The Golden and Mariana albatrosses, new species of pierid butterflies, with a review of subgenus Appias (Catophaga) (Lepidoptera)

Authors


Abstract

This paper presents an overview of the subgenus Appias (Catophaga) Hübner (Pieridae). A beautiful golden-yellow member of the group, endemic to the Indonesian island of Sulawesi, Appias (Catophaga) aurosa Yata & Vane-Wright sp.n., is described as new. A small white species, Appias (Catophaga) mariana Yata & Chainey sp.n., is described as new from the Marianas. Four other taxa, A. (C.) athama (Blanchard), A. (C.) galba (Wallace) stat.rev., A. (C.) galene (Felder & Felder) and A. (C.) wardii (Moore), treated in most recent literature as subspecies, are recognized here as distinct, increasing the number of Catophaga species generally recognized from nine to 15. A brief review is given for each, with notes on their diagnosis, general distribution and known hostplants. An annotated synonymic checklist indicating subspecies, type localities and four new synonyms, and nine lectotype designations, one neotype designation and two type locality restrictions necessary to stabilize usage conclude the paper. Online Supporting Information provides an extensive discussion concerning the possible evolution of these butterflies with respect to polymorphism, speciation, coloration and hostplant relationships, a comprehensive list of type material for all available species group names belonging to the subgenus, and a complete bibliography for all citations in both the printed and online material.

Introduction

This work does not constitute a fully comprehensive revision. Its purpose is three-fold: to provide an overview of the species of subgenus Appias (Catophaga) Hübner, including the description of two new members of the group, and a key to adult males and females; to review known larval hostplant relationships, distribution and, especially, the spectacular polymorphism of certain species, with a view to stimulating further research into their ecology, genetics, molecular systematics and evolutionary biology; and to provide a fully synonymic list of all currently accepted subspecies to facilitate the future revisionary taxonomy that will probably be needed once extensive molecular investigations have been made into the numerous populations of these insects.

The butterflies assigned currently to the genus Appias Hübner comprise more than 40 species of ‘whites’ (Pieridae). Although found largely in the old world tropics, the genus is also represented in the Americas. Appias is placed, together with Saletara Distant (three or four species: Parsons, 1998; Vane-Wright & de Jong, 2003), Udaiana Distant (one species) and Aoa de Nicéville (one species), in the Appiadina Kusnezov, a subtribe of the Pierinae: Pierini (Braby et al., 2006).

Although the species of Appias were divided by Klots (1933: 208) into four subgenera, Yata (1981) proposed seven subdivisions, five of which were recognized as subgenera (Appias s.s., Catophaga Hübner, Phrissura Butler, Hiposcritia Geyer, Glutophrissa Butler, plus the lyncida and sylvia species groups). Braby et al. (2006: 263) suggested that the collective genus Appias, as accepted currently, is almost certainly a paraphyletic group (notably with respect to the exclusion of Saletara: Yata, 1981: 392).

Among these divisions, the Indo-Australian subgenus Catophaga Hübner is remarkable for the inclusion of several brightly coloured species in which the adult males, instead of the typical white or yellow of most Appias, are brilliant flame orange, chocolate brown or powder blue. The females of these butterflies are variously coloured, either white, yellow, polymorphic white or yellow, polymorphic white, yellow or male-like, or, in some cases, simply male-like with respect to their dominant colour. Catophaga is also notable because some, if not all, of its species seem capable of switching larval hosts between Capparaceae (a family of the Brassicales, now often placed within the Brassicaceae) and certain Malpighiales (e.g. Drypetes, family Putranjivaceae – formerly included in the Euphorbiaceae), apparently due to the common possession of glucosinolates in these plants (Braby & Trueman, 2006).

Methods

Materials

The greatest part of the material examined for this study is preserved in the collection of the Natural History Museum, London (BMNH), with significant studied material also in the Biosystematics Laboratory, Kyushu University, Fukuoka (BLKU). During this work, more than 8500 museum specimens were examined, and approximately 200 genitalia dissections prepared. In addition, research was undertaken on the extensive type material of these butterflies held in the BMNH collections to ensure that, as far as possible, the species group names applied are typified correctly and appropriate to employ.

Other material examined is located in the Entomological Laboratory, Faculty of Agriculture, Kyushu University (AGKU), Muséum National d’Histoire Naturelle, Paris (MNHN), Oxford University Museum of Natural History (OUMNH) and the Bishop Museum, Honolulu (BPBM).

Genitalia preparation and terminology

For the preparation of genitalia, either the entire abdomen or posterior half of the abdomen was removed, macerated in 10% aqueous KOH, and dissected in water using a binocular microscope. Except where noted, genitalia drawings were executed using a camera lucida from the entire genitalia or single parts submerged in a Petri dish of water, without any compression by glass slide and cover slip. For better contrast, some preparations were stained with Chlorazol Black. Terminology for male genitalia is based on Shirôzu's (1960: 1–10) extensive account, except that we use the term phallus instead of the more frequent ‘aedeagus’, as endorsed by Kristensen (2003). Terminology for female genitalia mainly follows van Son (1949), with some additions from Kusnezov (1915) and Yamauchi & Yata (2004).

Wing venation terminology

The Comstock–Needham wing-vein and cell nomenclature adopted in the descriptions is based on Nielsen & Common (1991) and Smith & Vane-Wright (2001). This terminology, together with the numerical system employed by Yata (1981) and many other lepidopterists (e.g. Corbet & Pendlebury, 1992), is illustrated in Fig. 1.

Figure 1.

Wing venation of Appias (Catophaga) paulina, showing both the Comstock–Needham terminology and the numerical system (small ciphers) for the long veins. The short cross-veins closing the discal are notated according to common lepidopterological practice: upper, middle and lower discocelluar veins (udc, mdc, ldc). The cells are notated using the Comstock–Needham system only. Dotted lines in the discal cells indicate ‘folds’ (probable courses of proximal parts of veins M1–M3), and in CuA2 the lost vein CuB (which supposedly appears during early development but is later resorbed). Based in part on Smith & Vane-Wright (2001: 513, fig. 7).

Taxonomic results Appias Hübner

Appias Hübner, 1819: 91. Type species by selection of Butler (1870a: 49): Papilio zelmira Stoll, 1780. [P. zelmira is considered to represent the same species group taxon as the older nominal species, Papilio libytheaFabricius, 1775; Appias (Appias) libythea occurs widely in the Oriental Region. Butler (1870a: 49) gave priority, as first reviser, to Appias Hübner over Catophaga Hübner.]

Subgenus Appias (Catophaga) Hübner

Catophaga Hübner, 1819: 93. Type species by selection of Scudder (1875: 136): Papilio paulinaCramer, 1777.

Trigonia Geyer, 1837: 21, 35. Type species by selection of Scudder (1875: 286): Papilio neroFabricius, 1793. (Invalid name: junior homonym of Trigonia Brugière, 1789.)

Tachyris Wallace, 1867: 361. Type species by selection of Scudder (1875: 274): Papilio neroFabricius, 1793.

Diagnosis

Habitus. Forewing apex generally acute, often sharply pointed, especially in male. Ground colour varies from white to yellow, orange, red, brown, blue and bluish-grey. Marginal and postdiscal band usually present, especially in female, but dark markings otherwise absent (at least in the basal half of the hindwing underside).

Venation. Forewing upper discocellular 1.5–2× length of middle discocellular; middle discocellular strongly curved; lower discocellular oblique (Corbet & Pendlebury, 1992). Forewing cell discal cell with three wing folds, but the most anterior does not reach the discocellular vein.

Male genitalia. Uncus long (the free part half the length of whole uncus, or longer), thickened dorsoventrally; valva broad, not strongly produced posteriorly, with an internal process (near the middle of ventral margin of costa + ampulla region); saccus short and oblong with rounded anterior tip; phallus strongly curved with anterovental end strongly developed and arched ventrally, almost the same length as coecum (coecumpenis). A well-developed black genital hair tuft on eighth abdominal sternite.

Female genitalia. Signum transverse, more or less pointed laterally, with many spines. Females also have a well-developed black genital hair tuft on the distal margin of the eighth sternite (Yata, 1981); this character is only known from one Appias species not included in subgenus Catophaga: Appias lalassis Grose-Smith (type species of Lade de Nicéville, 1898).

Three subgroupings of subgenus Appias (Catophaga)

The 15 species of subgenus Appias (Catophaga) recognized here are divisible into three groups, but we are uncertain if these all comprise monophyletic units. The paulina complex, which comprises eight closely related species in which the males are white, yellow or bluish, is represented throughout the entire Indo-Australian region (Fig. 23). The nero group, comprising three very closely related, nonoverlapping species in which the males are reddish-orange, is Oriental, extending from north-east India eastwards to the Philippines, Lombok and Buru (Central Maluku), and is entirely parapatric with respect to the third group, the melania series (four allopatric species in which the males are brown or bluish), confined to the Papuan subregion, including Australia (Fig. 24).

Figure 23.

Range map for all currently recognized species of the Appias (Catophaga) paulina complex.

Figure 24.

Range map for all currently recognized species of the Appias (Catophaga) nero group and the Appias (Catophaga) melania series.

paulina complex: galene, wardii, albina, aurosasp.n., athama, paulina, marianasp.n., mata

nero group: galba, nero, zarinda

melania series: placidia, clementina, celestina, melania

The paulina complex

Diagnosis. Male: upperside ground colour usually white, but rarely yellow or bluish-grey; forewing sometimes with a dark free spot in cell M3; uncus narrow and spindle- or spatula-shaped viewed dorsally, slightly swollen laterally, with its tip bluntly pointed; angle between vinculum and saccus approximately 100–120°; valve rounded posteriorly or weakly produced posteroventrally; phallus thick and strongly arched dorsally. Female: upperside ground colour usually white, but more rarely yellowish or bluish-grey (e.g. mata, paulina), or even almost black (e.g. one form of albina from Sulawesi); forewing with postdiscal black band strongly arched outwardly in cell CuA1, often forming a free dark spot in cell M3.

