Tijdschrift voor Entomologie 161 (2018) 39–62

Revision of the Cyphura Warren, 1902. Part II: The caudiferaria-, phantasmah- and pardata-group from the Indo-Pacific region with eight new species (: : Uraniinae) Siep Sinnema & Jannie Sinnema-Bloemen

The caudiferaria-, phantasmah- and pardata-group of the genus Cyphura Warren, 1902 (Lepidoptera: Uraniidae) are revised. Eight new species are described. There are six new species in the caudiferaria-group: Cyphura angusta, C. continua, C. dilatata, C. interrupta, C. laeensis and C. tobeloensis. In this group, a holotype is identified for Srophidia clarissima Butler, 1879 and a lectotype is designated for Urapteroides latimarginata Swinhoe, 1902. In the phantasmah-group, two new species are described: C. astrolabensis and C. mastrigti. In this group a lectotype and paralectotypes are designated for Strophidia phantasmah Felder & Rogenhofer, 1875. In the pardata-group, a lectotype and paralectotypes are designated for Cyphura pardata Warren, 1906. Keywords: Lepidoptera; Uraniidae; Uraniinae; Cyphura; Xysterophora S.G. Sinnema*, J.W Sinnema-Bloemen, Sparjeburd 29, NL-8409 CK Hemrik, The Netherlands. [email protected]

Introduction Because of external and genital differences, the sub- Cyphura Warren, 1902 is a probably monophyletic genus Xysterophora Sinnema & Sinnema-Bloemen,­ genus (Uraniidae, Uraniinae) (Lees & Smith, 1991) 2016 (type species Cyphura albisecta Warren, 1907) with about 30 species; it is almost totally restricted was introduced. Based on external differences,­ to New Guinea. A few species occur in the Moluc- the remaining species in the genus Cyphura (sub- cas, Sulawesi and northern Australia. All species in genus Cyphura s.s.) are divided into five species the genus have white wings with a more or less broad groups: the geminia-group, the caudiferaria-group, blackish band along the termen of the forewing and the phantasmah-group, the semiobsoleta-group and a submarginal blackish or brown band on the hind- the pardata-group. In this paper the members of the wing. All species have a more or less extended tail caudiferaria-,­ phantasmah- and pardata-group will be on the hindwing, with in most species two or three discussed and eight new species will be described. black dots at the base of the tail. On the hindwing The members of the caudiferaria-group have black there is always a fairly large black spot between veins or brown margins on the white forewing and a black M3 and CuA1 and between CuA1 and CuA2. Some or brown submarginal band on the white hindwing. species have two brown transverse bands on the fore- There is no trace of brown transverse bands on both wing, other species have no additional markings on wings. In the male genitalia, the valves are broad, the forewing; in some there is a brown transverse hairy and slightly hooked at the end of the costa. The band on the hindwing. harpe is slender with a thorn at its apex. The phallus

Tijdschrift voor Entomologie 161: 39–62, Figs 1–54. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 15 December 2018. DOI 10.1163/22119434-00002073 *Corresponding author Downloaded from Brill.com10/06/2021 06:56:51AM via free access

40 Tijdschrift voor Entomologie, volume 161, 2018 is tubular, slender, more or less hooked at the apex RMNH Naturalis Biodiversity Center (for- and with a small sclerotized tooth near the apex. In merly Rijksmuseum van Natuur­ the female genitalia the corpus bursae is oblong with lijke Historie or Nationaal Natuur­ a long ductus bursae; signa on the corpus bursae are historisch Museum), Leiden, The absent. The caudiferaria-group as interpreted here Netherlands consists of nine species: C. caudiferaria (Boisduval, SINNEMA private collection of the authors, 1832), C. latimarginata (Swinhoe, 1902), C. claris- Hemrik, The Netherlands sima (Butler, 1879) and six new species, which are ZMAN Naturalis Biodiversity Center (for- described in this paper. merly Zoölogisch Museum van Am- The members of the phantasmah-group are almost sterdam), Leiden, The Netherlands completely white, the forewing with a very narrow brown marginal band and the costa striated with Methods brown scales. The hindwing has a diffuse brown sub- Genitalia preparations were made according to the marginal band and the black dots near the tail are standards of the Natural History Museum in London more or less diffuse. In the male genitalia the valves (Robinson 1976). At least one male and one female are broad, hairy and slightly hooked at the end of specimen for each species were dissected. After dis- the costa. The harpe is slender with a thorn at its section the genitalia were macerated for 12 hours in apex. The phallus is tubular, slender, and more or less cold KOH 10%, cleaned in ethanol 30% and stained hooked at the apex, where there is a small sclerotized with a chlorazol black solution in absolute ethanol. tooth. In the female genitalia the corpus bursae is The genitalia were temporarily stored in ethanol short and club-shaped and there is no signum on the 70% to allow study of their three-dimensional struc- corpus bursae. The phantasmah-group as interpreted ture. Finally the genitalia were mounted on glass in here consists of three species: C. phantasmah (Felder euparal. Most of the adults were photographed with & Rogenhofer, 1874) and two new species, which a Canon Powershot SX30 IS digital camera. are described in this paper. Genitalia slides were studied with a WILD M3 The species of the pardata-group have yellow binocular microscope at magnifications of 60–400×. markings on both fore- and hindwing. In the male Digital automontage photographs were made with genitalia the valves are broad, hairy and not hooked a motorized Zeiss V20 binocular microscope and a at the end of the costa. The harpe is slender with a digital Axio MRc5 camera controlled by Axioman- thorn at its apex. The phallus is tubular, slender and ager M2 software. not pointed at the apex. In the female genitalia the Wing length was measured from wing base to corpus bursae is globular and bears a signum. The apex in males and females. pardata-group as interpreted here consists of one Morphological terminology of the external struc- species: C. pardata Warren, 1906. tures (excluding the genitalia) mainly follows Scoble (1992) and Holloway et al. (2001). The terminology of the genitalia mainly follows Jordan (1939), Tuxen Material and methods (1970) and Kôda (1987). In the plates the following abbreviations are used: Material R = recto (dorsal) and V = verso (ventral); the scale For this study specimens were examined from nine line represents a length of 1.0 cm. collections. This study is based on adults only; we have no information on immature stages, hostplants or life Institutional codens and abbreviations used histories. ANIC Australian National Collec- tion, Canberra, Australia KSP Kelompok Serangga Papua (former Checklist collection of Br. Henk van Mas- trigt), Jayapura, Papua, Indonesia Genus Cyphura Warren, 1902 MNHB Museum für Naturkunde der Humboldt-Universität zu Berlin, The caudiferaria-group. Germany Cyphura caudiferaria (Boisduval, 1832) [Indonesia: NHMUK Natural History Museum (formerly Papua; Papua New Guinea] British Museum for Natural His- = Asthenia mundaria Walker, 1866 [Indonesia: tory), London, United Kingdom Moluccas, Halmaheira] OMNH Oxford University Museum of Nat- = Strophidia pannata Felder & Rogenhofer, 1874 ural History, United Kingdom [Indonesia: Papua, Salawati Island] QMBA Queensland Museum, Brisbane, Cyphura latimarginata (Swinhoe, 1902) [Indonesia: Australia Papua; Papua New Guinea]Downloaded from Brill.com10/06/2021 06:56:51AM via free access

Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 41

Cyphura clarissima (Butler, 1879) [Papua New Guin- – Submarginal band on the upperside of hind- ea: New Ireland] wing not interrupted between tornus and anal Cyphura tobeloensis sp. n. [Indonesia: Moluccas, angle (Figs 11, 12) �����������������������C. continua Halmaheira] 7 The upperside of forewing with narrow pale Cyphura angusta sp. n. [Indonesia: Biak Island] brown marginal band, the upperside of hind- Cyphura interrupta sp. n. [Papua New Guinea: New wing with narrow, straight pale brown sub- Ireland] marginal band, on underside of hindwing Cyphura continua sp. n. [Papua New Guinea: New submarginal band hardly visible (Fig. 8) Britain] ������������������������������������������������������ C. angusta Cyphura laeensis sp. n. [Papua New Guinea: Lae] – The upperside of forewing with broad mar- Cyphura dilatata sp. n. [Papua New Guinea: Manki­ ginal band, more or less broadening towards Ridge] the apex, hindwing underside with distinct submarginal band ���������������������������������������� 8 The phantasmah-group. 8 All three black spots in the tail on the upper- Cyphura phantasmah ([C. & ? R.] Felder & Rogen- side of hindwing embedded in submarginal hofer, 1875) [Indonesia: Moluccas, Gebeh Island] band (Figs 3, 4) ��������������������� C. latimarginata Cyphura mastrigti sp. n. [Indonesia: Numfor Island] – At least one black spot in tail on the upperside Cyphura astrolabensis sp. n. [Papua New Guinea, As- of hindwing isolated from submarginal band trolabe Bay] �������������������������������������������������������������������� 9 9 The upperside of forewing with very broad The pardata-group. dark brown marginal band, strongly broad- Cyphura pardata Warren, 1906 [Indonesia: Foja ening towards the apex, together with costal Mountains; Papua New Guinea, mainland] margin covering about half the surface of the forewing (Fig. 14) ��������������������������C. dilatata Key to the Cyphura species of the – The brown outer margin together with the caudiferaria-, phantasmah- and costal margin covering less than half the surface of the the upperside of forewing pardata-groups ������������������������������������������������������������������ 10 1 The upperside of forewing and hindwing with 10 On the upperside of forewing the margins yellow markings (Figs 49, 50) ��������C. pardata and on the upperside of hindwing the sub- – The upperside of forewing and hindwing marginal band are pale brown (Figs 6, 7) white with only brown or black markings �������������������������������������������������� C. tobeloensis �������������������������������������������������������������������� 2 – On the upperside of forewing the margins and 2 The upperside of forewing white with a very on the upperside of hindwing the submarginal narrow brown marginal band ���������������������� 3 band are dark brown or black �������������������� 11 – The upperside of forewing white with a 11 Submarginal band on underside of hindwing distinct brown or black marginal band hardly visible near the anal angle (Fig. 5) �������������������������������������������������������������������� 5 ���������������������������������������������������C. clarissima 3 The upperside of hindwing white with brown – Submarginal band on underside of hindwing submarginal band from apex to anal angle continuing to the anal angle ���������������������� 12 (Fig. 37) ��������������������������������� C. phantasmah 12 The black spot in the tail on the upperside – The upperside of hindwing white with a of the hindwing is free from the submarginal trace of a very narrow brown submarginal band, the second spot touches the submargin- band, visible near the costa and the anal angle al band (Figs 1, 2) ��������������������C. caudiferaria �������������������������������������������������������������������� 4 – The black spot in the tail on the upperside 4 The upperside of forewing costa heavily stri- of the hindwing is free from the submargin- ated with brown (Figs 38, 39) ������ C. mastrigti al band, the second spot is isolated from the – The upperside of forewing costa slightly stri- submarginal band by a distinct white border ated with pale brown (Fig. 40) ���������������������� (Fig. 13) �����������������������������������������C. laeensis ����������������������������������������������� C. astrolabensis 5 The upperside of forewing with subcostal dark brown band ������������������������������������������������ 6 The caudiferaria-group – The upperside of forewing with brown mar- gin along the costa, more or less striated with Cyphura caudiferaria (Boisduval, 1832) brown or black �������������������������������������������� 7 Figs 1, 2, 15, 16, 31 6 Submarginal band on the upperside of hind- Urapteryx caudiferaria Boisduval, 1832: 255. Holo- wing interrupted between tornus and anal type: Indonesia (Papua, Birdshead Peninsula, Ma- angle (Figs 9, 10) ���������������������� C. interrupta nokwari) (depositoryDownloaded unknown) from Brill.com10/06/2021 [not examined] 06:56:51AM via free access

42 Tijdschrift voor Entomologie, volume 161, 2018

Asthenia mundaria Walker 1866: 1919. fringes are white; near the black dots the fringes are Lectotype mundaria (designated here) ♂, [Indonesia, black. ­Moluccas, Halmahera,] Gilolo (OMNH, from Mr. Underside of forewing identical to the upperside, Saunders Collection, coll.nr. 2062A) [examined] but the costa not striated with black. The marginal Strophidia pannata Felder & Rogenhofer 1874: Tf. band is broader than on the upperside. 128 fig. 39. Underside of hindwing identical to the upperside, Holotype (fixed by monotypy) ♂, [Indonesia, but the submarginal band is broader near the tornus; Papua Barat, Raja Ampat Islands,] Salawati Is- the two black spots are embedded in the submarginal land [NHMUK, coll. nr. BMNH(E)1325348] band. [examined] Abdomen — Dorsally and ventrally white. Female. Forewing length 34 mm. Antennae filiform, Additional material examined ventrally pale brown, dorsally white and brown C. mundaria: Paralectotypes: 2♀, [Indonesia, Mo- checkered, length 12 mm. luccas, Halmahera,] Gilolo (OMNH, coll.nr. 2062B Forelegs brown, tibiae and tarsi white. Other legs and 2062C) [examined] white, including tibiae and tarsi. Identical to male, C. caudiferaria: 1♂, Indonesia (Papua), Birdshead but the margins on the forewing slightly paler. Peninsula, Arfak Mountains, Mokwam, 1510 m, Male genitalia (Figs 15, 16). Valves (Fig. 15) nar- 8.xi.2011, 1°06′ S, 133°54′ E, leg. S. & J. Sinnema row, almost rectangular, strongly haired, slightly (coll. RMNH, prep. nr. RMNH-RV1383) [exam- hooked at the apex. Harpe slender, obliquely cross- ined]; 1♀, Indonesia (Papua), Birdshead Peninsula, ing the valve, reaching the edge of the valve. Small Arfak Mountains, Mokwam,1510 m, 8.xi.2011, thorn at the apex of the harpe. Sacculus narrow, bent 1°06′ S, 133°54′ E, leg. S. & J. Sinnema (coll. at one third from base. RMNH, prep. nr. RMNH-RV1384) [examined]; Phallus (Fig. 16) bent, tapering towards apex. 1♀, Indonesia (Papua), Birdshead Peninsula, Arfak Apex curved, hook-like, very slender, underside with Mountains, Mokwam, 1510 m, 24.x.2011, 1°06′ S, small sclerotized tooth close to the apex. The up- 133°54′ E, leg. S. & J. Sinnema (coll. SINNEMA) perside of apex almost straight. Coecum rather long, [examined] slender. Female genitalia (Fig. 31). Corpus bursae oblong, length corpus bursae is about the same as length duc- Description (Figs 1, 2) tus bursae. No signum on corpus bursae. Antrum Male. Head — Vertex dark brown, frons white with rather broad, small sclerotized papillae anales with dark brown spot on the upperside. Antennae pale many setae. brown, filiform, length 12 mm. Labial palpus: dor- sally white with black at the base and a small black spot at the apex, ventrally pale white. Distribution Thorax — Thorax white, patagia dark brown, te- Cyphura caudiferaria is widespread in New Guinea. gulae white. Thorax ventrally white. In Indonesia it is found in many places along the Forelegs femur brown in front, tibiae brown in coast of Papua Barat and the Birdshead Penninsula front, other parts and tarsi white. Other legs white, and on the Raja Ampat Islands, Biak and other is- including tibiae and tarsi. lands near the coast of Papua Barat. It is also found Forewing length 28 mm. Forewing white, with at higher altitudes like Mokwam in the Arfak Moun- broad dark brown marginal band, broadening to- tains and in the Cyclops Mountains. In the Moluccas wards apex. Costa with black margin, broadening it is known from the island Seram. In Papua New towards apex and slightly striated with black at the Guinea the distribution is limited to the mainland. basal part. The angle between the marginal band and the marginal band along the costa is slightly round- Cyphura latimarginata (Swinhoe, 1902) ed. Fringes black. Figs 3, 4, 17, 18, 32 Hindwing white with a brown submarginal band Urapteroides latimarginata Swinhoe, 1902: 599. of equal width, starting from the anal angle and end- Lectotype: ♂, Papua New Guinea, Kaiser Wilhelms- ing at the tornus, isolating a white margin at the mar- land (NHMUK, coll. nr. BMNH(E)1325344) ginal band. A small tail near the tornus. Two black [examined] dots near the tornus, the one in the tail rounded and not embedded in the submarginal band. The other Additional material examined black dot is smaller and elongated and touching the 1♂, Papua New Guinea, Stephansort, leg. Cotton and submarginal band. Near the anal angle the white is Webster (NHMUK, coll. nr. BMNH(E)1325345, slightly suffused with pale brown. Fringes black and prep. nr. BMNH1325345) [examined]; 1♀, Pap- pale on the outside, on the upperside of the tail the ua New Guinea, Astrolabe Bay, leg. C. Wahnes

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Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 43

Figs 1–4. Adults of Cyphura spp. — 1, C. caudiferaria, male (coll. nr. BMNH(E)1325348); 2, C. caudiferaria, female (coll. nr. RMNH-RV1384); 3, C. latimarginata, male (coll. nr. BMNH(E)1325345); 4, C. latimarginata, female (coll. nr. BMNH(E)1325346).

