Revision of the Nyctemera clathratum complex (Lepidoptera: Arctiidae)

Revision of the Nyctemera clathratum complex (Lepidoptera: Arctiidae) Rob de Vos The Nyctemera clatrathum complex (Lepidoptera: Arctiidae, Arctiinae, Callimorphini) is revised. It consists of seven species, occurring on New Guinea and surrounding islands, of which three are new to science: Nyctemera giloloensis sp. n., N. oninica sp. n. and N. dauila sp. n. Two taxa are synonymized: Leptosoma absurdum Swinhoe, 1892 syn. n. of Leptosoma clathratum Snellen van Vollenhoven, 1863 and Deilemera pratti Bethune-Baker, 1904 syn. n. of Nyctemera mesolychna Meyrick, 1889. Lectotypes are designated for Deilemera pratti Bethune-Baker, 1904 and Deilemera absurdum latimargo Rothschild 1915. Adults and genitalia of all species are figured and distribution maps are presented. Rob de Vos, Zoölogisch Museum Amsterdam, dept. Entomology, Plantage Middenlaan 64, NL-1018 DH Amsterdam, The Netherlands. rvos@science.uva.nl. Introduction The genus Nyctemera Hübner, [1820] s.l. is a large heterogeneous taxon with many species in Southeast Asia. The few species in Africa that previously were considered to belong to Nyctemera recently have been revised and seem to belong to other genera (Dubatolov 2006). Apart from the taxa which have recently been shown to belong to Utetheisa Hübner, [1819] (De Vos 2007), Nyctemera s.l. can be divided into several subgenera and species groups. The taxonomical problems and mysteries are far from unravelled, although a start has recently been made (De Vos 1995a, 1995b, 1996, 1997a, 1997b, 2002, De Vos & Cerny 1999). In the present study the Nyctemera clathratum complex is revised. It is a group of seven species found on New Guinea and the surrounding islands. Three species are new to science and described below. Based on genitalic morphology these species clearly belong to the nominotypical subgenus Nyctemera s.s. The valves in the male genitalia all possess three processes. The N. clathratum complex is closely related to the baulus species group, which is shown by the general structure of the genitalia and the wing pattern. The N. clathratum complex is distinguished from the baulus complex by the more or less slender and short valva processes of almost equal length, while in the baulus complex the costal process and apical process are much larger and thicker (and curved towards each other, resembling the claws of a lobster) than a third and smaller process on the sacculus. Furthermore the banded pattern on the abdomens of the species of the clathratum complex is brown-white (the white only narrow), while in the baulus complex the species have black-yellow-white banded abdomens (the white bands much broader than in the clathratum complex). Externally, some species are so hard to distinguish that they easily could be considered conspecific, whereas other species are so different that they seem to belong to different species groups. However, study of the genitalia revealed that all seven species indeed belong to a single complex, because they all share diagnostic male and female genitalic characters. In most cases, they have complementary, not overlapping distributions. Tijdschrift voor Entomologie 150: 39 54, Figs 1 62. [ISSN 0040 7496]. http://www.nev.nl/tve 2007 Nederlandse Entomologische Vereniging. Published 1 June 2007.

40 Tijdschrift voor Entomologie, volume 150, 2007 Materials and methods Material For this study 249 specimens were examined in the following 12 collections: BMNH Natural History Museum (formerly British Museum for Natural History), London, UK BPBM Bernice P. Bishop Museum, Gressitt Center for Research, Honolulu, Hawaii, USA KSP Koleksie Serangga Papua (Private collection Henk van Mastrigt), Jayapura, Papua, Indonesia CKC Private collection Karel Cerny, Zirl, Austria CMWM Museum Thomas Witt (assigned to Zoologische Staatssammlung München), Munich, Germany ISNB Institute Royal des Sciences Naturelles de Belgique, Brussels, Belgium OXUM Hope Entomological Collections, University Museum, Oxford, UK RMNH Nationaal Natuurhistorisch Museum Naturalis, Leiden, The Netherlands SNSD WAU Staatliche Naturhistorische Sammlungen, Dresden, Germany Agricultural University, Wageningen, The Netherlands ZMAN Zoölogisch Museum, University of Amsterdam, The Netherlands ZMHB Museum für Naturkunde der Humboldt Universität, Berlin, Germany Methods Genitalia preparations were made according to the standards of the Natural History Museum in London (Robinson 1976). At least one male and female specimen for each species was dissected, from N. clathratum, leopoldi and mesolychna some more. After dissection the genitalia were macerated for 12 hours in cold KOH 10%, cleaned in ethanol 30% and stained with chlorazol black which was dissolved in absolute ethanol. The genitalia were temporarily stored in ethanol 70% to allow the study of the three dimensional structure. Finally the genitalia were mounted on glass slides in euparal. Most of the adults were photographed with a conventional Nikon F400 camera, with Sigma AF52 Macro-objective and Sunpack ring flashlight, using Fuji 100 ASA colour transparency film. The specimens of Nyctemera giloloensis and the holotype of N. leopoldi were photographed with a digital Nikon D50 camera with the same lens and flashlight. The genitalia were photographed through a Leica MZ16 binocular microscope with a fixed DFC 320 digital camera which is controlled by Leica Firecam 1.9.1 software on a Macintosh Power PC G4 with operating system 10.4.1. Wing length was measured from wing base to apex in males and females. Morphological terminology of the external structures (excluding the genitalia) mainly follows Scoble (1992) and Holloway et al. (2001). The terminology of the genitalia mainly follows Jordan (1939), Tuxen (1970) and Kôda (1987), see also Figs 1 4. Results Nyctemera Hübner sensu stricto As mentioned above, the group of species presently considered to form the genus Nyctemera, still is rather heterogeneous. Study of the genitalia of the approximately 65 Indo-Australian species show that at least six species groups (termed subgenera hereafter) can be recognized. The nominotypical subgenus Nyctemera, is in the male genitalia distinguished from all others by the rather short valvae with three processes, while the other subgenera have only one or two processes. Androconial scales and hairs can be found on and around the ciliated process of the sacculus (i.e. in Coleta Roepke, 1949 these scales are in a brush on the foreleg, in Arctata Roepke, 1949 as long hairs at the dorsum of the hindwing and shorter scales on the sacculus, and in Deilemera Hübner, [1820] as very long hairs in a modified fold on the the rounded dorsum at the underside of the hindwing). Furthermore it is recognized by its wing pattern with a clearly defined spool- or oval-shaped fascia, often crossed by darker veins, and longitudinal white stripes in the basal field of the forewing but this is also common feature in some other subgenera. The shape of the forewing is stretched triangular (in Coleta and Trypheromera Butler, 1881 broader triangular, in Arctata even longer stretched and in Deilemera almost ovalshaped in males). It is clear that the genus needs a thorough revision, not only on the species level but also on (sub)genus level. The complexity of the taxonomy of Nyctemera s.l. is emphasized by the recognition of species groups within the subgenera. One of these groups in the nominotypical subgenus Nyctemera is the Nyctemera clathratum complex. The Nyctemera clathratum complex Diagnosis Head and patagia are yellow with a black dot on each. Male antennae are bipectinate with long cilia, in females, these cilia are much shorter. The shaft is covered with black scales in both sexes. Ground