Distribution. (Fig. 23). This complex is represented throughout the Indo-Australian realm (including the western Pacific).

The nero group

Diagnosis. Male: upperside ground colour usually orange-red (rarely straw-coloured). Uncus narrow and nib-shaped viewed dorsally, gradually narrowed posteriorly, with its apex usually sharply pointed. Angle between vinculum and saccus approximately 90°; valve gradually narrowed posteriorly, distinctly produced posteroventrally, with distinct hooked spines present near apex; phallus thick but only weakly to moderately arched dorsally. Female: upperside ground colour usually dull orange-red (but polymorphic in Sulawesi and Palawan, where it has white, yellow and red forms, and without red forms in the rest of the Philippines), forewing black postdiscal band not strongly arched outwardly in cell CuA1, not forming a free dark spot in cell M3.

Distribution. (Fig. 24). This group of three parapatric species is widely distributed in the Oriental Region, from northern India to the Greater Sundas, Bali, the Philippines, Sulawesi and Buru (Central Maluku).

The melania series

Diagnosis. Male: upperside ground colour usually greyish-blue or dark brown, forewing never with a dark free spot in cell M3. Uncus somewhat broad and spatula-shaped viewed dorsally, slightly swollen laterally, with its tip bluntly pointed; angle between vinculum and saccus approximately 120°; valve evenly rounded posteriorly, distinctly incurved ventromedially; phallus slender and moderately arched dorsally. Female: forewing black postdiscal band not strongly arched outward in cell CuA1, not forming free dark spot in cell M3.

Distribution. (Fig. 24). The four allopatric species that make up this series are distributed from northern and central Maluku throughout much of the Papuan subregion, including north-eastern Australia.

Overview of species of the subgenus Appias (Catophaga) Appias (Catophaga) galene (Felder & Felder, 1865)

Sri Lanka albatross (Figs 2, 19D–F)

Figure 2.

Male genitalia of Appias (Catophaga) galene (Sri Lanka: NHM genitalia preparation no. 4669). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Figure 19.

Adult Appias (Catophaga) butterflies (halved: left hand upperside/right hand underside; all in BMNH unless otherwise stated). (A) A. aurosa Yata & Vane-Wright sp.n., male, whole upperside (Sulawesi: Macassar; #229202; forewing length 36.2 mm, holotype) (see also Fig. 20A); (B) A. mariana Yata & Chainey sp.n., male, whole upperside (Guam; #229237; forewing length 24.5 mm, holotype) (see also Fig. 20M); (C) A. mariana Yata & Chainey, sp.n., yellow female, whole upperside (Guam: Machanao; Bishop Museum; forewing length 25.5 mm, paratype) (see also Fig. 20P); (D) A. galene, male (Sri Lanka: Kandy; #665170; forewing length 30.0 mm); (E) A. galene, yellow female (Sri Lanka: #665171; forewing length 30.0 mm); (F) A. galene, white female (Sri Lanka: Newara Eliya; #665172; forewing length 28.3 mm); (G) A. wardii, male (southern India: North Kanara; #665191; forewing length 35.0 mm); (H) A. wardii, female (southern India: Nilgiris; #665174; forewing length 32.0 mm); (I) A. albina agatha, male (Philippines: Mindanao; #665175; forewing length 34.6 mm); (J) A. albina agatha, yellow female (Philippines: Mindanao, Davao; #665176; forewing length 27.8 mm); (K) A. albina agatha, white female (Philippines: Mindanao, Davao; #665177; forewing length 32.0 mm); (L) A. albina ambigua, female (East Timor: Dili; #665180; forewing length 28.0 mm); (M) A. albina ambigua, female (Indonesia: Lombok, Sapit; #135780; forewing length 26.3 mm); (N) A. albina ambigua, female (Indonesia: Lombok, Sapit; #665179; forewing length 27.8 mm); (O) A. albina albina, yellow female (Indonesia: Obi; #665178; forewing length 28.7 mm); (P) A. a. albina black female (Sulawesi: Palu; BLKU collection; forewing length 30.0 mm); (Q) A. athama manaia, male (Samoa: Upolu, Aleipata; #142269; forewing length 31.0 mm); (R) A. athama manaia, female (Samoa: Upolu, #142270; forewing length 29 mm); (S) A. athama wallacei, female (New Caledonia; #142268; forewing length 29.1 mm).

Pieris galene Felder & Felder, 1865: 165.

In several recent publications (e.g. Yata, 1981; D’Abrera, 1982, 1998), and for many years previously, A. galene has been regarded as a subspecies of A. paulina. However, Wynter-Blyth (1957: 430) (as ‘paulina’) and Kunte (2000: 101) treated it as a separate species, a status confirmed by the present study. Appias galene is endemic to Sri Lanka.

Diagnosis

Male. (Fig. 19D). Forewing somewhat acute at apex. Upperside ground colour white. Both wings usually without markings, but sometimes with a narrow black marginal border on forewing upperside. Hindwing with oval and bright whitish sex-patch posteromedially. Genitalia (Fig. 2): uncus very narrow and spindle-shaped viewed dorsally, with broadly blunt apex (also when viewed dorsally). Valva somewhat narrowed and rounded posteriorly, with a thick and arched interior process, directed dorsolaterally. Phallus somewhat short, thick and strongly arched dorsally, with rather broad coecum lacking a dorsal ridge; broadened basal prong as long as coecum.

Female. (Fig. 19E, F). Upperside ground colour white. Forewing black apical area usually with three submarginal whitish spots in cells R5–M2 (spot in cell M2 often reduced); black-dusted basal area with distal margin usually almost perpendicular to posterior margin of forewing. Underside hindwing yellow or white, with or without a dark submarginal band.

Distribution

Restricted to Sri Lanka. According to Woodhouse (1950) and d’Abrera (1998), a very common butterfly that frequently makes mass migrations across the island. Wynter-Blyth (1957: 430) stated that it occurs up to approximately 6000 ft (ca 1800 m); possibly absent in the far north of the island (H. Gaonkar, unpublished data).

Foodplants

Apparently unknown [Woodhouse, 1950: 153; d’Abrera, 1998; no entry in Robinson et al. (2001)].

Appias (Catophaga) wardii (Moore, 1884)

Ward's albatross (Figs 3, 19G, H)

Figure 3.

Male genitalia of Appias (Catophaga) wardii (South India: Nilgiri Hills; NHM genitalia preparation no. 4633). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Catophaga wardii Moore, 1884: 43.

In many recent accounts (e.g. Yata, 1981) and for many years earlier, wardii (like galene) was regarded as a subspecies of A. paulina. Bell (1913), who studied its life history, was uncertain regarding its separation from paulina (as leis), and referred to it as ‘leis-wardi’ [sic]. However, A. wardii was treated by Bingham (1907: 214), Talbot (1939: 406), Wynter-Blyth (1957: 431), Larsen (1987: 49) and by Kunte (2000: 101) as a distinct species, and this status has been confirmed during the present investigation. Appias wardii is endemic to the Western Ghats of southern India.

Diagnosis

Male. (Fig. 19G). Forewing somewhat obtuse at apex. Upperside ground colour white. Both wings always with distinct markings like the typical form of albina female (cf. Fig. 19K). Forewing black apical area usually with five submarginal light spots in cells R2 and R5–M3, those in M3 and CuA1 often reduced; black-dusted basal area with distal margin usually almost perpendicular to posterior margin of forewing, not oblique as in albina. Hindwing with oval and bright whitish sex-patch posteromedially. Genitalia (Fig. 3): uncus narrow and spindle-shaped viewed dorsally, apex blunt. Valva broad and rounded posteriorly, with a slender and almost straight interior process directed almost laterally. Phallus thick and strongly arched dorsally, with coecum lacking a dorsal ridge; basal prong as long as coecum.

Female. (Fig. 19H). General facies similar to male. Pale submarginal spots of forewing sometimes almost obsolete. Underside ground colour white.

Distribution

Restricted to western South India, from approximately 18°S to the far south, where it is fairly common along the Western Ghats. According to Bell (1913: 344) it is found from ‘sea-level up to 2500 or 3000 ft (ca 900 m). They are not found in the plain country being seemingly confined to the hill jungles where the rain is heavy’. Bell's statement is thus self-contradictory, at least regarding the lower altitudes at which this species can be found. Wynter-Blyth (1957: 431) states that it is ‘mainly confined to forest below 2500 ft'.

Foodplants

Putranjivaceae and Capparaceae. Bell (1912: 1145; 1913: 330, 341–344), although not stated directly, gives ‘Hemicyclia venusta’ (= Drypetes, Putranjivaceae) as the plant on which he bred this species. Talbot (1939: 408) notes Capparis heyneana.Robinson et al. (2001) list Capparis and Drypetes. Mathew & Binoy (2002) give Drypetes venusta.

Appias (Catophaga) albina (Boisduval, 1836)

White or common albatross (Figs 4, 19I–P)

Figure 4.

Male genitalia of Appias (Catophaga) albina (Indonesia: Sulawesi; NHM genitalia preparation no. 4686). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Pieris albina Boisduval, 1836: 480.