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44 Tijdschrift voor Entomologie, volume 161, 2018

Figs 5–8. Adults of Cyphura spp. — 5, C. clarissima, male, holotype; 6, C. tobeloensis, male, holotype; 7, C. tobeloensis, female, paratype (coll. nr. BMNH(E)1325373); 8, C. angusta, female, holotype.

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Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 45

Figs 9–12. Adults of Cyphura spp. — 9, C. interrupta, male, holotype; 10, C. interrupta, female, paratype; 11, C. continua, male, holotype; 12, C. continua, female, paratype.

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Figs 13, 14. Adults of Cyphura spp. — 13, C. laeensis, male, holotype; 14, C. dilatata, male, holotype.

(NHMUK, coll. nr. BMNH(E)1325346, prep. equal width. A small tail near the tornus. Three black nr. BMNH1325346)] [examined]; 1♂, Indone- dots near the tornus, all embedded in the submar- sia, coast between Geelvinck Bay and Humboldt ginal band. The black dot in the tail and the middle Bay, x–xii.1896, leg. Doherty (NHMUK, coll. one are rounded. The black dot near the anal angle nr. BMNH(E)1325350) [examined]; 1♀, Indone- is elongated. Fringes black and distally pale, on the sia, coast between Geelvinck Bay and Humboldt the upperside of the tail white, near the black dots Bay, x–xii.1896, leg. Doherty (NHMUK, coll. nr. fringes are black. BMNH(E)1325351) [examined] Underside of forewing identical to the upperside, but the dark marginal band is broader. The dark Description (Figs 3, 4) margin along costa fainting towards base, not stri- Male. Head — Vertex brown, frons white with black ated with black. spot on the upperside. Antennae pale brown, fili- Underside of hindwing identical to the upperside, form, length 15 mm. Labial palpus: dorsally white but the submarginal band is broader, isolating a very with black at the base and pale brown at the apex, narrow white margin along the marginal band, and ventrally white. also broader near the tornus. The three black spots Thorax — Thorax white, patagia brown, tegulae are totally embedded in the submarginal band. white. Thorax ventrally white. Abdomen — Dorsally and ventrally white. Forelegs femur brown in front, tibiae brown in Female. Identical to male, but dark margins and front, other parts and tarsi white. Other legs white, submarginal band on the hindwing slightly paler. including tibiae and tarsi. Forewing length 38 mm. Forewing length 32.0 mm. Forewing white, with Head — Vertex dark brown, frons white with broad dark brown marginal band, slightly broaden- large dark brown spot on the the upperside. Anten- ing towards apex. Costa with black margin, slightly nae pale brown, filiform, length 14 mm. broadening towards apex and slightly striated with Male genitalia (Figs 17, 18) (prep. nr. black at the basal part. The angle between the mar- BMNH1325345). Valves (Fig. 17) rounded, strong- ginal band and the marginal band along the costa is ly haired, slightly hooked at the apex. Harpe broad almost right and not rounded. Fringes black. at the base, tapering towards but not reaching the Hindwing white with a broad brown submarginal edge of the valve. Very small thorn at the apex of the band, starting from the anal angle, isolating a very harpe. Sacculus narrow, slightly bent. narrow white margin at the marginal band. The Phallus (Fig. 18) almost straight, tapering towards submarginal band is narrow near the tornus, along apex. Small sclerotized tooth close to the apex. Coe- the marginal band of the hindwing broader and of cum short and broad.Downloaded from Brill.com10/06/2021 06:56:51AM via free access

Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 47

Figs 15–22. Male genitalia of Cyphura spp. — 15, C. caudiferaria (prep. nr. RMNH-RV1383) (after removal of phallus); 16, C. caudiferaria, phallus; 17, C. latimarginata (prep. nr. BMNH1325345) (after removal of phallus); 18, C. latimarginata, phallus; 19, C. clarissima (prep. nr. BMNH-RV1582) (after removal of phallus); 20, C. claris- sima, phallus; 21, C. tobeloensis, holotype (after removal of phallus); 22, C. tobeloensis, phallus.

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Figs 23–30. Male genitalia of Cyphura spp. — 23, C. interrupta, holotype (after removal of phallus); 24, C. interrupta, phallus; 25, C. continua, holotype (after removal of phallus); 26, C. continua, phallus; 27, C. laeensis, holotype (after removal of phallus); 28, C. laeensis, phallus; 29, C. dilatata, holotype (after removal of phallus); 30, C. dilatata, phallus.

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Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 49

Figs 31–36. Female genitalia of Cyphura spp. — 31, C. caudiferaria (prep. nr. RMNH-RV1384), habitus; 32, C. latimarginata (prep. nr. BMNH1325346), habitus; 33, C. tobeloensis (prep. nr. BMNH1325373), habitus; 34, , paratype, habitus; 35, , paratype, habitus; 36, , holotype, habitus. C. interrupta C. continua C. angustaDownloaded from Brill.com10/06/2021 06:56:51AM via free access

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Figs 37–40. Adults of Cyphura spp. — 37, C. phantasmah, male, lectotype; 38, C. mastrigti, male, holotype; 39, C. mastrigti, female, paratype (coll. nr. BMNH(E)1325326); 40, C. astrolabensis, female, holotype.

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Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 51

Female genitalia (Fig. 32). (prep. nr. BMNH Forelegs: femur brown in front, tibiae brown in 1325346). Corpus bursae globular, slightly oblong. front, other parts and tarsi white. Other legs white, Length corpus bursae is about 80% of length ductus tarsi pale brown. bursae. No signum on corpus bursae. Antrum short, Forewing length 38 mm. Forewing white, with small sclerotized papillae anales with some setae. broad dark brown marginal band, slightly broad­ ening towards apex. Costa with dark brown margin, Distribution slightly broadening towards apex and slightly ­striated Cyphura latimarginata is restricted to New Guinea with black at the basal part. The angle between and not as widespread as C. caudiferaria. In Indo- the marginal band and the margin along the costa nesia it is found in places along the coast of Papua is almost straight and not rounded. Fringes dark Barat and the Birdshead Penninsula and on islands brown. near the coast of Papua Barat. It is also found at Hindwing white with a broad brown submarginal higher altitudes in the Cyclops Mountains and in band, starting from the anal angle, isolating a rather the Mamberamo district. In Papua New Guinea the narrow white margin at the marginal band. The sub- distribution is limited to some coastal areas of the marginal band is narrow near the tornus, while along mainland, such as Kaiser Wilhelmsland, Astrolabe the marginal band of the hindwing it is broader and Bay and Stephansort. of equal width. A small tail near the tornus. Three black dots near the tornus. The black dot in the tail is Diagnosis rounded and does not touch the submarginal band, Both C. caudiferaria and C. latimarginata are while the black dot in the middle is triangular and widespread over Papua Indonesia and Papua New touches the submarginal band. The black dot near Guinea; the most important difference is found on the anal angle is hardly visible, elongated and em- the the upperside of the hindwing: in C. caudifer- bedded in the submarginal band. Fringes dark brown aria the dark submarginal band isolates a rather and distally pale, on the upperside of the tail white, broad white margin, whereas in C. latimarginata near the black dots black. this white margin is very narrow. In the male geni- Underside of forewing identical to the upperside, talia the apex of the harpe reaches the edge of the but the dark marginal band is broader. The dark valve in C. caudiferaria, whereas in C. latimarginata margin along the costa somewhat fainter towards the harpe ends well before the edge of the valve. In base, not striated with black. C. caudiferaria the coecum is longer and the apex of Underside of hindwing identical to the upperside, the phallus more slender and stronger hooked than but the submarginal band is broader, isolating a nar- in C. latimarginata. row white margin along the marginal band, and also broader near the tornus. The three black spots are all Cyphura clarissima (Butler, 1879) embedded in the submarginal band. Figs 5, 19, 20 Abdomen — Dorsally and ventrally white. Strophidia clarissima Butler, 1879: 166. Holo- Female not known. type (fixed by monotypy) ♂, Papua New Guinea: Male genitalia (Figs 19, 20) (prep. nr. BMNH- New RV1582). Valves (Fig. 19) rounded, strongly haired, Ireland, leg. G. Brown (NHMUK, coll. nr. hooked at the apex. Harpe slender, not reaching BMNH(E)1325349) [examined] the edge. Very small thorn at the apex of the harpe. Note: The type specimen is labeled as “New Guin- Sacculus narrow, not bent. ea”, but Butler (1879) refers to “New Ireland” as the Phallus (Fig. 20) almost straight, tapering towards origin of the type specimen. apex, hooked at the apex. Small sclerotized tooth close to the apex. Coecum short and broad. Additional material examined 1♂, “New Guinea”, leg. C.J. Brooks B.M. 1936-681 Distribution (NHMUK, prep. nr. BMNH-RV1582) [examined] Papua New Guinea: New Ireland.

