Revision of the Nyctemera clathratum complex (Lepidoptera

IntroductionThe genus Nyctemera Hübner, [1820] s.l. is a large heterogeneous taxon with many species in South-east 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 di-vided 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 com-plex 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 spe-cies clearly belong to the nominotypical subgenus Nyctemera s.s. The valves in the male genitalia all possess three processes. The N. clathratum com-plex 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 proc-esses 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. Further-more the banded pattern on the abdomens of the species of the clathratum complex is brown-white (the white only narrow), while in the baulus com-plex 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. How-ever, 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 complemen-tary, not overlapping distributions.

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. [email protected].

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, UKBPBM Bernice P. Bishop Museum, Gressitt Center

for Research, Honolulu, Hawaii, USAKSP Koleksie Serangga Papua (Private collec-

tion Henk van Mastrigt), Jayapura, Papua, Indonesia

CKC Private collection Karel Cerny, Zirl, Austria

CMWM Museum Thomas Witt (assigned to Zo-ologische Staatssammlung München), Munich, Germany

ISNB Institute Royal des Sciences Naturelles de Belgique, Brussels, Belgium

OXUM Hope Entomological Collections, Univer-sity Museum, Oxford, UK

RMNH Nationaal Natuurhistorisch Museum Naturalis, Leiden, The Netherlands

SNSD Staatliche Naturhistorische Sammlungen, Dresden, Germany

WAU Agricultural University, Wageningen, The Netherlands

ZMAN Zoölogisch Museum, University of Amsterdam, The Netherlands

ZMHB Museum für Naturkunde der Humboldt Universität, Berlin, Germany

MethodsGenitalia 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 con-ventional Nikon F400 camera, with Sigma AF52 Macro-objective and Sunpack ring flashlight, using Fuji 100 ASA colour transparency film. The speci-mens 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 struc-tures (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 approxi-mately 65 Indo-Australian species show that at least six species groups (termed “subgenera” hereafter) can be recognized. The nominotypical subgenus Nyctem-era, is in the male genitalia distinguished from all others by the rather short valvae with three process-es, 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). Further-more it is recognized by its wing pattern with a clear-ly 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 Try-pheromera Butler, 1881 broader triangular, in Arctata even longer stretched and in Deilemera almost oval-shaped 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 Nyctem-era 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

DiagnosisHead 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

41De Vos: Nyctemera clathratum complex (Lepidoptera: Arctiidae)

colour of thorax and tegulae is pale yellow or bone-white. 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

Figs 1-4. Important genital structures of the Nyctemera clathratum group. – 1, Three processes of the male val-va; 2, lateral view of the male uncus with keel, apex and ventrum; 3, aedeagus with coecum; 4, female genitalia structures.

apical process

ventrum

2costal process

process on sacculus

1

coecum

3

4 ostium

ductus bursae (sclerotized)

pockets

signum

bursa copulatrix

ductus bursae (unsclerotized)

ductus seminalis


5 6

Figs 5-12. Nyctemera clathratum group, adult habitus. – 5, N. clathratum, male, Ambon (SNSD); 6, N. clath-ratum, 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).

7 8

9 10

11 12


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 dis-cal 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 saccu-

lus, 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 surround-ing 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 di-agnostic. The bursa copulatrix is large, globular, and

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.

13 14

15 16

17 18


possesses one long spade-shaped signum, except for giloloensis, whose signum is divided into two boat-shaped parts.

Checklist Nyctemera Hübner, [1820]: 178 Type species Phalaena lacticinia Cramer, 1777Subgenus Nyctemera s.s.

Nyctemera clathratum complexclathratum (Snellen van Vollenhoven, 1863) absurdum (Swinhoe, 1892) syn. n.giloloensis sp. n.leopoldi Tams, 1935oninica 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 saccu-lus 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.

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.

19 20

21

22


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 slen-der, 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 accu-rate, 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.

The only known specimen from Roon has the suffu-sion 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 misidentifica-tion]

Nyctemera clathrata clathrata: Roepke 1949: 66 [partly misidentification]

Type material. Holotype ?, Indonesia: “Indone-sia, NW Halmahera, blind road Baru-Basale, Gn. Talagarama, 550 m, 3–5.iii.1997, S. Naumann” (zman). Paratypes: 5?, 2/: Indonesia: 1?, “Hal-mahera, Van Diejen”, (rmnh); 1?, “Halmahera, To-belo, 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 narrow-ing towards base but not fading.Male genitalia (Figs 23–26). Uncus with a strong dor-sal 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 down-wards.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 record-ed 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.