Diagnosis

Male. (Fig. 19I). Forewing distinctly acute at apex. Upperside ground colour white. No markings, but sometimes with a narrow black marginal border on forewing upperside. Hindwing with oval, bright, whitish sex-patch posteromedially. Genitalia (Fig. 4): uncus narrow and spindle-shaped viewed dorsally, apex blunt. Valva broad and rounded posteriorly, very different in outline to A. paulina (Yata, 1981: 377; Parsons, 1998: 292, fig. 48), with a thick and arched interior process, directed dorsolaterally. Phallus thick and strongly arched dorsally, with long coecum lacking a dorsal ridge; basal prong as long as coecum.

Female. (Fig. 19J–P). Forewing fairly acute at apex. Upperside ground colour typically white, but sometimes yellow, and in one form found on Sulawesi, infuscated, almost black (Fig. 19P). Forewing black apical area with at least four submarginal whitish spots in cells R5–M3, those in cells M3 and M2 usually reduced; black-dusted basal area with distal margin oblique.

Distribution

Widely distributed over much of the Indo-Australian Region, from Sri Lanka and southern India (Kunte, 2000: 101) to Taiwan, Ryukyu Islands (Takara, 1956: 67), Sundaland (including Siberut: Corbet, 1941), Wallacea, Lesser Sunda Islands (Rawlins, 2007), New Guinea and Kiriwina, and coastal areas of parts of Northern Territory and Queensland in Australia (Braby, 2000: 328; Braby et al., 2009). It does not occur in the Bismarcks (Parsons, 1998: 291) or Solomons (Tennent, 2002: 63), and appears to be replaced by A. athama in the western Pacific (New Caledonia to Samoa).

Foodplants

Capparis, Crateva (Capparaceae) and Drypetes (Putranjivaceae) (Corbet & Pendlebury, 1992; Parsons, 1998; Bascombe et al., 1999; Kunte, 2000, 2006; Robinson et al., 2001). Igarashi & Fukuda (2000: 394) record Drypetes litttoralis as the foodplant in Palawan, and D. poilanei in Laos, while noting an old record of Capparis heyneana for the Malay Peninsula. Braby et al. (2010) consider A. albina to be monophagous on Drypetes deplanchei in northern Australia.

Appias (Catophaga) aurosa Yata & Vane-Wright sp.n.

Golden albatross (Figs 5, 19A, 20A)

Figure 5.

Male genitalia of Appias (Catophaga) aurosa Yata & Vane-Wright sp.n. (Indonesia: south Sulawesi; NHM genitalia preparation no. 4597). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Figure 20.

Adult Appias (Catophaga) butterflies (halved: left hand upperside/right hand underside; all in BMNH unless otherwise stated). (A) A. aurosa, male (Sulawesi: Macassar; #229202; forewing length 36.2 mm) (see also Fig. 19A); (B) A. paulina galathea, male (India: Nicobar Is, Car Nicobar; #665185; forewing length 32.3 mm); (C) A. paulina galathea, yellow female (India: Nicobar Is, Car Nicobar; #665186; forewing length 32.0 mm); (D) A. paulina galathea, white female (India: Nicobar Is, Kondul; #665187; forewing length 31.5 mm); (E) A. paulina galathea, piebald female (India: Nicobar Is, Kondul; #665188; forewing length 31.7 mm); (F) A. paulina grisea, male (Malaysia: P. Tioman; #665183; forewing length 27.5 mm); (G) A. paulina grisea female (Malaysia: Pulau, Tenggol, Trengganu; #665184; forewing length 29.8 mm); (H) A. paulina sawela, male (Indonesia: Lombok, Sapit; #229105; forewing length 27.3 mm); (I) A. paulina sawela, female (syntype) (Indonesia: Lombok, Sapit; #149947; forewing length 29.0 mm); (J) A. paulina ega, male (Australia: Queensland, Cairns; #665181; forewing length 33.0 mm); (K) A. paulina ega, female (Australia: Queensland, Mackay; #665182; forewing length 30.5 mm); (L) A. paulina adamsoni, male (Thailand: Ban Takum; #665189; forewing length 29.0 mm); (M) A. mariana, male (holotype) (Guam: #229237; forewing length 24.5 mm) (see also Fig. 19B); (N) A. mariana, male (Saipan: #229242; forewing length 24.5 mm); (O) A. mariana, female (Marianas; #229245; forewing length 23.3 mm); (P) A. mariana, yellow female (Guam: Tarague; BPBM collection; forewing length 26.5 mm) (see also Fig. 19C); (Q) A. mariana, white female (Guam: Machanao; BPBM collection; forewing length 25.5 mm); (R) A. mata mata, male (Indonesia: Nias; #665146; forewing length 27.5 mm); (S) A. ?mata mata, female (Indonesia: ‘Sumatra’; previously identified as mata, but identity uncertain; #665196; forewing length 27.0 mm); (T) A. mata caeca, female (Indonesia: Sipora; BLKU collection; forewing length 28.0 mm).

Tachyris nero zarinda ab. aurosaFruhstorfer, 1899: 84. Original male specimen,‘Celebes, Macassar, W. Doherty, 1896’, ‘Type’, ‘zarinda ab. aurosa Fruhst'. In BMNH (examined), BMNH(E) #229202. (Unavailable name.)

Appias nero zarinda ab. aurosa Fruhstorfer; Fruhstorfer, 1910: 151; Jurriaanse & Lindemans, 1920: 9.

Appias zarinda ab. aurosa Fruhstorfer; Martin, 1919: 85.

Appias zarinda f. aurora [sic]; Talbot, 1923: 9.

Appias nero zarinda male form aurosa Fruhstorfer; Talbot, 1932: 161.

Appias zarinda zarinda male ab.; Yata, 1981: 373, pl. 63, fig. 4.

Appias sp.n. Yata & Vane-Wright, inVane-Wright & de Jong, 2003: 50, 109, pl. 6, fig. 13.

Diagnosis

Male. (Figs 19A, 20A). Forewing somewhat elongate apically and distinctly acute at apex. Ground colour of wings golden cream yellow. Upperside of wings without markings. Hindwing with pale orange oval androconial patch posteromedially. Genitalia (Fig. 5): uncus narrow and spindle-shaped viewed dorsally, apex blunt. Valve gradually narrowed posteriorly to rounded end, with a thick and arched interior process, directed dorsolaterally. Phallus thick and strongly arched dorsally, with coecum bearing a dorsal ridge; basal prong as long as coecum.

Female. Unknown, or unrecognized.

Distribution

This Indonesian endemic is known only from southern, central and south-east Sulawesi: Ujung Pandang (type series), Palopo (Yata, 1981), Camba (South Sulawesi, 2004, ex Nishiyama 2004, type series) and Buton island (Jurriaanse & Lindemans, 1920).

Foodplants

Unknown.

Description

Male. (Figs 19A, 20A). Forewing length: 36–40 mm (n = 13; mean = 37.35 mm; standard deviation = 1.197). Upperside: ground colour cream to yellow, with brighter yellow tinge, especially on discal cell, and along costal and distal margins. All dark markings obsolete, except basal areas of both wings dusted black, extensively so along basal half of forewing costa, and a black anticiliary line from apex to tornus. Ciliary fringe yellow. Hindwing with oval, faintly differentiated pale orange androconial patch located posteromedially across cubital cells, just extending into anal cells and discal cell. Underside: both wings almost same as upperside, but more extensively bright, especially on the forewing discal cell and hindwing. In some specimens postdiscal dark band weakly indicated, running obliquely from origin of vein M1 to near tip of vein CuA2 on forewing, and more rarely recognizable in cells M1–M3 of the hindwing. Forewing costa edged with some scattered black scales, but basal areas of wings not black dusted. Black anticiliary line from apex to tornus; ciliary fringe yellow. Forewing acute at apex, outer margin almost straight, but weakly incurved between veins M1–CuA1, and bluntly angled near tornus; middle discocellular almost half the length of upper discocellular, and about quarter length of lower discocellular. Hindwing evenly rounded on costal to outer margin, strongly curved near tornus; Rs arising free from cell; middle discocellular subequal to upper discocellular and lower discocellular. Antenna approximately 0.41× length of forewing, black and white chequered except on posterodorsal surface and a few apical segments; club subcylindrical. Thorax above and abdomen above near its base clothed with pale, yellowish hairs. Abdomen above blackish, with pale whitish-yellow scales beneath.

Male genitalia. (Fig. 5). Dorsum somewhat narrow, triangular in dorsal aspect, compressed medially, somewhat concave dorsomedially, with crossbow-like membranous area medially; valvenansatz (term from Droshin: see Shirôzu, 1960) broad, becoming more membranous ventrally; vinculum not strongly arched posteriorly; saccus short and rather slender (0.4× ring height); angle between vinculum and saccus approximately 110–120°. Uncus moderately long (0.45× ring height), narrow and spindle-shaped in dorsal view, bluntly pointed apically, with dorsal edge gradually narrowing towards apex; beak-shaped when viewed laterally, somewhat deeper medially, slightly hooked at tip. Valva nearly twice as long as high, gradually narrowing posteriorly to rounded tip, weakly produced posteroventrally, with a long and somewhat arched interior process arising near the middle at ventral margin of costa + ampulla region, directed dorsolaterally; patches of hooked spines present along ventral margin at posterior end and just beyond middle. Phallus moderately long, rather thick and strongly arched dorsally, subzonal sheath approximately one-fifth length of phallus; coecum with a small dorsal ridge; basal prong flattened laterally and somewhat swollen apically. Juxta arrowhead-like, somewhat swollen lateromedially when viewed from behind.

Female. Unknown.

Type material

Holotype male and 12 paratype males in BMNH, and one paratype male in BLKU, all of which have been labelled accordingly.