Description (Fig. 5) Diagnosis Male. Head — Vertex brown, frons white with Cyphura clarissima differs from both C. caudiferaria small black spot on the the upperside. Antennae and C. latimarginata in the narrower dark margin on pale brown, filiform, length 13 mm. Labial palpus: the upperside of forewing and the narrower dark sub- dorsally white with black at the base and pale brown marginal band on the upperside of hindwing. Near at the apex, ventrally white. the anal angle, this submarginal band is hardly vis- Thorax — Thorax white, patagia brown, tegulae ible. In C. clarissima the black dots on the ­hindwing white. Thorax ventrally white. are clearly isolated from the submarginal band,

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52 Tijdschrift voor Entomologie, volume 161, 2018 whereas in C. latimarginata the dots are embedded Forewing length 31 mm. Forewing white, a trace in this band, and in C. caudiferaria the dot between of a pale brown marginal band near the costa. Basal tornus and anal angle touches the submarginal band. half of the costa striated with pale brown and dark brown. Fringes dark brown. Cyphura tobeloensis sp. n. Hindwing white with a narrow pale brown sub- Figs 6, 7, 21, 22, 33 marginal band of equal width, starting from the Holotype: ♂, Indonesia, Moluccas, Halmahera, To- apex, near the tornus bending to the anal angle. belo (NHMUK, bought from E. le Moult, coll. nr. The submarginal band isolates a broad white margin BMNH(E)1325374, prep. nr. BMNH1325374) at the marginal band. A small tail near the tornus. Paratype: 1♀, same as holotype (coll. nr. Three black dots near the tornus, the one in the tail BMNH(E)1325373, prep. nr. BMNH(E)1325373); rounded, the one in the middle is triangular. None paratypes: 7♂♂, 1 ♀, same as holotype (NHMUK) of these are embedded in the submarginal band. The black dot near the anal angle is very small and hardly Description (Figs 6, 7) visible and included in the submarginal band. Fring- Male. Head — Vertex dark brown, frons white with es pale brown and distally white between the apex two small dark brown spots on the upperside. Anten- and tornus, on the upperside of the tail the fringes nae pale brown, filiform, length 12 mm. Labial pal- are white. Between tornus and anal angle the fringes pus: dorsally white with black at the base and dark are white, but near the black spots they are black and brown at the apex, ventrally pale white. distally white. Thorax — Thorax white, patagia white, tegulae Underside of forewing like male, but margins very white. Thorax ventrally white. pale brown. Forelegs femur pale brown in front, tibiae pale Underside of hindwing identical to the upperside, brown, other parts and tarsi white. Other legs white, but the submarginal band is very pale and inter- including tibiae and tarsi. rupted between apex and tornus. The submarginal Forewing length 31.0 mm. Forewing white, with band is broader between the tornus and anal angle, broad pale brown marginal band, slightly broaden- the three spots are small and pale brown, and only ing and fading towards apex. Costa with pale brown the spot in the tail is not embedded in the submar- margin, only present at the middle of the costa and ginal band. near the apex. The basal part of this margin is stri- Male genitalia (Figs 21, 22) (prep. nr. BMNH(E) ated with black. Fringes dark brown. 1325374). Valves (Fig. 21) almost rectangular, strongly Hindwing white with a pale brown submarginal haired, slightly hooked at the apex. band of equal width, starting from the apex. Near Harpe slender, slightly bent, obliquely crossing the tornus, it bends to the anal angle and narrows. the valve, reaching the edge of the valve. Small thorn The submarginal band isolates a broad white margin at the apex of the harpe. Sacculus pronounced, bent at the marginal band. A small tail near the tornus. in its middle. Three black dots near the tornus, the one in the tail Phallus (Fig. 22) strongly bent in the middle and rounded, the one in the middle triangular. Both are near the apex while strongly tapering towards apex, not embedded in the submarginal band. Black dot giving the impression of the beak of a duck. A small near the anal angle small, elongated and included in tooth close to the apex. Coecum short and narrow. the submarginal band. Fringes brown and distally Female genitalia (Fig. 33) (prep. nr. BMNH(E) pale brown near the apex, on the marginal band and 1325373). Corpus bursae oblong, its length about on the upperside of the tail the fringes are white. twice the ductus bursae, and small sclerotized papil- Between tornus and anal angle, the fringes are black lae anales with some setae. No signum on corpus and distally white. bursae. Underside of forewing identical to the upperside, but the brown margin does not fade towards the Distribution apex. The costa is not striated with black. There is Indonesia: Halmahera (Moluccas). no brown margin along the costa, only a pale brown spot at the middle of the costa. Underside of hindwing identical to the upperside, Etymology but the submarginal band is broad near the tornus, The name refers to the origin of the holotype, the the three spots are dark brown and all embedded in village of Tobelo (Halmahera). the submarginal band. Abdomen — Dorsally and ventrally white. Cyphura angusta sp. n. Female. Like male, but patagia dark brown, vertex Figs 8, 36 pale brown, frons white with black spot on the upper Holotype: ♀, Indonesia, Biak, 16.v.1954, ex coll. half. All tarsi brown. Veldhuyzen (RMNH, prep. nr. RMNH-RV1500) Downloaded from Brill.com10/06/2021 06:56:51AM via free access

Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 53

Description (Fig 8) forewing and the narrow brown submarginal band Female. Head — Vertex white, frons white with on the hindwing. black spot on the the upperside. Antennae filiform, pale brown, length 12 mm. Labial palpus: dorsally Cyphura interrupta sp. n. white with black at the base and dark brown at the Figs 9, 10, 23, 24, 34 apex, ventrally pale white. Holotype: ♂, Papua New Guinea, New Ireland, Thorax — Thorax white, patagia brown, tegulae xi.1923, leg. A.F. Eichhorn (NHMUK, coll. nr. white. Thorax ventrally white. BMNH(E)1325352, prep. nr. BMNH1325352) Forelegs femur pale brown in front, tibiae pale Paratype: 1♀, Papua New Guinea, New Ireland, brown in front, tarsi pale brown. Other legs white, xi.1923, leg. A.F. Eichhorn (NHMUK, coll. nr. but tarsi pale brown. BMNH(E)1325353, prep. nr. BMNH1325353) Forewing length 37 mm. Forewing white, with narrow pale brown marginal band of equal width. Costa with diffuse brown margin, slightly broaden- Description (Figs 9, 10) ing towards apex. The basal half of the costa is stri- Male. Head — Vertex white, frons white with black ated with dark brown. Fringes dark brown. spot. This black spot on the upperside invaded with Hindwing white with a narrow pale brown sub- white in the middle. Antennae filiform, dorsally pale marginal band, starting from the apex and of equal brown, ventrally white and checkered dark brown, width. Near the tornus the submarginal band is length 13 mm. Labial palpus: dorsally white with bent towards the anal angle and the part between black at the base and dark brown at the apex, ven- the tornus and anal angle is narrower. The submar- trally pale white. ginal band isolates a broad white margin at the mar- Thorax — Thorax white, patagia brown, tegulae ginal band. A small tail near the tornus. Two black white. Thorax ventrally white. dots near the tornus, the one in the tail rounded, Forelegs femur pale brown in front, tibiae pale the other one elongated. The black spot in the tail brown in front, other parts and tarsi white. Other is free from the submarginal band, while the other legs white, including white tibiae and tarsi. black spot touches the submarginal band. On the Forewing length 35 mm. Forewing slightly fal- marginal band, the fringes brown and distally pale cate, white, with narrow dark brown marginal band brown, while on the upperside of the tail the fringes of equal width. Along the veins marginal band are white. Between tornus and anal angle the fringes slightly invaded with white. Costa with a brown sub- are black and distally brown. costal band, isolating a narrow white margin along Underside of forewing identical to the upperside, the costa, which is covered with some dark brown but the pale brown marginal band broadens towards scales. The basal half of the costa is striated with the apex, while the pale brown margin along the cos- black. Fringes dark brown. ta is fading at the base and in the middle of the costa. Hindwing white with a dark brown submarginal The costa is not striated with dark brown. band, starting from the apex and slightly broaden- Underside of hindwing identical to the upperside, ing towards the tornus. Between the tornus and the but the submarginal band between the apex and tor- anal angle the submarginal band is interrupted, near nus is reduced to some pale brown streaks. The pale the anal angle it is continued, but very narrow and brown submarginal band is solid near the anal angle. almost black. The submarginal band isolates a broad The two spots are dark brown. white margin at the marginal band. A small tail near Abdomen — Dorsally and ventrally white. the tornus. Two black dots near the tornus, the one Male not known. in the tail rounded, the other one is triangular. Both Female genitalia (Fig. 36) (prep. nr. RMNH- are free from the submarginal band. The part of the RV1500). Corpus bursae oblong, length ductus submarginal band near the anal angle gives the im- bursae about 1.5 times as long as corpus bursae, and pression of a third, elongated black dot. On the mar- small sclerotized papillae anales with some setae. No ginal band the fringes dark brown and distally pale signum on corpus bursae. brown, while on the upperside of the tail the fringes are white. Between tornus and anal angle the fringes Distribution are black distally with brown. Underside of forewing with brown margin, slight- Only known from Biak Island in Cenderawasih Bay, ly broadening towards apex. Near the tornus the Papua, Indonesia. brown margin is broader until CuA1. There is a very narrow brown margin along the costa, near the base Etymology consisting of a few brown dots. Near the apex the The name angusta [Latin for narrow, thin], an adjec- brown margin is broad, starting at one third of the tive, refers to the narrow outer brown margin on the costa from the apex. Downloaded from Brill.com10/06/2021 06:56:51AM via free access