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 41 1 costal process 2 apical process process on sacculus ventrum 4 ostium pockets ductus bursae (sclerotized) ductus bursae (unsclerotized) signum ductus seminalis bursa copulatrix 3 coecum Figs 1-4. Important genital structures of the Nyctemera clathratum group. 1, Three processes of the male valva; 2, lateral view of the male uncus with keel, apex and ventrum; 3, aedeagus with coecum; 4, female genitalia structures. colour of thorax and tegulae is pale yellow or bonewhite. Prothorax, mesothorax and tegulae possess a longitudinal black stripe, whereas the metathorax has a large black spot. Abdomens in all species are grey-brown with a narrow white ring of scales for each segment. Forewing ground colour is brown except for the broader or narrower white fascia located subapically. Where wing veins cross the white fascia, they are more or less suffused with brown scales. In contrast, in the basal half of the wing, veins are covered with white scales that sometimes extend significantly into

42 Tijdschrift voor Entomologie, volume 150, 2007 5 6 7 8 9 10 11 12 Figs 5-12. Nyctemera clathratum group, adult habitus. 5, N. clathratum, male, Ambon (SNSD); 6, N. clathratum, female, holotype; 7, N. clathratum, male, holotype absurdum; 8, N. clathratum, male, Misool (BMNH); 9, N. giloloensis, male, Halmahera, paratype (CMWM); 10, N. giloloensis, female, Halmahera, paratype (CMWM); 11, N. leopoldi, male, holotype; 12, N. leopoldi, female, Arfak Mts., Papua (ZMAN).

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 43 13 14 15 16 17 18 Figs 13-18. Nyctemera clathratum group, adult habitus. 13, N. oninica, male, holotype; 14, N. oninica, female, Roon Island, Papua, paratype (ZMAN); 15, N. mesolychna, female, holotype; 16, N. mesolychna, male, lectotype pratti; 17, N. latimargo, male, lectotype; 18, N. dauila, male, holotype. adjacent areas (Figs 9 10, Fig. 14). The wingfold between the anal vein (A2+3) and cubital vein (Cu2) is white. A white longitudinal line occurs in the discal cell in some species (e.g., N. giloloensis sp.n., and N. oninica). The hindwings are white with contrasting brown scales on the veins. The width of the brown, apical hind margin band is specific as is the degree of the brown suffusion of the veins. Genitalia are typical for the subgenus Nyctemera Hübner, [1820]. In males, the valve possesses three processes (a strongly ciliated process on the sacculus, a less ciliated apical process formed by fusion of the harpe and ampulla (Kôda 1987) and a less ciliated costal process) (Fig. 1). The uncus is broad and dorso-ventrally flattened with a beak-shaped apex. The shape of the dorsal keel, when present, is very diagnostic (Fig. 2). Females have surrounding the ostium two small lateral pockets (Fig. 4). A funnel-shaped antrum is continued by a partially sclerotized and flattened ductus bursae. The ductus bursae curls upon itself when entering the bursa copulatrix. The shape and number of curls can be diagnostic. The bursa copulatrix is large, globular, and