Holotype male and 12 paratype males in BMNH labelled: ‘Celebes, Macassar, W. Doherty, 1896’ (Ujung Pandang, South Sulawesi, Indonesia) (specimen register nos BMNHE 149984, 229197–229207, 229236). No. 229202 is labelled ‘zarinda aurosa Fruhstorfer in Seitz, p. 151’ (forewing length 40.0 mm) and is selected as holotype of Appias (Catophaga) aurosa Yata & Vane-Wright. No. 229207 (forewing length 37.0 mm), which carries the label ‘zarinda, ab. aurosa Fruhst./type’, is also labelled ‘Tachyris nero zarinda ab. aurosa Fruhstorfer, type specimen, det. O.Yata & R.I. Vane-Wright, 1998’. This last specimen apparently reached the BMNH from the Fruhstorfer Collection [Fruhstorfer (1899) had noted that Oberthür had sent him one specimen]; the other 12 all came from the C. Oberthür Collection, and are so labelled. One male paratype, labelled ‘S. Sulawesi, Camba, 2004’ (forewing length 35.5 mm), in BLKU.

Comments

Appias aurosa was described first as a male aberration of Appias zarinda from south Sulawesi by Fruhstorfer (1899), and the same status has been accorded to this insect by subsequent authors, as listed in the synonymy. However, we consider that ab. aurosa represents a distinct species – described and named here as new because Fruhstorfer's name is clearly infrasubspecific (and thus unavailable). On the basis of external phenotype and male genital morphology, this species is distinct from A. zarinda, and appears to be more closely related to A. albina.

Appias aurosa shares a number of potential synapomorphies with A. albina, including the spindle-shaped uncus with bluntly pointed apex, the thick and strongly arched phallus, and the oval androconial patch on the hindwing upperside. Appias aurosa is similar to A. zarinda in the narrowed valve, the tegumen with crossbow-like membranous area, and the sharply pointed forewing without markings. However, such a tegumen is probably symplesiomorphic, whereas the unique wing characters are almost certainly homoplasious (geographical parallelism). Large size coupled with a strongly arched forewing costa or pointed forewing tip affects many different butterflies endemic to Sulawesi (‘Celebes forewing’), a striking and mysterious convergence first noted by Alfred Russel Wallace (1865, 1867, 1869: 215–217) and discussed by Vane-Wright & de Jong (2003: 23). A sharply pointed, unmarked forewing affects three Catophaga taxa occurring on Sulawesi: A. zarinda zarinda, A. aurosa and A. paulina albata. Appias aurosa effectively replaces A. albina in Sulawesi, but coexists with A. zarinda. Appias albina is very rare in Sulawesi, even though it is quite common in neighbouring regions, such as Borneo, Palawan, Lesser Sundas and New Guinea. It is possible that albina has migrant status on Sulawesi (although the unique ‘black’ females of A. albina found there perhaps argue against this: Fig. 19P), where aurosa and albina might form a ‘species duplex’ (Corbet & Pendlebury, 1992). The systematic position of A. aurosa should become clearer once the female has been found and studied.

Appias (Catophaga) athama (Blanchard, 1848)

Pacific albatross (Figs 6, 19Q–S)

Figure 6.

Male genitalia of Appias (Catophaga) athama (New Hebrides: NHM OY genitalia preparation no. 7). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Pieris athama Blanchard, 1848: pl. 1, figs 10, 11

Much confusion has surrounded this species, including the question of its authority [the name Pieris athama was introduced separately by Lucas (1852)], and its type locality, ‘Balaou’. We now know that this represents the Fijian island of Ovalau (Appendix 1).

Pieris athama is treated in many publications as a subspecies or synonym, either of A. albina (e.g. D’Abrera, 1971, 1990; Holloway & Peters, 1976) or A. paulina (e.g. Yata, 1981) – although this has always been questioned by D’Abrera (1971, 1990). As dealt with by Tennent (2004, 2006), and confirmed by the present study, A. athama is a distinct species endemic to New Caledonia and adjacent regions of the western Pacific.

Diagnosis

Male. (Fig. 19Q). Forewing broad and weakly falcate near apex. Upperside ground colour white with weak lustre. No markings, but costa to outer margin edged with a distinct, fine, black margin. Hindwing with oval, whitish androconial patch posteromedially. Underside ground colour usually dark or bright yellow on forewing apex and on hindwing. Genitalia (Fig. 6): tegumen moderately swollen anterodorsally; uncus narrow and spindle-shaped viewed dorsally, with blunt apex and not abruptly curved ventrally. Valva broad and rounded posteriorly. Phallus thick and moderately arched dorsally, with long coecum with a weak dorsal ridge; basal prong somewhat shorter than coecum.

Female. (Fig. 19R, S). Forewing broad and weakly falcate near apex. Upperside ground colour white to yellow with reddish tinge. Forewing with pale submarginal spots in cells R5, M1 and M3, but rarely traceable in cell M2. Hindwing black distal border evenly broad with marginal triangular pale spots. Underside markings almost as upperside, but with forewing apex and margin of hindwing border paler.

This species closely resembles A. albina, but is readily distinguished from the latter in the male by the falcate forewing distinctly edged with black and the bluntly pointed uncus not abruptly curving ventrally at apex, and in the female by the even, black hindwing border with marginal triangular pale spots.

Distribution

Recorded from New Caledonia, Loyalty Is., Isle of Pines, Vanuatu, Banks Islands, Fiji, Tonga and Western and American Samoa [for details see Tennent (2006)]. This species occurs sympatrically with A. paulina in the eastern part of its range.

Foodplants

Apparently unknown. Both Capparis and Drypetes occur in the western Pacific, including Samoa (Amerson et al., 1982).

Appias (Catophaga) paulina (Cramer, 1777)

Yellow albatross (Figs 7, 20B–L)

Figure 7.

Male genitalia of Appias (Catophaga) paulina (Myanmar: Karen Hills; NHM genitalia preparation no. 4651). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Papilio paulina Cramer, 1777: 21, 150, pl. 110, figs E,F

The published type locality was ‘Côte de Coromandel, à Tranquebar & dans l’isle de Java, près de Batavia’. This taxon was long considered to be based on material from Sri Lanka (Fruhstorfer, 1910: 155; Talbot, 1939: 404; Wynter-Blyth, 1957: 430; Edwards et al., 2001: 466), or south-east India (e.g. Parsons, 1998: 291). However, the most similar Appias from Sri Lanka is the distinct A. galene (see above). An authentic ex van Lennep ‘Cramer specimen’ has been located in the BMNH (Chainey, 2005: 329, fig. 40), and is here designated lectotype of Papilio paulina Cramer (Appendix 2). With very little doubt it is Javanese in origin, and we consider that the nominate subspecies of A. paulina represents the race found on Java. The idea that the original locality was Sri Lanka (and/or southern India) appears erroneous.

Diagnosis

Male. (Fig. 20B, F, H, J, L). Forewing somewhat obtuse at apex, which may or may not be narrowly darkened. Upperside ground colour white, both wing surfaces usually with a free dark spot in cell M3. Hindwing without oval sex-patch. A group of long and very prominent scales on dorsum of eighth tergite (Parsons, 1998: 290) (other Catophaga species have similar scales in this position, but they are shorter and less prominent). Genitalia (Fig. 7): uncus spatula-shaped viewed dorsally and blunt-ended, with its dorsal ridge well developed. Valva broad and rounded posteriorly, with a thick and arched interior process, directed dorsolaterally. Phallus thick and strongly arched dorsally, with elongate coecum bearing a dorsal ridge; broadened basal prong shorter than coecum; common stem of coecum and basal prong well developed.

Female. (Fig. 20C–E, G, I, K). Upperside ground colour white, sometimes yellow. Forewing black apical area usually with five submarginal whitish spots in cells R2 and R5–M3, those in cells M2 and M3 often reduced; heavily black-dusted basal area with distal margin almost perpendicular to posterior margin of forewing (not oblique). As in other members of the subgenus, a black, well-developed genital hair tuft arises on the distal margin of the eighth sternite (Yata, 1981).

Comment. Many subspecies of A. paulina are currently recognized (Appendix 1). It seems plausible that future work, including the potential application of molecular methods, will show that some represent distinct species. For example, races such as A. paulina eurosundana from Alor, A. p. paula from Wetar and A. p. galathea (Fig. 20B–E) from the Andamans and Nicobars, appear significantly different to nominate paulina.

Distribution

As currently conceived, A. paulina is a widely distributed species found throughout much of the Indo-Australian Region, including New Guinea and parts of Australia (where it has even been recorded as far south as Tasmania: Braby, 2000: vol. 1, 325; Braby 2005), Christmas Island and Lord Howe, and from northern India to New Caledonia and Vanuatu (Tennent, 2006: 31). However, it does not occur in Pakistan, peninsular India and Sri Lanka, or in the Bismarck Archipelago (Parsons, 1998: 291) or the Solomons (Tennent, 2002: 63) and appears unrecorded from the Mentawai Islands. Appias paulina minato occurs in Taiwan and Kagoshima Prefecture of Japan (Ryukyu or Loocho Islands), with its northern limit being Akuseki-jima, in the Tokara Is., a little to the north of Amami-ohshima. Prior to the 1960s, in the Ryukyus it was known from Okinawa, Ishigaki, Iriomote, Hateruma and Yunaguni (Takara, 1956: 66, as Appias melania minato). It has also been recorded from Yaeyama and Miyako, and apparently spread to Amami-ohshima in 1980 and Akuseki-jima in 1975 or 1982. Further northerly movement, including to Kyushu, is limited by the absence of its larval foodplant, Drypetes matsumurae Kaneh (H. Fukuda, personal communication, 2007).