54 Tijdschrift voor Entomologie, volume 161, 2018

Underside of hindwing identical to the upperside, Thorax – Thorax white, patagia brown, tegulae but the narrow submarginal band extends to a large white. Thorax ventrally white. triangular field near the tornus. The spots are dark Legs brown, including tibiae and tarsi. brown and all embedded in the submarginal band. Forewing length 31 mm. Forewing slightly fal- Between the dark brown dots, two small white spots cate, white, with a broad dark brown marginal band, are isolated between the tornus and the anal angle. along the veins slightly invaded with white. Near the Abdomen — Dorsally and ventrally white. apex, only a few scattered dark brown scales. Costa Female. Identical to male, but patagia dark brown, with dark brown subcostal band, near the apex only vertex pale brown, frons white with black spot on the a few scattered brown scales. The basal half of the upper half. All tarsi brown. costa is striated with black. Fringes dark brown. Forewing length 36 mm. Forewing like the male, Hindwing white with a broad dark brown sub- but the margin and costa are paler brown. Hindwing marginal band of equal width, starting from the like the male, but submarginal band slightly broader apex. Near the tornus the submarginal band is bent and paler brown. towards the anal angle. The submarginal band iso- Underside of forewing identical to the male, but lates a broad white margin at the marginal band. A brown marginal band not broadened near the tor- small tail near the tornus. Two rounded black dots nus, with a trace of a pale brown subcostal band near near the tornus, the one in the tail free from the sub- the middle of the costa. marginal band, the upper half of the second one em- Underside of hindwing identical to the male. bedded in the submarginal band. On the marginal Male genitalia (Figs 23, 24). Valves (Fig. 23) broad, band the fringes dark brown and distally pale brown, not rounded, strongly hairy, hooked at the apex. while white on the upperside of the tail. Between Harpe stout, almost straight, obliquely crossing tornus and anal angle, the fringes are black, distally the valve, not reaching the edge of the valve. Thorn brown. at the apex of the harpe. Sacculus stout, bent at one Underside of forewing with broad dark brown third from the base. margin, slightly broadening towards apex. Near the Phallus (Fig. 24) bent, tapering in direction of tornus the brown margin is broader until CuA1. apex. A small sclerotized tooth close to the apex. There is a very narrow brown margin along the costa, Coecum short and narrow. consisting of a few brown dots. A trace of a brown Female genitalia (Fig. 34). Corpus bursae oblong, subcostal band near the middle of the costa. Near length corpus bursae 1.6 times as long as ductus bur- the apex, the brown margin is broad, starting at one sae, and small sclerotized papillae anales with many third of the costa from the apex. setae. No signum on corpus bursae. Underside of hindwing identical to the upper- side, but the broad dark brown submarginal band is Distribution extended to a large triangular field near the tornus. Papua New Guinea: New Ireland, New Hanover. The veins are visible as white lines in the submar- ginal band. The spots are dark brown and are all embedded in the submarginal band. Between the Etymology dark brown dots, two small white spots are isolated The name interrupta, an adjective, refers to the inter- between the tornus and the anal angle. rupted submarginal band between tornus and anal Abdomen — Dorsally and ventrally white. angle on the hindwing. Female. Identical to male, but forewing length 34 mm. The margins of the forewing are brown, the Cyphura continua sp. n. costa more striated with brown. On the hindwing, Figs 11, 12, 25, 26, 35 the submarginal band is slightly broader. Holotype: ♂, Papua New Guinea, Talasea, New Brit- Underside identical to male, but margins and sub- ain, iii-iv.1925, leg. A.F. Eichhorn (NHMUK, prep. marginal band brown. nr. BMNH-RV1584) Male genitalia (Figs 25, 26) (prep. nr. BMNH- Paratype: 1♀, Papua New Guinea, Talasea, New Brit- RV1584). Valves (Fig. 25) broad, not rounded, ain, ii.1925, leg. A.F. Eichhorn (NHMUK, prep. nr. strongly hairy, hardly hooked at the apex. Harpe BMNH-RV1585) stout, almost straight, obliquely crossing the valve, reaching the edge of the valve. Thorn at the apex of Description (Figs 11, 12) the harpe. Sacculus stout, slightly bent at one third Male. Head — Vertex white, frons white with from the base. black spot on the upperside. Antennae: length 11 Phallus (Fig. 26) bent, near the apex strongly mm, filiform, pale brown. Labial palpus: dorsally curved, hook-like, giving the impression of the beak white with black at the base and dark brown at the of a bird of prey. A small sclerotized tooth close to apex, ventrally pale white. the apex. Coecum short and narrow. Downloaded from Brill.com10/06/2021 06:56:51AM via free access

Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 55

Female genitalia (Fig. 35) (prep. nr. BMNH- apex, striated with black at the basal part. The angle RV1585). Corpus bursae oblong, length corpus between the marginal band and the margin along the bursae about twice the length of ductus bursae, and costa is slightly rounded. Fringes black. small sclerotized papillae anales with many setae. No Hindwing white with a dark brown submarginal signum on corpus bursae. band, starting at the apex, bent slightly outwards in the direction of the costa. Near the costa the submar- Distribution ginal band is more narrow and bends towards the Papua New Guinea: New Britain: Talasea, Umboi anal angle. The submarginal band isolates a white Island. margin at the marginal band. A small tail near the tornus. Two black dots near the tornus, the one in the tail rounded, the other dot is triangular. The Etymology black dot in the tail does not touch the submarginal The name continua [Latin for continuous], an ad- band. The submarginal band is almost interrupted jective, refers to the submarginal band on the hind- near the position of the second black dot, leaving wing, which is not interrupted between tornus and this black dot also free from the submarginal band. anal angle. Fringes dark brown and distally pale, while on the upperside of the tail the fringes are white. Between Diagnosis tornus and anal angle, the fringes are black and dis- Cyphura interrupta and C. continua are rather similar. tally pale brown. The main difference can be found in the ­submarginal Underside of forewing identical to the upperside, band on the hindwing’s upperside: in C. interrupta, but the costa with a brown margin, not striated with this band is interrupted between the tornus and the black. anal angle, leaving the black spot in the middle iso- Underside of hindwing identical to the upperside, lated from the submarginal band. In C. continua, the but the submarginal band is extended to a large tri- submarginal band is not interrupted and the black angular field near the tornus. On the outside, the spot in the middle is embedded in this band. The submarginal band is invaded with white at the veins. margin of the forewing and the submarginal band The two spots near the tornus are dark brown and on the hindwing are slightly broader in C. continua both embedded in the submarginal band. Between than in C. interrupta. In the male genitalia, the valve the dark brown dots, two small white spots are iso- of C. continua is more strongly hooked at the apex lated between the tornus and the anal angle. than in C. interrupta. By contrast, the apex of the Abdomen — Dorsally and ventrally white. phallus is more strongly hooked in C. interrupta than Female not known. in C. continua. Male genitalia (Figs 27, 28). Valves (Fig. 27) nar- row, rounded, strongly hairy, hooked at the apex. Cyphura laeensis sp. n. Harpe stout, almost straight, obliquely crossing the Figs 13, 27, 28 valve, not reaching the edge of the valve. Small thorn Holotype: ♂, Papua New Guinea, Lae (PNG), at the apex of the harpe. Sacculus stout, bent in the 13-xii-1967, Coode 88 (RMNH, prep. nr. middle. RMNH- RV1501) Phallus (Fig. 28) slightly bent, near the apex strongly hooked and at the apex slightly bending up- Description (Fig. 13) wards. A small sclerotized tooth close to the apex. Male. Head — Vertex brown, frons white with nar- Coecum broad and short. row black margin on the upperside. Antennae fili- form, pale brown, length 13 mm. Labial palpus: dor- Distribution sally white with black at the base and dark brown at Only known from Papua New Guinea: Lae, on the the apex, ventrally pale white. northern coast. Thorax — Thorax white, patagia brown, tegulae white. Thorax ventrally white with pale brown streak in the middle. Etymology Forelegs femur pale brown in front, tibiae pale The name laeensis refers to the origin of the holo- brown in front, tarsi pale brown. Other legs white, type, the village of Lae (PNG). but tarsi pale brown. Forewing length 37 mm. Forewing white, with Diagnosis broad dark brown marginal band, broadening to- Cyphura laeensis is rather similar to C. caudifer- wards apex. Width of the brown margin near the apex aria, but the dark submarginal band on the up- is about one third of forewing length. Costa with perside of the hindwing is more narrow and the dark brown subcostal margin, broadening towards black spot between tornus and anal angle is clearly Downloaded from Brill.com10/06/2021 06:56:51AM via free access