44 Tijdschrift voor Entomologie, volume 150, 2007 19 20 21 22 Figs 19-22. Male genitalia of Nyctemera clathratum (prep. RV1180). 19, outside of right valve; 20, dorsal view of uncus; 21, lateral view of uncus; 22, aedeagus. possesses one long spade-shaped signum, except for giloloensis, whose signum is divided into two boatshaped parts. Checklist Nyctemera Hübner, [1820]: 178 Type species Phalaena lacticinia Cramer, 1777 Subgenus Nyctemera s.s. Nyctemera clathratum complex clathratum (Snellen van Vollenhoven, 1863) absurdum (Swinhoe, 1892) syn. n. giloloensis sp. n. leopoldi Tams, 1935 oninica sp. n. mesolychna Meyrick, 1889 pratti (Bethune-Baker, 1904) syn. n. latimargo (Rothschild, 1915) dauila sp. n. Nyctemera clathratum (Snellen van Vollenhoven) Figs 5 8, 19 22, 47 48, 61 Leptosoma clathratum Snellen van Vollenhoven, 1863: 48 Holotype /, Indonesia: Forsten, Amboina; TYPE Sn.v.Voll.; Museum Leiden, Nyctemera clathratum Sn.v.Voll. /, Det: RvE [= R. van Eecke] (RMNH) [examined] Leptosoma clathratum; Butler 1880: 673; Pagenstecher 1888: 113. Pitasila clathrata: Kirby 1892: 424. Nyctemera clathratum: Pagenstecher 1901: 137; Bryk 1937: 58. Deilemera (Tripheromera) clathrata: Swinhoe 1903: 70. Nyctemera clathrata: Seitz 1915: 269; Eecke 1927: 222 (in part). Nyctemera clathrata clathrata: Roepke 1949: 66 (in part). Leptosoma absurdum Swinhoe 1892: 143 Holotype? Indonesia, Salawatti, Wallace, (OXUM) syn. n. [examined] Nyctemera absurdum: Pagenstecher 1901: 135; Bryk 1937: 47. Deilemera (Tripheromera) absurda: Swinhoe 1903: 79. Deilemera absurdum: Rothschild 1915a: 78. Nyctemera absurda: Seitz 1915: 272. Deilemera absurdum latimargo: Rothschild 1915b: 215 (nec Rothschild 1915a). Nyctemera absurdum latimargo: Bryk 1937: 48 (in part). Material examined. 18 specimens in addition to cited types. Indonesia: 1?, Ambon [no data], coll. Staudinger & Bang-Haas (SNSD); 1?, Ambon, x.1923, C.J. Brooks; 1/, Ambon [without further details]; 1?, Ambon, x.1907, Pratt; 1?, Ambon, ii.1892, W. Doherty; 1?, Ceram [Seram], Amahai, 13.xii.1929, miss Longfield; 3?, S. Ceram [Seram], Amahai, ix.1926 [no collector]; 1?, Ron [Roon], Geelvink Bay [no data], W. Doherty; 1/, Saparua [no data], Fruhstorfer; 1?, Waigeu [Waigeo, no data], Waterstradt; 1?, Waigeu [Waigeo], 23 26, J.J. Joicey coll.; 2?, 1/, Waigeu [Waigeo], Camp Nok, 2500 ft, iv.1938, v.1938, L.E. Cheesman; 1?, Misool [no data], Tauern (all BMNH). Diagnostic characters. Length of forewing 20 21 mm. Forewing colour brown. Basal half of wing with rather broad white veins, white wingfold and longitudinal line in cell narrow. Fascia broad, crossed by broad brown veins. Hindwing with rather narrow brown hind margin and with brown veins gradually fading to white or very narrowly brown towards base, except for the anal vein which is completely brown. Male genitalia (Figs 19 22). Uncus without keel (Figs 20 21), apex long and curved at an angle of 45 with ventral edge (Fig. 21), ventrum straight. Valve with costal process and process on the sacculus nearly equal in length and shorter than the apical process (Fig. 19). Aedeagus long, slender and curved, coecum (Figs 3 & 22) long and curved downwards.