Foodplants

Capparis (Capparaceae), Drypetes, Putranjiva (Putranjivaceae) (Braby, 2000: 466; Robinson et al., 2001). Igarashi & Fukuda (2000: 392) noted Drypetes matsumurae as the foodplant in Japan and Taiwan, D. littoralis in Palawan and D. deplanchei in Australia, and commented that an old record for Capparis heyneana‘requires reconfirmation’.

Appias (Catophaga) mariana Yata & Chainey sp.n.

Mariana albatross (Figs 8, 9, 19B, C, 20M–Q)

Figure 8.

Male genitalia of Appias (Catophaga) mariana Yata & Chainey sp.n. (Marianas: Guam; NHM specimen register number BMNH(E)# 229237). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Figure 9.

Female genitalia (signum) of Appias (Catophaga) mariana Yata & Chainey sp.n. (Marianas: Rota: Tatacho-Songsong; AGKU OY genitalia preparation no. 1189). Posterior aspect. 0.5 mm.

Appias leis subtuslutea Roepke; Swezey, 1942: 66, pl.9, figs. 12, 15. Misidentification

Appias paulina jacquinotii Lucas; Yata, 1981: 379. Misidentification

Appias paulina Cramer; Fukuda & Nicho, 2001: 31, 33. Misidentification

Diagnosis

Male. (Figs 19B, 20M, N). Small species (forewing length below 27.0 mm). Forewing somewhat acute at apex, more pointed than in paulina. Upperside ground colour white, often with pale greenish-yellow tinge. Black postdiscal band present in cells M1–M3, and usually expressed in cells R4+5 and CuA1, never reduced to an isolated spot, even in cell M3. On underside forewing postdiscal black band usually present from costa to cell M3, but barely visible in cell CuA1, as on upperside. Genitalia (Fig. 8): uncus somewhat short (0.5× ring height), narrow and spatula-shaped viewed dorsally, blunt-ended, with dorsal ridge well developed. Valva broad and weakly rounded posteriorly. Phallus moderately long, thick and strongly arched dorsally.

Female. (Figs 19C, 20O–Q). Upperside forewing with distal margin of black basal area somewhat oblique. Underside forewing postdiscal black band well marked from costa to tornus. Signum broad, diamond-shaped when viewed posteriorly, with many spines (Fig. 9).

Distribution

Endemic to Mariana Islands (Guam, Saipan, Rota).

Foodplants

Unknown. Hostplants that should probably be considered are Drypetes dolichocarpa Kanehira, endemic to the Marianas, and/or Capparis cordifolia Lamarck (Fosberg et al., 1975).

Description

Male. (Figs 19B, 20M, N). Forewing length: 19–27 mm (n = 10, mean 24.15 mm; standard deviation = 1.817). Upperside: ground colour white, usually with pale greenish-blue tinge. Forewing black distal border almost regular and gradually narrowed towards tornus, outwardly concave in cells M1–CuA1, and almost obsolete in cell CuA2. Black postdiscal band present in cells M1–M3, and usually traceable in cells R4+5 and CuA1 as much narrower and faint streak close to distal border, and not reduced to an isolated spot even in cell M3. Black anticiliary line recognizable from apex to tornus; fringe black, but whitened towards tornus. Hindwing black distal border generally poorly developed, usually reduced to marginal vein-dots, often obsolete. Black anticiliary line traceable from apex to tornus; ciliary fringe white. Basal portions of both wings black dusted, extensively so towards apex along costal margin of forewing. Underside: ground colour almost the same as on upperside, but somewhat darker, rarely yellowish on forewing apex and on hindwing. Postdiscal black band usually present from costa to cell M3, and barely traceable in cell CuA1, as on upperside. Black anticiliary line barely traceable from apex to tornus on forewing; fringe white. Forewing somewhat acute at apex; outer margin almost straight, but weakly rounded between veins CuA1 and CuA2; bluntly angulate near tornus; middle discocellular subequal to upper discocellular, almost half the length of lower discocellular. Hindwing evenly rounded on costal to outer margin, strongly curved near tornus; Rs arising free from cell; middle discocellular subequal to or less than upper discocellular or lower discocellular. Antenna approximately 0.41× length of forewing, black and white-checkered, except on the posterodorsal surface and a few apical segments; club cylindrical. Thorax above and abdomen above near base clothed with whitish hairs. Abdomen above blackish with scales beneath whitish.

Male genitalia. (Fig. 8). Dorsum elongate-triangular in dorsal aspect, compressed medially, concave dorsomedially, fairly swollen anterodorsally; valvenansatz broad, becoming membranous ventrally; vinculum not strongly arched posteriorly; saccus short and relatively slender (0.25× ring height); angle between vinculum and saccus approximately 100–110°. Uncus somewhat short (0.5× ring height), narrow and spatula-shaped viewed dorsally, with a blunt tip, the dorsal ridge well developed, beak-shaped viewed laterally, and distinctly deeper medially, and slightly hooked at its tip. Valva broad and weakly rounded posteriorly, with a thick and arched interior process directed dorsolaterally; a batch of curled, short setae present along ventral margin of valva just beyond its middle. Phallus moderately long, thick and strongly arched dorsally, subzonal sheath approximately one-fifth length of phallus, with elongate coecum bearing a weak dorsal ridge; broadened basal prong as long as coecum; common stem of coecum and basal prong well developed. Basal prong flattened laterally and somewhat swollen apically. Juxta arrowhead-like, but ending bluntly ventrally, and somewhat swollen lateromedially when viewed from behind.

Female. (Figs 19C, 20O–Q). Forewing length: 22–26 mm (n = 3, mean = 24.77 mm; standard deviation = 1.27). Upperside: ground colour white, with greenish-yellow tinge, sometimes yellowish on hindwing. Forewing black costal border broad, with its inner margin indistinct basally; black distal border generally broad, fused with black postdiscal band, with its inner edge irregular from costa to vein CuA1, obtuse-angled at vein CuA1, deeply excavated in cell CuA1; a series of submarginal white spots appearing in cells R2 and R5–M3. Ciliary fringe black. Basal portion of forewing extensively black dusted, fused with costal border; black-dusted basal area with distal margin somewhat oblique. Hindwing black distal border broad, with its inner edge moderately defined, zigzag-shaped. Whitish anticiliary line recognizable from apex to tornus. Ciliary fringe black. Basal area and anal margin of hindwing black dusted. Underside: pale ground colour almost the same as upperside, but darker, especially on forewing apex and on hindwing; rarely dark yellow (Fig. 20P). Black anticiliary line barely traceable from apex to tornus on forewing. Ciliary fringe whitish. Forewing postdiscal black band well marked from costa to tornus. Forewing somewhat obtuse at apex; outer margin almost straight, but weakly rounded between veins CuA1 and CuA1; bluntly angulate near tornus. Hindwing evenly rounded on costal to outer margin, curved near tornus. General features of thorax, abdomen and venation similar to male.

Female genitalia. (Fig. 9). The papilla analis almost elliptical with apical lobe, some hairs and a slender apophysis posterioris. The lobulus vaginalis (Kusnezov, 1915: 132, fig. 72) weakly sclerotized, with an apophysis anterioris, and divided into upper and lower lobes, of which the ventral is broad and evenly curved posteriorly. The ductus bursae is moderately long, and twisted near border of cervix bursae. The ductus seminalis attaches to the tip of the cervix bursae. Corpus bursae large and subglobular, with a smaller appendix bursae (as in many other genera of Pieridae, including Cepora Billberg). Signum broad, diamond-shaped when viewed posteriorly, strongly constricted medially, with spines covering entire surface.

Type material

Holotype male and eight paratype males in BMNH, 15 paratypes in AGKU and four paratypes in BPBM, all of which have been labelled accordingly.

Holotype male and four paratype males in BMNH labelled: Guam, Rothschild Bequest, B.M. 1939-1 (specimen register nos BMNH(E) 229237–229241). No. 229237 is dated [18]95 (forewing length: 24.5 mm) and is selected as holotype of Appias (Catophaga) mariana Yata & Chainey. Nos 229238 and 229240 are dated March [18]95. No. 229239 is dated 10.x.[18]94. No. 229241 is dated ‘11–10’ [18]94. One paratype male in BPBM labelled: Guam, Ritidian pt., 22.iv.1936, E.J. Bryan, Jr. One paratype male and one paratype female in BPBM labelled: Guam, Tarague, 17.v.1936, O.H. Swezey. One paratype female in BPBM labelled: Guam, Machanao, 2.vi.1936, O.H. Swezey. Three paratype males in BMNH labelled: Saipan, Rothschild Bequest, B.M. 1939-1 (specimen register nos BMNH(E) 229242–229244). No. 229242 dated viii.[18]95. Nos 229243–229244 dated 13.ix.[18]95. One paratype female in BMNH labelled ‘Museum Paris Mariannes M.A. Marche 194-88’, ex Oberthür coll., Brit. Mus. 1927–3 (specimen register no. BMNH(E) 229245). One paratype male in AGKU labelled: Rota Is. South Coast, 2.v.1936, T. Esaki. Nine paratype males and two paratype females in AGKU labelled: Rota, Tatacho-Sonson, 8.ii.1936, T. Esaki. One paratype male and two paratype females in AGKU: labelled: Rota, Teteto- Tatacho-Sonson, 5.xi.1937, T. Esaki.

Comments

This species closely resembles A. paulina, but is distinguishable from the latter by its smaller size (male forewing length not known to exceed 27.0 mm), more sharply pointed forewing apex, the distinct forewing postdiscal black band present from costa to cell M3 on both wing surfaces of the male, and its shorter uncus with better developed dorsal ridge.

