hesperiden. − Entomologische This study was only possible thanks to the gener- Zeitschrift 23: 171-174. ous help of C.G. Treadaway (Limbach, Germany), Hewitson, W.C., 1868. Descriptions of one hundred new species of Hesperiidae (2): 26-56. − Author, London. who made all Hesperiidae from his incomparably rich Igarashi, S. & Fukuda, H., 1997. The life histories of Asian collection of Philippine available for study Butterflies, vol. 1. − Tokai University Press, Tokyo, to the author. The help of P.R. Ackery (The Natural xix+549 pp. History Museum, London) and Dr R.I. Vane-Wright Jong, R. de, 1975. An��������������������������������������� abdominal scent organ in some female (formerly The Natural History Museum, London) Pyrginae (Lepidoptera, Hesperiidae). ����������������− Entomologische during numerous visits to the museum collections Berichten Amsterdam 35: 166-169. is gratefully acknowledged. Dr. W. Nässig (Sencken- Jong, R. de, 1982. ������������������������������Secondary sexual characters in Celaenorrhinus and the delimitation of the genus berg Museum und Forschungsinstitut, Frankfurt-am- (Lepidoptera, Hesperiidae). − Journal of Natural History Main) is thanked for the loan of the type of O. helias 16: 695-705. helisa Semper. Dr J.M. Burns and Dr M. Schilthuizen Jong, R. de, & Treadaway, C.G., 1993. The Hesperiidae are thanked for their constructive comments. (Lepidoptera) of the Philippines. − Zoologische Verhan- delingen 288:1-125. Maruyama, K., 1991. Butterflies of Borneo, vol. 2(2), References Hesperiidae. − Tobishima Corporation, Tokyo, 89+83 p. Nicéville, C.L.A. de, 1890. On new and little-known But- Bascombe, M.J., G. Johnston & F.S. Bascombe, 1999. The terflies from the Indian region, with description of three Butterflies of Hong Kong. −���������������������������� Academic Press, San Diego, new genera of Hesperidae. − Journal Bombay natural x+422 pp. History Society 5: 199-225. Bridges, C.A., 1994. Catalogue of the -group, Piepers, M.C., & Snellen, P.C.T., 1910. The Rhopalocera Genus-group and Species-group names of the Hesperiidae of Java. Hesperiidae. Martinus Nijhoff, The Hague, (Lepidoptera) of the World. −������������������������������������������������� Bridges, Urbana, xlvii+ xxvi+60 pp. 598 pp. Semper, G., 1892. Die Schmetterlinge der philippi- Burns, J.M., 1964. Evolution in butterflies of the nischen Inseln 1 (7): 271-380. − C.W. Kreidel’s Verlag, genus Erynnis. – University of California Publications in Wiesbaden. Entomology 37: 1-217. Treadaway, C.G., 1995. Checklist of the butterflies of the Doherty, W., 1891. Butterflies of Sumba and Smbawa [sic!], Philippine Islands (Lepidoptera: Rhopalocera). − Nach- with some Account of the Island of Sumba. − Journal asi- richten des entomologische Vereins Apollo, Suppl. 14: atic Society of Bengal (II)60:141-197. 7-118. Eliot, J.N., 1992. 4th revision of A.S. Corbet & H.M. Pend- lebury, The Butterflies of the Malay Peninsula. − Malayan Nature Society, Kuala Lumpur, xiv+595 pp. Evans, W.H., 1949. A catalogue of the Hesperiidae of Europe, Asia and in the British Museum (Natural History). − Trustees of the British Museum, Received: 30 August 2006 London, xix+502 pp. Accepted: 22 September 2006

159 Downloaded from Brill.com09/27/2021 09:27:07AM via free access