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 45 23 24 The only known specimen from Roon has the suffusion of the veins in the forewing fascia even broader and the dark veins in the hindwing are not fading or narrowing. Nyctemera giloloensis sp. n. Fig. 9 10, 23 26, 49 50, 61 Nyctemera clathrata: Eecke 1927: 222 [partly misidentification] Nyctemera clathrata clathrata: Roepke 1949: 66 [partly misidentification] 25 26 Figs 23-26. Male genitalia of Nyctemera giloloensis (prep. RV1149). 23, outside of right valve; 24, dorsal view of uncus; 25, lateral view of uncus; 26, aedeagus. Female genitalia (Figs 47 48). Small pockets inside ostium wide and shallow. Antrum broad, sclerotized part of ductus bursae long with 120 curl. Ductus seminalis arises from the ductus bursae at one third distance from the bursa copulatrix. Signum (Fig. 48) with distal end tongue-shaped and caudal end slender, elongated trowel-shaped. Distribution (Fig. 61, dots). The species is recorded from the Moluccan islands Ambon, Saparua and Seram and from the Raja Ampat Islands (Papua Indonesia) Waigeo, Salawati and Misool. One old specimen from W. Doherty (in BMNH) is from the Island Roon, north from Wandammen Peninsula. In general Doherty s labelling proved to be rather accurate, but including Roon in the distribution area of clathratum seems very peculiar. Variation. On the Raja Ampat Islands the species is darker brown (Fig. 8) with even stronger brown veins crossing the fascia. On the hindwings the dark coloured veins are hardly fading towards base. Type material. Holotype?, Indonesia: Indonesia, NW Halmahera, blind road Baru-Basale, Gn. Talagarama, 550 m, 3 5.iii.1997, S. Naumann (zman). Paratypes: 5?, 2/: Indonesia: 1?, Halmahera, Van Diejen, (rmnh); 1?, Halmahera, Tobelo, bought from E. Le Moult, (bmnh); 2?, 2/, Molukken, Halmahera, Mt. Talagaranu, 600m, 15 km SE Baru, primärwald, 1 12' N 127 32' E, 22 31.i.1996, leg. Sinjaev & Tarasov (cmwm); 1?, Morotai, leg. Bernstein (rmnh). Diagnostic characters. Length of forewing 23 mm. Forewing dark grey-brown. Basal half of wing with white veins, a broad white wingfold, longitudinal line in cell broad. Fascia very broad and crossed by broad suffused veins, with costal and tornal patch large. Hindwing with narrow hindmargin and veins very broadly suffused with brown, gradually narrowing towards base but not fading. Male genitalia (Figs 23 26). Uncus with a strong dorsal keel (Fig. 24) running from base to apex. Apex of uncus short and slender (Fig. 25), and ventrum of uncus with one shallow and one deep bulge. Valve processes (Fig. 23) short, thick and almost equal in length, the apical process straight with an oblique apex, the others curved with a blunt apex. Aedeagus (Fig. 26) straight, coecum broad and curved downwards. Female genitalia (Figs 49 50). Ostium with two small and deep narrow pockets. Sclerotized part of ductus bursae long with two curls of 180. Ductus seminalis originating from the middle of the ductus bursae, just after the curls, and becomes enlarged. Two boat-shaped signa (Fig. 50) with faint remnant of a connection between the two. Distribution (Fig. 61, squares). The species is recorded from the northern Moluccan Islands Halmahera and Morotai. Etymology. The species name, an adjective, refers to the old name for Halmahera, Gilolo, which is also the name for a village on the island.

46 Tijdschrift voor Entomologie, volume 150, 2007 27 28 29 30 Figs 27-30. Male genitalia of Nyctemera leopoldi (prep. RV1150). 27, outside of right valve; 28, dorsal view of uncus; 29, lateral view of uncus; 30, aedeagus. Nyctemera leopoldi Tams Fig. 11 12, 27 30, 51 52, 61 Nyctemera leopoldi Tams 1935: 39 Holotype?, Indonesia: Arfak, Sakoemi, (Nouvelle Guinée), 11.iii.1929, H.R.H. Prince Leopold (ISNB) [examined] Nyctemera leopoldi: Bryk 1937: 68 Material examined. 83 specimens in addition to cited types. Indonesia, Papua: 3?, Mafor [Numfor, no data], ex coll. Fruhstorfer; 1/, Nieuw Guinea, 1929, v.d. Bergh (ZMHB); 1?, 1/, New Guinea, Ambubaki, L. Laglaize; 2/, New Guinea, Ditschi, Arfak, 1200 1500 m, v-vi.1928, E. Mayr; 2/, Dorey [Manokwari], iv.1897, W. Doherty; 5?, 6/, New Guinea, Mt. Siwi, Arfak, 800 m, iv-vi.1928, 23.iv.1928, 4.v.1928, 7.v.1928, 13.v.1928, E. Mayr; 1?, 2/, Dutch New Guinea, Ninay Valley, Central Arfak mts., 3500 ft, xi.1908-i.1909, ii-iii.1909; 11?, 10/, Irian Jaya, New Guinea, Arfak, Ngat Biep, 850 m, 18 19.xii.1993, Brechlin & Cerny (CKC); 3?, Irian Jaya, Arfak, Warkapi, 200 m, 16.xii.1993, Brechlin & Cerny (CKC); 2/, Irian Jaya, Warkapi, Arfak Mts., 500 m, 12.xi.1993, A.J. de Boer, A.L.M. Rutten & R. de Vos (ZMAN); 14?, 1/, Noord Nieuw Guinea, 1929, P.J. van den Bergh (BMNH: 1?, RMNH); 1?, Dutch New Guinea, Wai Sai River, Weyland Mts. [Kobowre Mts.], 1000 ft, vi.1920, C.F. & J. Pratt; 4?, 3/, Dutch New Guinea, Weyland Mts. [Kobowre Mts.], Dewaro Village, 3500 ft, vi.1920, C.F. & J. Pratt; 1?, 2/, 1/, Nieuw Guinea, Vogelkop [Doberai], 1929, P.J. van den Bergh (WAU); Irian Jaya, Senopi, Vogelkop [Doberai], 2.vii.1983, H. van Mastrigt (KSP); 2?, Irian Jaya, Subyo, Vogelkop [Doberai], 10.vii.1983, H. van Mastrigt (KSP); 1?, Ned. Nieuw Guinea, Achterland Hollandia [Jayapura], iv.1928, W. Stüber (ZMAN) (collection BMNH unless otherwise mentioned). Diagnostic characters. Length of forewing 19 22 mm. Forewing colour pale grey-brown. Basal half of wing with veins only narrowly white, wingfold and longitudinal line in cell narrowly white. Fascia not extremely broad with suffused crossing veins darker than wing colour. Forewing apex somewhat darker than rest of wing. Hindwing with rather broad pale grey-brown hind margin fading into the white central area. Veins suffused with brown, not fading or narrowing. Male genitalia (Figs 27 30). Uncus with short arched dorsal keel (Fig. 29). Apex of uncus thick and at an angle of 45 (Fig. 29), ventrum straight. Three valval processes long, slender with pointed apex (Fig. 27), the costal process very slender and longer than the other two. Aedeagus long (Fig. 30), slightly curved, coecum long and straight. Female genitalia (Figs 51 52). Pockets inside ostium very wide and shallow. Antrum broad, sclerotized part of ductus bursae short with 180 curl. Signum with slender distal part and pointed apex, caudally becoming broad, spade-shaped with pronounced apices. Distribution (Fig. 61, triangles). The species seems to prefer mountainous areas and is found in Papua Indonesia on the Birdshead Peninsula, mainly in the Arfak Mountains, and known from a few records from the Kobowre (Weyland) Mountains. It has also been recorded from the island Mefor (now Numfor) (BMNH). A single male specimen is known from an area south of Jayapura in Northeast Papua (ZMAN). Nyctemera oninica sp. n. Figs 13 14, 31 34, 53 54, 62 Type material. Holotype?, Indonesia: Fak- Fak, Dutch New Guinea, i-ii.1908, [A.E.] Pratt, (BMNH). Paratypes: 20?, 17/: Indonesia: 5?, 3/, Fak-Fak, Dutch New Guinea, 1700 ft, xi.1907 (1?), xii.1907 (3?, 1/), i-ii.1908 (1?, 2/),