This butterfly has not been collected or recorded, as far as we are aware, since 1937, and could be extinct. Fukuda & Nicho (2001) did not encounter this species in the highly degraded environments of Guam during February and November 2000, and did not find evidence of further records of it from Guam since Swezey's fieldwork of 1936 (Swezey, 1942). Asou (2000) failed to find A. mariana on Saipan or Managaha during a visit in 1998. Although a small white butterfly is very likely to be passed over as a common species of little interest, this insect should be sought, especially on Saipan and Rota.

Appias (Catophaga) mata (Kheil, 1884)

Kheil's albatross (Figs 10, 20R–T, 22L)

Figure 10.

Male genitalia of Appias (Catophaga) mata (Indonesia: Sipora; BLKU OY genitalia preparation no.1187). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Figure 22.

Adult Appias (Catophaga) butterflies (halved: left hand upperside/right hand underside; all in BMNH). (A) A. nero nero, male brown aberration (Malaysia: Perak, Tapah Hills; #665154; forewing length 36.8 mm); (B) A. nero?, male yellow aberration (Malaysia: Perak; #665158; forewing length 37.0 mm); (C) A. nero nero, female (Malaysia: Taiping; #665156; forewing length 33.0 mm); (D) A. nero nero, mixed gynandromorph (Malaysia: Cameron Highlands; #665157; forewing length 36.5 mm); (E) A. zarinda, male (Indonesia: Sulawesi, Tanah Mateh Pa; #665132; forewing length 42.5 mm); (F) A. zarinda, white female (Indonesia: Sulawesi, Minahassa, #665135; forewing length 39.5 mm); (G) A. zarinda, pale yellow female (Indonesia: Sulawesi, Manado, #665133; forewing length 40.0 mm); (H) A. zarinda, orange female (Indonesia: Sulawesi, Minahassa, Tanggari; #665134; forewing length 39.4 mm); (I) A. placidia, male (Indonesia: Obi; #665136; forewing length 33.0 mm); (J) A. placidia, grey female (Indonesia: Ceram; Mansela; #665138; forewing length 37.2 mm); (K) A. placidia, yellow female (Indonesia: Ceram; #665137; forewing length 35.8 mm); (L) A. mata caeca, male (holotype) (Indonesia: Mentawi Is, Sipora; #142295; forewing length 29.6 mm); (M) A. clementina, male (Indonesia: Tenimber; #665139; forewing length 25.5 mm); (N) A. clementina, female (Indonesia: Tenimber; #665140; forewing length 26.7 mm); (O) A. celestina barea, male (Indonesia: Aru; #665141; forewing length 37.5 mm); (P) A. celestina barea, male (syntype) (Indonesia: Aru; #135576; forewing length 35.0 mm); (Q) A. celestina barea, white female (Indonesia: Aru; #665143; forewing length 35.1 mm); (R) A. celestina barea, yellow female (Indonesia: Aru; #665142; forewing length 33.3 mm); (S) A. melania, male (Australia: Queensland, Kuranda; #665192; forewing length 35.0 mm); (T) A. melania, female (Australia: Queensland, Kuranda; #665145; forewing length 33.0 mm).

Pieris mata Kheil, 1884: 34, pl. 4, fig. 31.

Diagnosis

Male. (Figs 20R, 22L). Forewing somewhat obtuse at apex. Upperside ground colour bluish-grey. Forewing usually with free dark spot in cell M3. Hindwing without oval sex-patch. Genitalia (Fig. 10): uncus spatula-shaped viewed dorsally, blunt-ended, with dorsal ridge well developed. Valva broad and rounded posteriorly, with a thick and arched interior process directed dorsolaterally. Phallus thick and strongly arched dorsally, with elongate coecum bearing a dorsal ridge; broadened basal prong shorter than coecum; common stem of coecum and basal prong well developed.

Female. (Fig. 20S, T). Apparently dimorphic. Upperside ground colour pale yellow, but almost whitish in forewing, or pale bluish, somewhat male-like (not clearly evident in Fig. 20T). Forewing black apical area with four submarginal whitish spots in cells R2 and R5–M2, that in cell M2 almost obsolete; black-dusted basal area with distal margin almost perpendicular to posterior margin of forewing.

Distribution

Known only from Nias and Sipora (Mentawai Islands). The two subspecies, A. m. mata (Nias) and A. m. caeca (Sipora) are very distinct, and it is possible they represent separate species.

Foodplants

Unknown. At least two species of Drypetes have been recorded from Siberut (Hadi et al., 2009).

Appias (Catophaga) galba (Wallace, 1867) stat.rev.

Wallace's albatross (Figs 11, 12A–C, 21A–F)

Figure 11.

Male genitalia of Appias (Catophaga) galba (India: Khasi Hills; NHM genitalia preparation no. 4603). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Figure 12.

Female genitalia (signum) of three Appias (Catophaga) species, posterior view. (A–C) A. galba (A, N.W. Myanmar: BLKU OY genitalia preparation no. 1190; B, India: Kashi Hills: NHM genitalia preparation no. 4604; C, Hainan, BLKU OY genitalia preparation no. 1191); (D–F) A. nero (D, Malay Peninsular; BLKU OY genitalia preparation no.1192; E, Philippines: Palawan, NHM genitalia preparation no. 4613; F, Indonesia: Bawean, NHM genitalia preparation no. 4609); (G, H) A. zarinda (G, Indonesia: Sulawesi; BLKU OY genitalia preparation no. 1193; H, Indonesia: Sulawesi; BLKU OY genitalia preparation no. 1194). Scale bar = 0.5 mm.

Figure 21.

Adult Appias (Catophaga) butterflies (halved: left hand upperside/right hand underside; all in BMNH). (A) A. galba, male (India: Meghalaya, Khasia Hills; #665195; forewing length 38.3 mm); (B) A. galba, male (Thailand/Vietnam: Black River, Nam Hou; #665148; forewing length 36.7 mm); (C) A. galba, male (Myanmar: Muenghi Hill Tracts, Hukawng Valley; #665149; forewing length 38.0 mm); (D) A. galba, female (India: Sikkim; #665151; forewing length 38.9 mm); (E) A. galba, female (India: Meghalaya, Cherrapunji; #665194; forewing length 40.5 mm); (F) A. galba, female (Vietnam: Laokay, Muong-Khuong; #665152; forewing length 38.0 mm); (G) A. nero soranus, male (Philippines: Negros, #665168; forewing length 35.6 mm); (H) A. nero soranus, female (Philippines: Negros, Amlan Falls; #665169; forewing length 34.1 mm); (I) A. nero palawanica, male (Philippines: Palawan; #665162; forewing length 35.0 mm); (J) A. nero palawanica, white female (Philippines: Palawan; #665163; forewing length 30.0 mm); (K) A. nero palawanica, piebald female (Philippines: Palawan; #665164; forewing length 35.0 mm); (L) A. nero palawanica, yellow female (Philippines: Palawan; #665167; forewing length 34.5 mm); (M) A. nero palawanica, orange female (Philippines: Palawan; #665166; forewing length 33.0 mm); (N) A. nero palawanica, red female (Philippines: Palawan; #665193; forewing length 34.0 mm); (O) A. nero flavius (syntype), male [‘N Born’ (= Philippines: Taganak); #229186; forewing length 33.5 mm]; (P) A. nero flavius, yellow female (Philippines: Taganak; #665159; forewing length 32.0 mm); (Q) A. nero flavius, deep yellow female (Philippines: Taganak; #665161; forewing length 34.5 mm); (R) A. nero flavius, orange female (Philippines: Taganak; #665160; forewing length 34 mm); (S) A. nero nero, male [Malaysia: ‘Camp Zor’ (?); #665153; forewing length 31.0 mm]; (T) A. nero nero, pale ‘sufflava’ male (Indonesia: Sumatra, Setinjak; #665155; forewing length 36.3 mm).

Tachyris galba Wallace, 1867: 378.

In all recent publications known to us (e.g. Yata, 1981; D’Abrera, 1982, Osada et al., 1999), and for many years previously (e.g. Talbot, 1939), galba has been treated as a subspecies of A. nero. As demonstrated by our present study, A. galba is a distinct species.

Diagnosis

Male. (Figs 21A–C). Forewing somewhat falcate and distinctly acute at apex. On upperside black distal border and a series of postdiscal black spots usually present. Hindwing fairly pointed near tornus. Genitalia (Fig. 11): uncus nib-shaped viewed dorsally with its apex somewhat pointed, and not so abruptly curved ventrally as nero. Valva somewhat narrowed posteroventrally with a thick and arched interior process dorsolaterally. Phallus weakly arched dorsally, with coecum usually having a dorsal ridge; basal prong as long as coecum.

Female. (Fig. 21D–F). Forewing somewhat falcate and distinctly acute at apex. On upperside forewing black distal border and postdiscal black zigzag band present. Hindwing fairly pointed at tornus. Upperside ground colour dull orange-red. Forewing black apical area usually with seven submarginal orange spots in cells R2–CuA1, representing the ground colour between the dark distal border and the dark postdiscal band; forewing upperside without any dark discocellular marking. Signum relatively broad, with many spines (Fig. 12A–C).

Comment

This species closely resembles A. nero, but is easily distinguished from the latter by the falcate forewing with distinctly acute apex, a series of postdiscal black spots on forewing upperside in the male, the distinctly angled hindwing near the tornus, and in the female the absence of any infuscation of the forewing discocellular veins, and broader signum. Some galba males have a slightly ‘two-toned’ look to the hindwing, with the base deeper red than the remainder, which is more orange and becomes yellowish towards the tornus (Fig. 21B), and there can be a suggestion of this two-tone effect on the forewing in some females (e.g. Fig. 21F). We have not observed this phenotype in A. nero or A. zarinda.