56 Tijdschrift voor Entomologie, volume 161, 2018 isolated from the submarginal band. in C. caudifer- costa. Slightly hooked at the apex. Harpe strongly aria the submarginal band is broader and the black haired, obliquely crossing the valve, not reaching spot between ­tornus and anal angle is not isolated the edge of the valve. Small thorn at the apex of the from the submarginal band. In the male genitalia of harpe. Sacculus narrow, slightly bent. C. laeensis, the phallus is hooked and slightly bent up Phallus (Fig. 30) tubular, slightly bent, tapering at the apex, whereas in C. caudiferaria the apex is also towards apex. Apex strongly curved, hook-like, very hooked but not bent up. slender, underside with small sclerotized tooth close to the apex. Coecum rather long, broad. Cyphura dilatata sp. n. Figs 14, 29, 30 Distribution Holotype: ♂, Papua New Guinea, Manki Ridge, Bu- The only specimen is from Papua New Guinea, lolo, 1700 m, 6.v.1956, I.F.T.A. coll., ex coll. Ernst Manki Ridge, Bulolo. Jugl (ZMHB, prep. nr. ZMHB-RV1566)

Description (Fig 14) Etymology Male. Head — Vertex white with black dot, frons The name dilatata [= extended or broadened] re- white with large black rectangular spot. Antennae fers to the broad dark brown marginal band on the pale brown, filiform, length 13 mm. Labial palpus: forewing and the broad submarginal band on the dorsally white with black at the base and dark brown hindwing. at the apex, ventrally pale white. Thorax — Thorax white, patagia pale brown, te- Discussion gulae white. Thorax ventrally white. Cyphura dilatata is rather similar to C. caudiferaria, Forelegs femur brown in front, tibiae brown in but on the forewing the dark brown marginal band front, other parts and tarsi white. Other legs white, strongly broadens towards the apex, resulting in the including tibiae and tarsi. impression that the whole apical field is dark brown. Forewing length 34 mm. Forewing white, with In C. caudiferaria, the dark brown marginal band very broad dark brown marginal band, strongly only slightly broadens to the apex. broadening towards apex. This dark brown margin meets the margin at the costa at 45% of length of costa from apex. Costa with broad dark brown mar- The phantasmah-group gin, broadening towards apex and striated with black and white, except for the part near the apex. Fringes Cyphura phantasmah ([C. & ? R.] Felder & Rog- black. enhofer, 1875) Hindwing white with a dark brown submarginal Figs 37, 41–43 band of equal width, starting from the apex, bent Strophidia phantasmah [C. & ? R.] Felder & Rogen- slightly outward, near the tornus bending to the anal hofer (1864–1875: Tf. 128 fig. 40). Lectotype angle and narrower. The submarginal band isolates (designated here): ♂, [Indonesia, Moluccas,] Ge- a white margin at the marginal band. A small tail beh (NHMUK, coll. nr. BMNH(E)1325319) near the tornus. Three black dots near the tornus, [examined] the one in the tail rounded and not embedded in the submarginal band. The black dot in the middle is smaller, elongated and touches the submarginal Additional material examined band. The black dot near the anal angle is elongated Paralectotypes: 1♂, [Indonesia, Moluccas,] Gebeh, and included in the submarginal band. A trace of Moore Collection 94-106 (NHMUK, coll. nr. brown just above the black dot near the anal angle. BMNH(E)1325327, prep. nr. BMNH1325327) Fringes black and distally pale brown, while on the [examined]; 3♂, [Indonesia, Moluccas,] Gebeh, leg. upperside of the tail the fringes are white. Bernstein (RMNH) [examined] Underside of forewing identical to the upperside, Note: in the Global Lepidoptera Names Index the but the costa not striated. name has been changed to C. phantasma; the only Underside of hindwing identical to the upperside, text available is the caption of the original figure and but the submarginal band is broad near the tornus we see no reason to change this name. and anal angle, and the three black spots are embed- ded in the submarginal band. Description (Fig. 37) Abdomen — Dorsally and ventrally white. Male. Head — Vertex pale brown, frons white with Female not known. small black round spot on the upperside. ­Antennae Male genitalia (Figs 29, 30). Valves (Fig. 29) broad, length 12 mm, pale brown, filiform.­Labial palp rounded, slightly hairy, but almost no hairs along the dorsally black with black apex, ventrally white. Downloaded from Brill.com10/06/2021 06:56:51AM via free access

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Thorax — Thorax white, patagia and tegulae Holotype: ♂, Indonesia, Suer, Numfor Island, v/ white. Thorax ventrally white. vi-1879, leg. W. Doherty (NHMUK, coll. nr. Forelegs dark brown, tibiae and tarsi pale brown. BMNH(E)1325325, prep. nr. BMNH1325325) The other legs, including tibiae and tarsi pale brown. Paratypes: 1♀, same as holotype (NHMUK, coll. nr. Forewing length 38.0 mm. Forewing white, with BMNH(E)1325326, prep. nr. BMNH1325326); very narrow pale brown margin, starting from tornus 7♂, 5♀, same as holotype (NHMUK); 1♂, 1♀, until one fourth of marginal band from apex. This Namber, Numfor Island, leg. H. van Mastrigt, 14– pale brown margin is bordered on the inner side by 17.vii.2005 (KSP, coll. nr. KSP30343 (male) and a very narrow brown submarginal band. Along the KSP30346 (female)). costa, a brown margin, speckled with white, almost reaching the apex. This margin is dark brown at one Description (Figs 38, 39, 44–46, 47) third from the basal part of the costa, while the other Male. Head — Vertex brown, frons white with up- part is pale brown. Fringes dark brown. per part black. Antennae length 9 mm, pale brown, Hindwing white with a tail at the tornus and a filiform. Labial palpi dorsally black with white apex, narrow brown submarginal band, isolating a narrow ventrally white. white margin at the marginal band. The submargin- Thorax — Thorax white, patagia and tegulae al band broadens and fades towards the anal angle, white. Thorax ventrally white. which it reaches. Two black dots near the tornus, All legs, including tibiae and tarsi, white. the one in the tail not embedded in the submargin- Forewing length 30.0 mm. Forewing white. Along al band, while the other is embedded. The spot in the costa, a brown margin, speckled with white, al- the tail is round, the other spot triangular. Fringes most reaching the apex. The margin is broadened brown, white at the upperside of the tail and black from the basal part to the middle of the costa, then near the black dots . fades towards the apex. The margin is dark brown at Underside of forewing white, with pale brown the basal half of the costa, the other part is brown. margin, slightly broadening at the apex. No margin Fringes brown. along the costa. Hindwing white with a very small tail at the tor- On the underside of the hindwing, the pale brown nus and a very faint brown submarginal band, only submarginal band is indistinct and interrupted with visible near the apex along one third of the mar- white at the upperside of the tail. Neither of the dots ginal band. The submarginal band is also visible are embedded in the submarginal band. near the anal angle. A trace of a black dot near the Abdomen — Dorsally and ventrally white. tornus. Fringes white, black at the underside of the Male genitalia (Figs 41–43) (prep. nr. BMNH(E) tail. 1325327). Valves (Fig. 41) rather narrow, right- Underside of forewing white, apex with a pale angled at the end of sacculus, slightly hooked at brown margin, starting at one third from the apex at the end of the costa, strongly hairy. Harpe narrow, the marginal band and ending at one sixth from the broadened at the base, rather long thorn at the apex apex on the costa. of harpe and harpe not reaching the edge of the Underside of hindwing white. valve. Sacculus rather broad and bent in the middle. Abdomen — Dorsally and ventrally white. Hairs on sacculus and outer edge of cucullus directed Female. Like the male, but antennae filiform, length towards base. 13 mm. Forewing length 33.0 mm. On the forewing Phallus (Figs 42, 43) strongly tapering towards there is a faint pale brown margin, fading in the di- apex, which is slender and curved. Small cornutus rection of the tornus, not reaching the tornus. At near apex. Coecum short and broad. the costa a pale brown margin, speckled with white. Female not known. Along the costa also a narrow dark brown margin, tapering towards the apex. The fringes are pale Distribution brown. Gebeh Island (Indonesia, Moluccas) Hindwing white with a tail at the tornus and a narrow brown submarginal band, isolating a white margin at the marginal band. The submarginal band Discussion is fading towards the tornus, not visible near the tail, Unfortunately in the NHMUK and RMNH col- but a trace of the submarginal band near the anal lections, we only found males, all from the island angle. A dark brown small spot in the tail. Fringes Gebeh. Until now we could not identify potential brown, white at the upperside of the tail and dark female specimens from Gebeh. brown, distally white at the underside of the tail. Underside as the male, but on the hindwing a very Cyphura mastrigti sp. n. small brown spot in the tail and a trace of a brown Figs 38, 39, 44–47 submarginal band near the anal angle. Downloaded from Brill.com10/06/2021 06:56:51AM via free access