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 47 31 32 33 34 Figs 31-34. Male genitalia of Nyctemera oninica (prep. RV1182). 31, outside of right valve; 32, dorsal view of uncus; 33, lateral view of uncus; 34, aedeagus. A.E. Pratt (BMNH); 4?, 1/, Kapaur [= Fakfak], xii.1896 (1? without date), W. Doherty (OXUM: 1?, BMNH: 3?, 1/); 1?, Kapaur, i.1897, low e.[levation], W. Doherty (BMNH); 1?, 1/, Dutch New Guinea, Wandammen Mts., 3 4000 ft, xi.1914, A.C. & F. Pratt (BMNH); 2?, 3/, Ile de Ron [= Roon], Baie de Geelwink, 1892, W. Doherty, (BMNH); 4?, 2/, Ron J., vii.1897, W. Doherty, (BMNH); 3?, 1/, Roon, W. Doherty, (BMNH); 5/, Dutch New Guinea, Roon, Geelvink Bay, vii.1920, C.F. & J. Pratt, (BMNH); 1/, Indonesia, Irian Jaya, Roon, Yende, at light, 60 m, 7.xi.1993, A.J. de Boer, A.L.M. Rutten & R. de Vos (ZMAN). Diagnostic characters. Length of forewing 21 22 mm. Forewing chocolate brown. Basal half of wing with veins narrowly white, wingfold and longitudinal line in cell narrowly white. Fascia very broad with crossing veins very narrow suffused with brown, costal and tornal patch of fascia large, the latter usually connected with the white wingfold. White hindwing with narrow brown hind margin. Veins not suffused, in wingbase only the cubital vein and anal vein with grey-brown. Male genitalia (Figs 31 34). Uncus with short and arched dorsal keel and dorsally two pronounced lateral ridges (Fig. 32). Apex of uncus short and thick (Fig. 33), ventrum straight. Valve with three processes very long, of equal length, with sharp apices (Fig. 31). Aedeagus rather short and curved, coecum straight (Fig. 34). Female genitalia (Figs 53 54). Inside of ostium with two very small pockets. Antrum broad, sclerotized part of ductus bursae short with an angle of about 70. Signum (Fig. 54) with long and very slender distal part, caudal part regular spade-shaped. Distribution (Fig. 62, dots). The species is found in Papua Indonesia on the Onin Peninsula, the Wandammen Peninsula and Roon Island. Variation. Specimens from Roon Island and Wandammen Peninsula have very narrow suffused veins on the hindwings (Fig. 14). Etymology. The name, an adjective, refers to the Onin Peninsula from where the holotype originates. Nyctemera mesolychna Meyrick Figs 15 16, 35 38, 55 56, 62 Nyctemera mesolychna Meyrick 1889: 466. Holotype /, Papua New Guinea: New Guinea, South Coast [= Mt. Obree], 1888, (BMNH) [examined] Nyctemera mesolychna: Kirby 1892: 419; Pagenstecher 1901: 116; Swinhoe 1903: 83; Seitz 1915: 271; Bryk 1937: 71. Deilemera pratti Bethune-Baker 1904: 412 Lectotype? (hereby designated), Papua New Guinea: British New Guinea, Dinawa, 4000 ft, ix.1902, A.E. Pratt, (BMNH) syn. n. [examined] Nyctemera pratti: Seitz 1915: 269; Bryk 1937: 75. Nyctemera baulus pratti: Roepke 1957: 169. Nyctemera baulus: Edwards 1996: 284 [partly misidentification] Material examined. 65 specimens in addition to cited types. Indonesia, Papua: 3?, Ned. Nieuw Guinea, Waris District, Ampas, 11.ix.1937, 12.ix.1937, W. Stüber, coll. J.M.A. van Groenendael (ZMAN). Papua New Guinea: 2?, New Guinea, Adelbert Mts., Wanuma, 800 m, 24.x.1958, 26.x.1958, J.L. Gressitt (BPBM); 2?, 1/, British New Guinea, Angabunga River, affl. of St. Joseph river, 6000 ft, xi.1904-ii.1905, A.S. Meek; 1?, 2/, New Guinea, Aroa River, A.S. Meek; 9?, 9/, British New Guinea, Upper Aroa River, i-iv.1903, ii.1903, A.S. Meek; 2?, 1/, British New Guinea, Dinawa, 4000 ft, vii.1902, ix.1902, xii.1902, A.E. Pratt; 2/, British Central New Guinea, Ekeikei, 1500 ft, iii-iv.1903, A.E. Pratt; 5?, 5/, British New Guinea, Hydrographer Mts., 2500 ft, i.1918, i-ii.1918, ii.1918, Eichhorn brothers; 1/ (paralectotype)