Distribution

Endemic to northern India (Sikkim), Burma, northern Thailand, northern Indochina and Hainan, including Laos (Osada et al., 1999: 202), and also recorded from Guangxi, China (Chou, 1994 vol. 1: 243). Appears to be parapatric to A. nero.

Foodplants

Apparently unknown.

Appias (Catophaga) nero (Fabricius, 1793)

Orange albatross (Figs 12D–F, 13, 21G–T, 22 A–D)

Figure 13.

Male genitalia of Appias (Catophaga) nero (Indonesia: Tanah Djampea; NHM genitalia preparation no. 4600). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Papilio nero Fabricius, 1793: 153.

Papilio nero was based on material in the British Museum, but without indication of origin. To stabilize current usage (due to Butler, 1870b), a neotype has been selected from West Java (Appendix 2).

Diagnosis

Male. (Figs 21G, I, O, S; 22A, B). Forewing somewhat acute at apex. Upperside ground colour orange-red, but occasionally straw-coloured (Fig. 21T), or brown in one unique example (Fig. 22A). Forewing upperside of male usually narrowly margined with black. Genitalia (Fig. 13): uncus nib-shaped viewed dorsally with its apex sharply pointed. Valva gradually narrowed posteroventrally, usually weakly produced apically, with a thick and arched interior process dorsolaterally. Phallus weakly arched dorsally, with coecum usually lacking a dorsal ridge; basal prong as long as coecum.

Female. 21H, J–N, P–R; 22C, D). Upperside ground colour usually dull orange red, but sometimes white to yellow; the main Philippine races do not have orange females, only white/yellow. Forewing black apical area usually with seven submarginal light spots in cells R2 and R5–CuA2, those in cells R2, M2 and CuA1 often reduced; forewing upperside with discocellular marking represented by a short black bar; dark basal area with distal margin usually diffuse, but more distinct and often almost perpendicular to posterior margin of forewing in main Philippine races. Signum narrow, with many spines (Fig. 12D–F).

Distribution

Widely distributed in Sundaland and the Philippines, but replaced in the Sulawesi region by A. zarinda. According to Corbet & Pendlebury (1992: 90), the female is rarely encountered in the Malay Peninsula, occurring mainly in forests above approximately 600 m. Judging by museum collections, females are more readily encountered in populations occurring in Wallacea, most notably the highly polymorphic race found on Palawan. In this context it may be notable that Jumalon (1970) recorded A. n. palawanica from human faeces used as bait. It seems possible that females of the more eastern, often highly polymorphic subspecies differ in their behaviour – and the same may be true for A. zarinda as well.

Foodplants

Capparis (Capparaceae) and Drypetes (Putranjivaceae) (Robinson et al., 2001). Dupont & Scheepmaker (1936: 46) recorded Drypetes macrophylla and, apparently, Pithecellobium lobatum (as ‘Pithecolobium’: Fabaceae) as foodplants in Java – but the latter is unconfirmed; Roepke (1935: 64) gave D. macrophylla only, based on earlier work by Leefmans.

Appias (Catophaga) zarinda (Boisduval, 1836)

Flame albatross (Figs 12G, H, 14, 22E–H)

Figure 14.

Male genitalia of Appias (Catophaga) zarinda (Indonesia: south Sulawesi; NHM genitalia preparation no. 4619). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Pieris zarinda Boisduval, 1836: 486, pl. 18, fig. 4.

Diagnosis

Male. (Fig. 22E). Forewing elongate apically and sharply pointed at apex. Upperside ground colour orange-red. Upperside of wings without markings. Genitalia (Fig. 14): uncus narrow and nib-shaped viewed dorsally, with apex sharply pointed. Valva gradually narrowed posteroventrally, produced apically, with a thick and arched interior process directed dorsolaterally. Phallus moderately arched dorsally; coecum usually with a dorsal ridge; basal prong as long as coecum.

Female. (Fig. 22F–H). Upperside heavily marked with black; ground colour dull orange-red, white or creamish-yellow. Forewing usually with seven pale submarginal spots in cells R2 and R5–CuA2, those in cells M2 and CuA1 reduced; hindwing median light band with both margins sharply defined; black basal area with distal margin somewhat oblique. Genitalia: signum large, asymmetrical, with left-hand end more extended, with many spines (Fig. 12G, H).

Comment

Recognized as a species by Yata (1981), A. zarinda closely resembles A. nero, but is readily distinguished from the male of the latter by the sharply pointed forewing apex, upperside of wings without markings and veins not black-dusted, and in the female by the larger, asymmetrical signum.

Distribution

Found only on Sulawesi and some adjacent islands, Sula, and Buru (Peggie et al., 1995; Vane-Wright & de Jong, 2003).

Foodplants

Apparently unknown.

Appias (Catophaga) placidia (Stoll, 1790)

Chocolate albatross (Figs 15, 22I–K)

Figure 15.

Male genitalia of Appias (Catophaga) placidia (Indonesia: Seram; NHM genitalia preparation no. 4614). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Papilio placidia Stoll, 1790: 133, pl. 28, figs 4,4c.

Diagnosis

Male. (Fig. 22I). Forewing somewhat acute at apex. Upperside ground colour dark brown, except for a very fine golden marginal fringe that does not occur in other Catophaga species, including nero. Hindwing with dark, oval sex-patch. Genitalia (Fig. 15): uncus somewhat broad and depressed, and spatula-shaped viewed dorsally, with apex bluntly pointed. Valva broad and rounded posteriorly, distinctly incurved ventromedially, with a thick and arched interior process, directed dorsolaterally. Phallus moderately arched dorsally with coecum bearing a dorsal ridge; somewhat broadened basal prong as long as coecum.

Female. (Fig. 22J, K). Upperside ground colour dark brown. Forewing usually with seven pale submarginal spots in cells R2 and R5–CuA2, those in cells M2, CuA1 and CuA2 often reduced. Ground colour beneath obscure greenish-white, or yellowish.

Distribution

Restricted to North and Central Maluku (but not found on Kep. Sula or Buru) (Peggie et al., 1995).

Foodplants

Apparently unknown.

Appias (Catophaga) clementina (Felder, 1860)

Clementine's albatross (Figs 16, 22M, N)

Figure 16.

Male genitalia of Appias (Catophaga) clementina (Indonesia: Tanimbar; BLKU OY genitalia preparation no. 1185). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Pieris clementina C. Felder, 1860: 448.

Diagnosis

Male. (Fig. 22M). Forewing length less than 30 mm. Forewing somewhat acute at apex. Upperside ground colour greyish-blue, underside yellowish-grey. Male usually with dark free spot in cell M2. Forewing usually with seven light submarginal spots in cells R2 and R5–CuA2, with that in cell M2 often reduced. Hindwing with pale oval sex-patch. Genitalia (Fig. 16): uncus somewhat broad and spatula-shaped viewed dorsally, with its apex bluntly pointed. Valva broad and rounded posteriorly, distinctly incurved ventromedially, with a thick and arched interior process, directed dorsolaterally. Phallus fairly arched dorsally with coecum lacking a dorsal ridge; somewhat broadened basal prong as long as coecum.

Female. (Fig. 22N). Forewing length less than 28 mm. Upperside ground colour greyish-blue, relieved by a white postdiscal band on both wings. Forewing usually with seven pale submarginal spots in cells R2 and R5–CuA2, that in cell M2 often reduced. Underside greenish-blue basally.

Distribution

Alor, Timor, Wetar, Roma, Moa, Damar, Teun, Sermata, Babar, Tanimbar (Selaru, Yamdena, Larat, Maru) (Rawlins, 2007).

Foodplants

Apparently unknown.

Appias (Catophaga) celestina (Boisduval, 1832)

Blue albatross (Figs 17, 22O–Q)

Figure 17.

Male genitalia of Appias (Catophaga) celestina (Indonesia: Waigeo; NHM genitalia preparation no. 4616). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Pieris celestina Boisduval, 1832: 46.

Diagnosis

Male. (Fig. 22O, P). Forewing somewhat acute at apex. Upperside ground colour greyish-blue, with curvilinear row of largely connected dark postdiscal spots running from base of cell R5 to margin of CuA1. Hindwing with oval sex-patch. Genitalia (Fig. 17): uncus very broad and spatula-shaped viewed dorsally, with apex bluntly pointed. Valva broad and rounded posteriorly, distinctly incurved ventromedially, with a thick and arched interior process directed dorsolaterally. Phallus moderately arched dorsally; coecum bearing a weak dorsal ridge; somewhat broadened basal prong as long as coecum; common stem of coecum and basal prong well developed.

Female. (Fig. 22Q, R). Upperside ground colour whitish, with some greyish-blue scales along inner margin of hindwing dark margin and basally, or yellow. Forewing usually with six pale submarginal spots in cells R2 and R5–CuA1, those in cells M2 and CuA1 often reduced; darkened basal area with distal margin oblique.

Distribution

Found in New Guinea, Aru, Kai, Bismarck Archipelago and Bougainville (but not in the Solomon Islands: Tennent, 2002), and in the rainforest at Iron Range, Cape York Peninsula, Queensland (one male in good condition recently recorded: Weir et al., 2005).

Foodplants

Unknown.

Appias (Catophaga) melania (Fabricius, 1775)

Grey albatross (Figs 18, 22S, T)

Figure 18.

Male genitalia of Appias (Catophaga) melania (Australia: Queensland; BLKU OY genitalia preparation no. 722). (A) Ring, lateral views; (B) dorsum, dorsal views; (C) right valva, inner view; (D) phallus, lateral and dorsal views; (E) juxta, anal view. Scale bar = 0.5 mm.

Papilio melania Fabricius, 1775: 475.