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.

23 24

25

26


Nyctemera leopoldi TamsFig. 11–12, 27–30, 51–52, 61

Nyctemera leopoldi Tams 1935: 39 Holotype ?, Indone-sia: “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 [Dobe-rai], 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, Achter-land 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 cen-tral 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 Num-for) (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/),

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.

27 28

29

30


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. Do-herty”, (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 longitu-dinal 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 usu-ally connected with the white wingfold. White hind-wing with narrow brown hind margin. Veins not suf-fused, 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 lat-eral ridges (Fig. 32). Apex of uncus short and thick (Fig. 33), ventrum straight. Valve with three proc-esses 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 dis-tal part, caudal part regular spade-shaped.Distribution (Fig. 62, dots). The species is found in Papua Indonesia on the Onin Peninsula, the Wan-dammen Peninsula and Roon Island.Variation. Specimens from Roon Island and Wan-dammen 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 MeyrickFigs 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 misidentifi-

cation]

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)

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.

31 32

33

34


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 Guin-ea, Mafulu, 4000 ft, i.1934, L.E. Cheesman; 2?, Brit-ish 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 other-wise 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

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, cau-dal 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 ar-eas at altitudes from sea level up to 1800 meters. Its westernmost records are just in the northeast of Pa-pua Indonesia near the border of Papua New Guinea, south of Jayapura.

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.

35 36

37

38

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.

39 40

41

42


Nyctemera latimargo (Rothschild)Figs 17, 39–42, 57–58, 62

Deilemera absurdum latimargo Rothschild 1915a: 78 Lec-totype ? (hereby designated), Indonesia: “Dutch New Guinea, Base Camp, Setakwa River, sea level, xi.1912, A.F.R. Wollaston” (BMNH). [examined] [Note: Roth-schild mentioned seven females, but in fact these are all males]

Nyctemera absurdum latimargo: Bryk 1937: 48 [partly misi-dentification].

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°. Ven-trum of uncus straight. Three valval processes short (Fig. 39), the apical process somewhat longer than the others. The process of the sacculus points poste-riorally, 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, sclero-tized 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 Moun-tains, 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 par-ticular 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 dentat-ing the inner edge of the hind margin. Dorsum suf-fused with grey-brown and connected with the grey-brown wingbase.Male genitalia (Figs 43–46). Uncus with long and

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.

43 44

45

46


Figs 47-48. Female genitalia of Nyctemera clathratum. 47, habitus; 48, signum (prep. RV1181).

47

48

Figs 49-50. Female genitalia of Nyctemera giloloensis. 49, habitus; 50, signum (prep. RV1222).

Figs 51-52. Female genitalia of Nyctemera leopoldi. 51, habitus; 52, signum (prep. RV335).

51

52

Figs 53-54. Female genitalia of Nyctemera oninica. 53, habitus; 54, signum (prep. RV1183).

53

54

49

50


strongly arched dorsal keel (Fig. 45). Apex of uncus thick and curved downwards at 45°. Ventrum of un-cus 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 slen-der 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. Sig-num (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.

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

Figs 59-60. Female genitalia of Nyctemera dauila. 59, habitus; 60, signum (prep. RV1189).

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).

55

5856

57

60

59


ductus bursae and the spade- or trowel-shaped sig-num, are characters of the clathratum-complex that are not found in other Nyctemera species groups in this combination. Nyctemera giloloensis takes a somewhat isolated posi-tion in this group. The most peculiar deviating char-acter 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, Lon-don, UK), Rienk de Jong (RMNH, Leiden, The Netherlands), Henk van Mastrigt (Jayapura, Indo-nesia), Wolfram Mey (ZMHB, Berlin, Germany), Scott Miller (BPBM, Honolulu, USA, now Smithso-nian 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 (Am-sterdam, The Netherlands) for taking the digital photographs of the genitalia, Godard Tweehuizen (NEV-library, Amsterdam) for finding all neces-sary publications and the Uyttenboogaart-Eliasen Foundation (Dutch Entomological Society) for financing most of the visits to the museum collec-tions.

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. la-timargo (diamonds), N. mes-olychna (stars) & N. dauila (triangles).

61

62


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Received: 27 December 2006Accepted: 7 April 2007

Revision of the Nyctemera clathratum complex (Lepidoptera

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