48 Tijdschrift voor Entomologie, volume 150, 2007 35 36 39 40 37 41 38 42 Figs 35-38. Male genitalia of Nyctemera mesolychna (prep. RV274). 35, outside of right valve; 36, dorsal view of uncus; 37, lateral view of uncus; 38, aedeagus. British New Guinea, Mt. Kebea, 6000 ft, iii-iv.1903, A.E. Pratt; 1/, British New Guinea, Mt. Kebea, 6000 ft, vii.1903, A.E. Pratt; 1?, 3/, British New Guinea, Mt. Kebea, 3000 ft, vi.1903, A.E. Pratt; 5?, New Guinea, Mafulu, 4000 ft, i.1934, L.E. Cheesman; 2?, British New Guinea, Owgarra, Upper Aroa River, ii.1905, A.S. Meek; 2/, German New Guinea, Simbang, Webster; 1?, 1/, New Guinea, Wau, 4000 ft, 9.xii.1961, 22.vii.1962, J. Sedlacek (BPBM); 2?, 2/, British New Guinea, Welsch River [Kwikila], Weiske (collection BMNH, unless otherwise mentioned). Diagnostic characters. Length of forewing 19 21 mm. Forewing brown. Basal half of wing with very narrow white veins, wingfold and longitudinal line in cell narrowly white, almost faded. White fascia rather narrow and slightly smoked with dark scales, crossed by suffused veins. Hindwing with rather broad hind margin. Veins only slightly suffused with brown, hardly visible. Dorsum of hindwing suffused with grey-brown. Male genitalia (Figs 35 38). Uncus with short, arched Figs 39-42. Male genitalia of Nyctemera latimargo (prep. RV1186). 39, outside of right valve; 40, dorsal view of uncus; 41, lateral view of uncus; 42, aedeagus. dorsal keel (Fig. 37), apex long, ventrum straight (Fig. 36). Three valval processes short, thick and of equal length (Fig. 35), the costal and apical processes curve inwards. Aedeagus (Fig. 38) long and slender, curved, with coecum long and straight. Female genitalia (Figs 55 56). Pockets inside ostium wide and stretched. Antrum broad, sclerotized part of ductus bursae rather long with 135 curl. Signum (Fig. 56) with slender distal part and sharp apex, caudal part broad spade-shaped with sharp apex. Distribution (Fig. 62, stars). The species is widely distributed through the eastern part of New Guinea, mainly in Papua New Guinea in mountainous areas at altitudes from sea level up to 1800 meters. Its westernmost records are just in the northeast of Papua Indonesia near the border of Papua New Guinea, south of Jayapura.

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 49 43 44 45 46 Figs 43-46. Male genitalia of Nyctemera dauila (prep. RV1188). 43, outside of right valve; 44, dorsal view of uncus; 45, lateral view of uncus; 46, aedeagus. Nyctemera latimargo (Rothschild) Figs 17, 39 42, 57 58, 62 Deilemera absurdum latimargo Rothschild 1915a: 78 Lectotype? (hereby designated), Indonesia: Dutch New Guinea, Base Camp, Setakwa River, sea level, xi.1912, A.F.R. Wollaston (BMNH). [examined] [Note: Rothschild mentioned seven females, but in fact these are all males] Nyctemera absurdum latimargo: Bryk 1937: 48 [partly misidentification]. Material examined. 30 specimens in addition to cited types. Indonesia: 1?, Dutch New Guinea, Canoe Camp, Setakwa River, xii.1912, A.F.R. Wollaston; 1?, Dutch New Guinea, nr Oetakwa [Utakwa] River, Snow Mts., 3500 ft, x-xii.1910, A.S. Meek; 6? (paralectotypes), Dutch New Guinea, Base Camp, Setakwa River, sea level, xi.1912 (4?), xi-xii.1912 (1?), i.1913 (1?), A.F.R. Wollaston; 9?, 6/, Dutch New Guinea, Upper Setekwa River, Snow Mts., 2000 3000 ft, vii.1910, A.S. Meek; 3/, Dutch New Guinea, Upper Setekwa River, Snow Mts., 2000 3000 ft, viii.1910, A.S. Meek (all BMNH); 1?, Irian Jaya, 45 km N. Wamena, 2100 m, 4.xii.1997, K. Cerny (CKC). Diagnostic characters. Length of forewing 20 21 mm. Forewing dark brown. Basal half of wing with white veins, wingfold and longitudinal line in cell narrow white. Fascia rather narrow, crossed by dark brown veins. Hindwing with very broad dark brown hind margin. Veins in the white centre of the wing with scarce brown scaling, hardly visible, only at the hind margin shortly running towards base with brown. Wingbase suffused with grey-brown, running along the cubital vein and anal vein. Male genitalia (Figs 39 42). Uncus with long and strongly arched dorsal keel (Fig. 41). Apex of uncus thick and curved downwards at nearly 90. Ventrum of uncus straight. Three valval processes short (Fig. 39), the apical process somewhat longer than the others. The process of the sacculus points posteriorally, the apical and costal process curve inwards. Aedeagus (Fig. 42) almost straight, coecum long and straight. Female genitalia (Figs 57 58). Inside ostium with two small wide and deep pockets. Antrum broad, sclerotized part of ductus bursae short with about 110 curl. Distal part of signum (Fig. 58) slender with sharp and curled apex, caudal part spade-shaped with sharp and curled apex. Distribution (Fig. 62, diamonds). Almost all records are from the Snow Mountains (now Sudirman Mountains, Papua Indonesia) along the Utakwa River and Setakwa River. One specimen was recently collected in the Baliem Valley (Jayawijaya Mountains), 45 km north from Wamena (CKC). It seems to have no particular altitude preference and is found from sea level up to 2100 meters. Nyctemera dauila sp. n. Figs 18, 43 46, 59 60, 62 Type material. Holotype?, Papua New Guinea: Goodenough Isl., 2500 4000 ft, iv.1913, A.S. Meek (BMNH). Paratypes: 4?, 3/, Papua New Guinea: Goodenough Isl., 2500 4000 ft, iv.1913 (3?, 1/), v.1913 (1?, 2/), A.S. Meek (BMNH). Diagnostic characters. Length of forewing 21 22 mm. Forewing dark brown. Basal half of wing with faded white veins, wingfold and longitudinal line in cell very narrowly white. Fascia narrow, crossed by dark brown veins, margin of fascia not sharp but faded. Hindwing with broad dark brown hind margin. Veins in white centre of wing not suffused, only dentating the inner edge of the hind margin. Dorsum suffused with grey-brown and connected with the greybrown wingbase. Male genitalia (Figs 43 46). Uncus with long and