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Male genitalia (Figs 44–46). Valves (Fig. 44) rather Description (Fig. 40) broad, right-angled at the end of sacculus, slightly Female. Head — Vertex white, frons white with hooked at the end of the costa. Harpe slightly broad- small black bar on the upperside. Antennae length ened at the base, small thorn at the apex of harpe and 13 mm, pale brown, filiform. Labial palpi dorsally harpe not reaching the edge of the valve. Hairs on black with black apex, ventrally white. sacculus and outer edge of cucullus directed towards Thorax — Thorax white, patagia and tegulae base, with many hairs at base of sacculus. white. Thorax ventrally white. Phallus (Figs 45, 46) curved, slightly tapering to- Forelegs dark brown, tibiae and tarsi pale brown. wards apex, which is slightly curved. Cornutus near The other legs, including tibiae and tarsi pale brown. apex. Coecum short and broad. Forewing length 34.0 mm. Forewing white, near Female genitalia (Fig. 47) Corpus bursae short, the tornus a trace of a pale brown marginal band. club-shaped. Length corpus bursae about the same Along the basal part of the costa a pale brown mar- as length ductus bursae. No signum on corpus bur- gin, striated with pale brown. Fringes pale brown. sae. Antrum narrow. Papillae anales short, and setae Hindwing white with a tail at the tornus and a on papillae anales. very narrow pale brown submarginal band, which is hardly visible in some specimens, isolating a nar- Distribution row white margin at the marginal band. Two small Only known from Numfor Island (Indonesia, Papua) black dots near the tornus, which are hardly visible. Fringes white, between tornus and anal angle pale brown. Etymology Underside of forewing white, with a very pale This species is named in honor of Henk van Mas- brown margin on the basal part of the costa. trigt, who collected a large number of Underside of hindwing white without markings. and in Papua Indonesia. In his collection, we Abdomen — Dorsally and ventrally white. found two paratypes of Cyphura mastrigti sp. n. Female genitalia (Fig. 48) Corpus bursae short, club-shaped. Length corpus bursae is about the same Discussion as length ductus bursae. No signum on corpus bur- Cyphura phantasmah and C. mastrigti are rather sae. Antrum narrow. Papillae anales short, a few setae similar. C. phantasmah has a forewing length of on papillae anales. 38 mm, all 14 specimen of C. mastrigti in the NHMUK have a forewing length of about 30 mm. Distribution The brown margin at the costa is of equal width Papua New Guinea: Astrolabe Bay, Stephansort. in C. phantasmah, whereas in C. mastrigti it broad- ens towards the middle of the costa. In C. mastrigti there is no brown margin at the marginal band. On Etymology the hindwing, the brown submarginal band is very The name refers to the origin of the holotype, Astro- narrow (less than 1 mm) in C. mastrigti , whereas in labe Bay (PNG). C. phantasmah it is about 2 mm wide. In C. phan- tasmah there are usually two black dots between Discussion the tornus and the anal angle of the hindwing, in The members of the phantasmah-group all have C. mastrigti, there are never two black spots, and almost the same externals: white, very narrow pale in some specimens there is a trace of a small dark brown margins. On the hindwing the submarginal spot in the tail. band is hardly visible and the black dots are less pro- In the male genitalia, the phallus is more slender nounced than in the other members of this genus. and bent in C. phantasmah. The cornutus near the Unfortunately, we found only males of Cyphura apex is smaller in C. phantasmah and the distance to phantasmah in the collections of NHMUK and the apex is larger. RMNH, all from the small island Gebeh on the east side of the Moluccas. C. mastrigti, only known from Cyphura astrolabensis sp. n. the island Numfor, is smaller than C. phantasmah Figs 40, 48 and differs from it by the very small tail on the hind- Holotype: ♀, Papua New Guinea, Stephansort wing and the broad striated margin at the costa of (NHMUK, coll. nr. BMNH(E)1325347, prep. nr. the forewing. We only found five females ofC. astro- BMNH1325347) labensis in the collection of NHMUK; it is not clear Paratypes: 2♀, Papua New Guinea, Stephansort if there are males in this collection that belong to [NHMUK]; 2♀, Papua New Guinea, Astrolabe Bay this species; DNA-barcoding could reveal the other (NHMUK) sex in the future.

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Figs 41–48. Male genitalia (Figs 41–46) and female genitalia (Figs 47, 48) of Cyphura spp. in the phantasmah group. — 41, C. phantasmah (prep. nr. BMNH1325327) (after removal of phallus); 42, C. phantasmah, phallus; 43, C. phantasmah, apex of phallus; 44, C. mastrigti, holotype (after removal of phallus); 45, C. mastrigti, phallus; 46, C. mastrigti, apex of phallus; 47, C. mastrigti (prep. nr. BMNH1325326) habitus;Downloaded 48. from C. atrolabensisBrill.com10/06/2021, holotype, 06:56:51AM habitus. via free access