50 Tijdschrift voor Entomologie, volume 150, 2007 47 49 48 50 Figs 47-48. Female genitalia of Nyctemera clathratum. 47, habitus; 48, signum (prep. RV1181). 51 Figs 49-50. Female genitalia of Nyctemera giloloensis. 49, habitus; 50, signum (prep. RV1222). 53 52 54 Figs 51-52. Female genitalia of Nyctemera leopoldi. 51, habitus; 52, signum (prep. RV335). Figs 53-54. Female genitalia of Nyctemera oninica. 53, habitus; 54, signum (prep. RV1183).

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 51 55 57 56 58 Figs 57-58. Female genitalia of Nyctemera latimargo. 57, habitus; 58, signum (prep. RV1187). Figs 55-56. Female genitalia of Nyctemera mesolychna. 55, habitus; 56, signum (prep. RV1190). 59 strongly arched dorsal keel (Fig. 45). Apex of uncus thick and curved downwards at 45. Ventrum of uncus almost straight. Valve (Fig. 43) with process of sacculus short and thick, apical and costal process of equal length but the costal process much more slender with a sharp apex. Aedeagus (Fig. 46) long and curved, coecum long and straight. Female genitalia (Figs 59 60). Inside of ostium with two small wide pockets. Sclerotized part of ductus bursae narrow and long with about 100 curl. Signum (Fig. 60) very long with slender distal part with blunt apex, caudal part elongated trowel-shaped. Distribution (Fig. 62, triangles). The species has only been found on Goodenough Island (Papua New Guinea, D Entrecasteaux Islands). Etymology. The name, a noun in apposition, refers to the local and old name of Goodenough Island, Dauila. 60 Figs 59-60. Female genitalia of Nyctemera dauila. 59, habitus; 60, signum (prep. RV1189). Discussion The Nyctemera clathratum-complex comprises seven species which are restricted to the Moluccas and New Guinea and some adjacent islands. The group is rather uniform in wing pattern and also the genitalia show many similarities suggesting monophyly of the group. In the male the similarly shaped uncus and three processes of the valve, and in the female the funnel-shaped antrum, the curled sclerotized part of

52 Tijdschrift voor Entomologie, volume 150, 2007 61 Figs 61-62. Distribution maps of the Nyctemera clathratum group. 61, Nyctemera clathratum (dots), N. giloloensis (squares) & N. leopoldi (triangles); 62, N. oninica (dots), N. latimargo (diamonds), N. mesolychna (stars) & N. dauila (triangles). 62 ductus bursae and the spade- or trowel-shaped signum, are characters of the clathratum-complex that are not found in other Nyctemera species groups in this combination. Nyctemera giloloensis takes a somewhat isolated position in this group. The most peculiar deviating character is the differently shaped signum in the female genitalia, which is divided into two boat-shaped parts (Fig. 50). Other genitalia and wing pattern characters, however, prove that it correctly belongs in the clathratum-group. Acknowledgements The author likes to thank all persons mentioned for allowing him to study the material needed for this revision: Behnaz van Bekkum-Ansari (RMNH, Leiden, The Netherlands), Karel Cerny (Zirl, Austria), Willem Hogenes (ZMAN, Amsterdam, The Netherlands), Martin Honey (BMNH, London, UK), Rienk de Jong (RMNH, Leiden, The Netherlands), Henk van Mastrigt (Jayapura, Indonesia), Wolfram Mey (ZMHB, Berlin, Germany), Scott Miller (BPBM, Honolulu, USA, now Smithsonian Institution), Matthias Nuss (SNSD, Dresden, Germany), Caroline Pepermans (RMNH, Leiden, The Netherlands), Chris O Toole (OXUM, Oxford, UK), Thomas Witt (Munich, Germany), Rob Zwart (WAU, Wageningen, The Netherlands), Patrick Grootaert (ISNB, Brussels, Belgium). Special thanks are due to Willem N. Ellis (Amsterdam, The Netherlands) for taking the digital photographs of the genitalia, Godard Tweehuizen (NEV-library, Amsterdam) for finding all necessary publications and the Uyttenboogaart-Eliasen Foundation (Dutch Entomological Society) for financing most of the visits to the museum collections.

De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae) 53 References Bethune-Baker, G.T., 1904. New Lepidoptera from British New Guinea. Novitates Zoologicae 11(2): 367 429. Bryk, F., 1937. Arctiidae, Subfam.: Callimorphinae et Nyctemerinae. Lepidopterorum Catalogus, 82: 1 105. Butler, A.G., 1880. On a second Collection of Lepidoptera made in Formosa by H.E. Hobson, Esq. Proceedings of the Zoological Society of London 1880: 666 691. Dubatolov, V.V., 2006. On the generic status of the Afrotropical Nyctemera species (Lepidoptera, Arctiidae). Atalanta 37(1/2):191 205. Edwards, E.D., 1996. Arctiidae. In: E.S. Nielsen, E.D. Edwards & T.V. Rangsi, 1996. Checklist of the Lepidoptera of Australia. Monographs on Australian Lepidoptera 4: 278 285. Eecke, R. van, 1927. Een en ander over de soorten van het genus Nyctemera Huebn. Entomologische Berichten, Amsterdam 7: 220 223. Holloway, J.D., G. Kibby & D. Peggie, 2001. 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Subfamilie: Callimorphinae & VII. Subfamilie: Nyctemerinae. In: A. Seitz (ed.). Die Gross-Schmetterlinge der Erde 10: Spinner und Schwärmer des Indo-Australischen Gebiets: 264 290. A. Kernen, Stuttgart. Snellen van Vollenhoven, S.C., 1863. Bijdrage tot de kennis van het vlindergeslacht Leptosoma Boisd. Tijdschrift voor de Dierkunde 1: 35 51. Swinhoe, C., 1892. Catalogue of Eastern and Australian Lepidoptera Heterocera in the collection of The Oxford Museum, part I, Sphinges and Bombyces. At The Clarendon Press, Oxford, 324 pp. Swinhoe, C., 1903. On the Genus Deilemera Hübner. Transactions of the Entomological Society of London 1903: 53 85. Tams, W.H.T., 1935. Résultats scientifiques du voyage aux Indes Orientales Néerlandaises de LL. AA. RR. 1e Prince et la Princesse Leopold de Belgique, Lepidoptera II, 2. Heterocera. Verhandelingen van het Koninklijk Natuurhistorisch Museum van Belgie (Buiten Reeks) 4(12): 31 64 (11 plates). Tuxen, S.L., 1970. Taxonomist s glossary of genitalia in insects. Munksgaard, Copenhagen, 359 pp. Vos, R. de, 1995a. Nyctemera groenendaeli spec.nov. from New Guinea (Lepidoptera: Arctiidae, Nyctemerinae). Nachrichten des Entomologischen Vereins Apollo 15(4): 481 489. Vos, R. de, 1995b. A revision of Nyctemera consobrina (Hopffer, 1874) with redescriptions of three subspecies (Lepidoptera: Arctiidae, Nyctemerinae). Nachrichten des Entomologischen Vereins Apollo 16(1): 81 93. Vos, R. de, 1996. Nyctemera pseudokala sp.nov. and N. mastrigti sp.nov., two new species from Indonesia (Lepidoptera: Arctiidae, Nyctemerinae). Nachrichten des Entomologischen Vereins Apollo 17(3): 301 311. Vos, R. de, 1997a. A revision of Nyctemera kebeae (Bethune-Baker, 1904) (with descriptions of two new subspecies) and N. warmasina (Bethune-Baker, 1910) (Lepidoptera: Arctiidae, Nyctemerinae). Nachrichten des Entomologischen Vereins Apollo 18(1): 1 12. Vos, R. de, 1997b. The identity of Nyctemera simulatrix Walker, 1864 (Lepidoptera: Arctiidae, Nyctemerinae). Nachrichten des Entomologischen Vereins Apollo 18(2 3): 205 210. Vos, R. de, 2002. Revision of the Nyctemera evergista group (= subgenus Deilemera Hübner) (Lepidoptera: Arctiidae, Arctiinae, Nyctemerini). Nachrichten des Entomologischen Vereins Apollo 23(1/2): 7 32. Vos, R. de, 2007. The Utetheisa species of the subgenera Pitasila Moore, Atasca Swinhoe and Raanya sg. n. (Lepidoptera: Arctiidae, Arctiinae). Aldrovandia 3 (in press). Vos, R. de & K. Cerny, 1999. A review of the Philippine

54 Tijdschrift voor Entomologie, volume 150, 2007 species of the genus Nyctemera Hübner, [1820] with descriptions of new species and subspecies (Lepidoptera: Arctiidae, Nyctemerinae). Nachrichten des Entomologischen Vereins Apollo 20(2): 133 188. Received: 27 December 2006 Accepted: 7 April 2007