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The pardata-group Hindwing white with a yellow suffusion in the outer half of the wing in the region of the anal angle. Cyphura pardata Warren, 1906 A small tail near the tornus. Between tornus and anal Figs 49–54 angle five black triangular spots. Starting from the Cyphura pardata Warren (1906: 66). Lectotype (des- anal angle, a small spot, two round spots close to ignated here): ♂, Papua New Guinea, Angabunga each other and two larger triangular spots near the River, affl. of St. Joseph River, 6000 ft upwards, tornus. A black submarginal stripe between vein two x-1904–ii.1905, leg. A.S. Meek (NHMUK, coll.nr. and vein five. A short black margin just above the BMNH(E)1325357) [examined] tail on the marginal band and a black margin in the neighborhood of the black dots between the tornus and the anal angle. Additional material examined Underside of forewing white, a dark brown mar- Paralectotypes: 1♂, same as lectotype (NHMUK, coll. gin with some small black dashes along the basal part nr. BMNH(E)1325358, prep. nr. BMNH1325358) of the costa. The three bands are visible on the hind- [examined]; 1♀, same as lectotype (NHMUK, coll. wing, the edges of the two outer bands suffused with Nr.BMNH(E)1325359, prep. nr. BMNH1325359) brown, mainly at the coastal part. The apical part [examined]. 1♀, Biagi, Mambare River, 5000ft, striated with brown. A dark brown margin at the i–iv.1906, leg. A.S. Meek (NHMUK, prep. nr. marginal band, starting at the apex and fading out in BMNH-RV1579) [examined]; 1♂, Indonesia the direction of the tornus. Fringes white, between (Papua): Hulu, Mamberamo, Foja Mts, 2°34.5′ S, tornus and anal angle yellow with black at the end. 138°42,9′ E, 23.xi–7.xii.2005, leg. H. van Mastrigt Underside of hindwing like the upperside, but the (KSP, coll.nr. KSP42844) [examined] black submarginal stripe reaches the anal angle and the three black dots near the anal angle embedded Description (Figs 49, 50) in this stripe. Male. Head —Vertex white, frons white with oval Abdomen — Dorsally and ventrally white. Dor- black spot in the middle. Antennae length 13 mm, sally a distinct black line, continued on the thorax. pale brown, filiform. Labial palpi dorsally black with Female. Like the male, but antennae filiform, length white apex, ventrally white. 14 mm. Thorax — Thorax creamy white, patagia and te- Forewing length 34 mm. gulae creamy white. A black line in the middle of the Hindwing like the male, but with a diffuse brown thorax, continued on the abdomen. Thorax ventrally streak, starting on the inner margin at one seventh white. from the anal angle, bent in the direction of the Forelegs dark brown in front, tibiae and tarsi middle of the costa and fading out in the middle of white. Other legs white, including tibiae and tarsi. the hindwing. Forewing length 30.0 mm. Forewing creamy Underside of hindwing like the upperside, but white. Along the basal part of the costa, short black the diffuse brown streak does not reach the inner dashes, reaching the subcostal vein. Three broad yel- margin. low-brown transverse bands, bordered with brown, Male genitalia (Figs 51, 52) (prep. nr. BMNH(E) starting from the inner margin and reaching the sub- 1325358). Valves (Fig. 51) square, not hooked at the costal vein. The inner edge of the basal band starts on apex of the costa. Harpe not reaching the edge of the inner margin at one fifth from the base and ends the valve, slightly broadening towards apex of harpe. at the base of the costa. The outer edge starts on the Thorn at apex of harpe of medium size. Sacculus inner margin at two third from the base and reaches narrow. the costa at one seventh from the base. The inner Phallus (Fig. 52) tubular, slender, slightly curved. edge of the middle band starts on the inner margin at Apex not pointed. Coecum very short, slender. one fifth from the tornus and reaches the costa at one Female genitalia (Figs 53, 54) (prep. nr. BMNH- third from the base. The outer edge of the middle RV1579). Corpus bursae (Fig. 53) globular, length band starts at the inner margin at one eight from corpus bursae about the same as length of ductus the tornus and reaches the costa at the middle of the bursae. One signum on the upper half of the corpus costa. The inner edge of the outer band starts at the bursae (Fig. 54), small, oblong, constricted and cov- inner margin at the tornus and reaches the costa at ered with a few rows of spines. Antrum small. Papil- one third from the apex. This outer band fills the lae anales with a few setae. apical part of the forewing. This apical part is stri- ated with black dashes, mainly in the costal part. A Distribution narrow black submarginal band along the marginal In Indonesia (Papua) only known from the Foja band, starting near the tornus and ending just before Mountains; in Papua New Guinea known from the apex. Fringes black. a few places on the mainland: Angabunga River, Downloaded from Brill.com10/06/2021 06:56:51AM via free access

Sinnema and Sinnema-Bloemen: Revision of the genus Cyphura Warren 61

Figs 49–54. Adults and genitalia of Cyphura pardata. — 49, male, lectotype; 50, female, paralectotype (coll. nr. BMNH(E)1325359); 51, male genitalia (after removal of phallus) (prep. nr. BMNH1325358); 52, male genitalia, phallus; 53, female genitalia, habitus (prep. nr. BMNH-RV1579); 54, female genitalia, signum.

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62 Tijdschrift voor Entomologie, volume 161, 2018

­Biagi (Mambare River), Bome, Matoko (Finisterre recueillis pendant le voyage. Première partie, Lépi- Range), Rawlinson Mts, Owgarra. doptères. — In: Voyage de découvertes de l’Astrolabe ­pendant les années 1826–1829. J.Tastu, Paris, France, Diagnosis pp 171–267. Butler, A.G., 1879. On the Heterocera in the collection Cyphura pardata differs from all other Cyphura spe- of Lepidoptera from New Ireland obtained by the Rev. cies by the yellow markings on both forewing and G. Brown. — Proceedings of the Zoological Society of hindwing; C. (X.) devosi also has yellowish hind- London 1879: 160–166. wings, but the forewings are black and white as in Felder, C., Felder, R. & Rogenhofer, A.F., 1864–1875. the other members of the subgenus Xysterophora Lepidoptera — In: Reise der Österreichischen Fre- (Sinnema & Sinnema-Bloemen 2016). gatte Novara um die Erde in den Jahren 1857, 1858, The genitalia show also a resemblance with Xys- 1859 unter den Befehlen des Commodore B. von terophora species: in the females there is a signum on Wüllerstorf-Urbair, Bnd. 2, abt. 2; Tf. 128 Figs 39, the corpus bursae as in all members of Xysterophora, 40. Kaiserlich-Königliche­ Hof- und Staatsdruckerei, but in other species of subgenus Cyphura the signum Vienna, Austria. on the corpus bursae is absent. In the male genita- Holloway, J.D., Kibby, G. & Peggie, D., 2001. The fami- lia of C. pardata the phallus is tubular, slender and lies of Malesian moths and butterflies. Fauna Malesiana not pointed, very similar to Xysterophora species. Handbook, Vol. 3. — Brill, Leiden, The Netherlands, The valvae are broad and the harpe is slender and 455 pp. does not reach the edge of the valve, very similar to Jordan, K., 1939. On the constancy and variability of the Xysterophora­ , but there is an important difference: differences between the Old World species of Utethei- the abdomen of the male of C. pardata does not sa (Lepid.; Arctiidae). — Novitates Zoologicae 41: show a trace of the “grater” on the third abdominal 251–291. segment, always present in the Xysterophora males. Kôda, N., 1987. A generic classification of the subfamily Arctiinae of the Palaearctic and Oriental regions based on the male and female genitalia (Lepidoptera, Arcti- Acknowledgements idae), Part I. — Tyô to Ga 38: 153–237. This revision would not have been possible without Lees, D.C. & Smith, N.G., 1991. Foodplant associations of the Uraniinae (Uraniidae) and their systematic, evo- the help and cooperation of the following persons lutionary, and ecological significance. – Journal of the and institutions. First of all we are indebted to Rob Lepidopterists’ Society 45: 296–347. de Vos (Naturalis Biodiversity Center, Leiden, The Robinson, G.S., 1976. The preparation of slides of Lepi- Netherlands) who inspired us to take a closer look doptera genitalia with special reference to the Micro- at and to revise the genus Cyphura and produced the lepidoptera. — Entomologist’s Gazette 27: 127–132. genitalia slides of all specimens we have described, Scoble, M.J., 1992. The Lepidoptera: Form, function and made the photographs of the genitalia and kindly diversity. — Oxford University Press, Oxford, UK, 404 commented on the manuscript. We thank the late pp. Henk van Mastrigt (Kelompok Serangga Papua, Jay- Sinnema, S. & Sinnema-Bloemen, J. 2016. Revision of the apura, Papua, Indonesia), Martin Honey (Natural genus Cyphura Warren, 1902. Part I: Introduction of the History Museum, London, United Kingdom) and new subgenus Xysterophora with two new species from Wolfram Mey (Museum für Naturkunde der Hum- New Guinea (Lepidoptera: Uraniidae: Uraniinae). —‘ boldt-Universität Berlin), who allowed us to study Tijdschrift voor Entomologie 159: 143–163. specimens from their collections, James Hogan, who Tuxen, S.L., 1970. Taxonomist’s glossary of genitalia in in- provided us with photographs of the type material sects. — Munksgaard, Copenhagen, Denmark, 359 pp. of the Oxford University Museum of Natural His- Swinhoe, C. 1902. New and little known species of tory, and Andreas Zwick, who gave us information Drepanulidae, Epiplemidae, Micronidae and Geome- about the collection of the Australian National In- tridae in the National Collection. —Transactions of the sect Collection CSIRO. The Uyttenboogaart-­Eliasen Entomological Society of London, 3: 585–677. Foundation (The Netherlands) supported the sur- Walker, F., 1866. List of the specimens of lepidopterous in the collection of the British Museum. Part veys to Papua, which enabled us to collect fresh ma- 35, supplement-part 5, Catalogue of Lepidoptera Het- terial and to study the KSP collection (Jayapura). erocera, seventh series, p 1919. We thank David Lees, Erik van Nieukerken (editor Warren, W., 1902. Drepanulidae, Thyridae, Uranidae, TvE) and an anonymous reviewer for reviewing the Epilpemidae and Geometridae from the Oriental manuscript. ­Region. — Novitates Zoologicae 9: 340–372. Warren, W., 1906. New Drepanulidae, Thyrididae, Ura- References niidae and Geometridae from British New Guinea. — Novitates Zoologicae 13: 61–161. Boisduval, J.A., 1832. Faune entomologique de l'Océan pacifique, avec l'illustration des insectes nouveaux Downloaded from Brill.com10/06/2021 06:56:51AM via free access