Diagnosis

Male. (Fig. 22S). Forewing somewhat elongate and somewhat acute at apex. Upperside ground colour grey-blue, with more whitish postdiscal area extending into apex of discal cell. Forewing with an oblique black band running from costa to origin of vein M1 to tip of CuA1. Forewing black apical area usually with six light submarginal spots in cells R2 and R5–CuA1, that in cell CuA1 often obsolete. Hindwing with pale oval sex-patch. Underside forewing with an extensive orange-yellow streak occupying anterior half of discal cell; hindwing extensively brownish, similar in colour to underside forewing apex. Genitalia (Fig. 18): uncus somewhat broad and spatula-shaped viewed dorsally, bluntly pointed apically, with dorsal ridge well developed. Valva broad and rounded posteriorly, distinctly incurved ventromedially, with a thick and arched interior process, directed dorsolaterally. Phallus strongly arched dorsally; coecum with a dorsal ridge; somewhat broadened basal prong as long as coecum.

Female. (Fig. 22T). General features similar to male, but ground colour paler and black areas broader.

Distrubution

Restricted to a small upland area of Queensland, north-eastern Australia, from the latitude of Cooktown to the southern end of the Paluma Range, inland from Townsville. Sometimes migrates to the coast at Townsville or even further south where, however, it does not breed (Dunn & Dunn, 1991: 55; Braby, 2000: 327).

Foodplants

Drypetes (Putranjivaceae) (Braby, 2000).

Key to species of Appias (Catophaga)

Note: Males of the Appiadina have a conspicuous apicoventral abdominal hair-tuft. However, females of Catophaga have a similar tuft, and this character cannot readily be used to separate the sexes of species belonging to this subgenus.

  • 1. Male: forewing apex relatively pointed, wings usually less heavily patterned with black .................................................................2

  • – Female: forewing apex relatively rounded, wings more heavily patterned with black (note: female of aurosa unknown) .................................. 16

  • 2. (Males) Wings extensively reddish-orange, rarely straw-coloured (nero group)..................................................................... 3

  • – (Males) Wings never reddish-orange......................................5

  • 3. Hindwing distinctly angled near tornus; upperside forewing apex darkened, usually with a row of dark postdiscal spots that form a weak band running midway between end of discal cell and wing apex; distal sections of hindwing veins darkened........................................................ galba (Wallace, 1867)

  • – Hindwing not distinctly angled near tornus...............................4

  • 4. Forewing sharply pointed at apex; upperside of forewing without black veins zarinda (Boisduval, 1836)

  • – Forewing not sharply pointed at apex; upperside of forewing usually with black veins (not in A. nero flavius from the Turtle Islands, Philippines); upperside sometimes straw coloured....................................nero (Fabricius, 1793)

  • 5. Wings dark brown or bluish (at least hindwings).........................6

  • – Wings not brown or bluish ............................................. 10

  • 6. Upperside ground colour dark chocolate brown...................placidia (Stoll, 1790)

  • – Upperside ground colour bluish or bluish-grey (at least on hindwings) .... 7

  • 7. Hindwing without oval whitish sex-patch; forewing usually with free dark spot in cell M3 (Mentawai Islands only) ............................... mata (Kheil, 1884)

  • – Hindwing with oval whitish sex-patch on upperside (sometimes indistinct); forewing without free dark spot in cell M3..................................8

  • 8. Small species (forewing length less than 30 mm) (eastern Lesser Sunda islands only)....................................................clementina (Felder, 1860)

  • – Larger species (forewing length usually more than 30 mm)............... 9

  • 9. Upperside forewing with apex of discal cell and extensive postdiscal area whitish, with a broad postdiscal curvilinear dark band that extends to costa; underside of forewing discal cell yellowish-orange at base........... melania (Fabricius, 1775)

  • – Upperside forewing without any extensive white area, almost entirely blue except for darkened apex and the often incomplete, narrow, dark postdiscal curvilinear band, which does not reach costa; underside of forewing discal cell yellowish or grey at base.......................................... celestina (Boisduval, 1832)

  • 10. Ground colour of wings bright golden-yellow aurosa Yata & Vane-Wright sp.n.

  • – Ground colour of wings whitish, at most slightly yellowish; rarely suffused grey ...................................................................... 11

  • 11. Both wings with distinct markings similar to typical female form of paulina (i.e. a more or less broad black border present on the hindwing (Fig. 19G), although sometimes indistinct) (peninsular India only)............... wardii (Moore, 1884)

  • – Hindwings with a few black spots along margin at most (Fig. 19B), thus not like typical female form of paulina .................................... 12

  • 12. Small species (forewing length less than 26.0 mm); forewing underside with curvilinear postdiscal black band running from costa to cell M3, and not reduced to isolated spots (Marianas only).........mariana Yata & Chainey sp.n.

  • – Larger species (forewing length greater than 27.0 mm); forewing underside without continuous black postdiscal band from costa to cell M3; if band present then reduced to isolated spots, or fails to reach costa......................13

  • 13. Both wings without upperside markings, except in some cases a black marginal border to forewing, including apex...............................14

  • – Both wings usually with more extensive dark markings (sometimes reduced to a free black spot in cell M3) on forewing upperside; forewing apex not distinctly acute, somewhat blunt..............................................paulina (Cramer, 1777)

  • 14. Forewing apex distinctly acute, posterior margin almost straight..albina (Boisduval, 1836)

  • – Forewing apex not distinctly acute but often falcate, with outer margin concave...................................................................15

  • 15. Forewing entirely white except for distinct black scales narrowly along margin; upperside ground colour lustrous white (eastern species: New Caledonia to Samoa) ............................................... athama (Blanchard, 1848)

  • – Forewing often with black or grey scales at apex; upperside ground colour matt white, without lustre (Sri Lanka only)................galene (Felder & Felder, 1865)

  • 16. (Females) Upperside ground colour extensively chocolate-brown placidia (Stoll, 1790)

  • – (Females) Upperside not chocolate-brown; ground colour white, yellow, red or bluish-grey, or even almost dark brown or black........................ 17

  • 17. Small species (forewing length typically less than 28 mm), with extensive bluish-grey areas on upperside and a rather narrow band postdiscal white band (Fig. 22N); underside largely brown, but with base of forewing greenish clementina (Felder, 1860)

  • – Not so; if extensively bluish-grey, then forewing underside orange at base18

  • 18. Hindwing upperside almost entirely blue-grey, contrasting with discal area of forewing, which is extensively white; forewing underside orange at base; hindwing underside mainly brown..................................melania (Fabricius, 1775)

  • – Hindwing not contrasting with forewing in this way; underside of hindwing not largely brown or underside of forewing not orange at base............. 19

  • 19. Hindwing white or yellow, without any dark scales at base, but with a broad well-defined and even black posterior band, narrowly edged with white or yellow triangles on margin (Fig. 19R, S) .................. athama (Blanchard, 1848)

  • – Hindwing not so; if with broad black posterior band, the band is more diffusely defined, and lacks the narrow pale margin........................20

  • 20. Hindwing largely dark brownish-black, with a rather narrow discal band and a row of distinct submarginal spots, these areas being either orange-red, yellow or white; the outer margin of the discal band passes through the apex of the discal cell (Fig. 22F, G, H)........................ zarinda (Boisduval, 1836)

  • – Hindwing not so; if both band and submarginal spots present, then outer margin of the postdiscal band passes well beyond the apex of cell, and the submarginal spots are less well defined.................................... 21

  • 21. Upperside ground colour extensively orange or red; if white or yellow, then vein at apex of forewing cell marked with a line of black scales (Figs 21D–F, H, J–N, P–R, 22C) ............................. 22

  • – Upperside ground colour white or yellow (occasionally largely black), without a line of black scales at the apex of the forewing discal cell................ 23

  • 22. Forewing acutely pointed at apex; hindwing distinctly angled at tornus; upperside ground colour always dull orange-red; postdiscal dark band on upperside forewing always clearly separate from end of discal cell, never extending to or touching it (Fig. 21D–F) ....................... galba (Wallace, 1867)

  • – Forewing only bluntly pointed at apex and hindwing only bluntly angled at tornus; upperside ground colour not always dull orange-red, sometimes white or yellow; postdiscal dark band on upperside forewing, if distinguishable, always extends to or touches end of discal cell (Figs 21D–F, H, J–N, P–R, 22C)......................................... nero (Fabricius, 1793)

  • 23. Forewing black apical area with a whitish or yellowish spot in cell R2, although this may be very small or only visible ventrally in occasional examples of A. paulina with very broad black margins (notably from Timor and Sula), but in these cases there is a distinct line of bluish-grey scales along the anterior edge of the black margins on the hindwing, or the black is expanded to leave only a narrow white band.................................................24

  • – Forewing black apical area without a whitish or yellowish spot in cell R2 29

  • 24. Forewing with a large well-defined white or yellow spot in cell M3, contrasting with a very small spot in cell M2; hindwing white or yellow, with a broad black margin that is edged anteriorly with an area of bluish-grey scales celestina (Boisduval, 1832)

  • – Forewing black apical area without a whitish or yellowish spot in cell M3; hindwing often without an area of bluish-grey scales ...................... 25

  • 25. Small species (forewing length usually less than 26.0 mm); forewing underside with black postdiscal band clearly reaching costa, but not touching the apex of the discal cell (although there are diffuse blackish scales in this area) (Fig. 20O–Q) (Marianas only).......mariana Yata & Chainey sp.n.

  • – Larger species (forewing length usually more than 27.0 mm); forewing underside with black postdiscal band often not reaching costa, but does include at least the anterior apex of the discal cell (not from Marianas) ........... 26

  • 26. Forewing black apical area with two or three submarginal pale spots in cells R2, R5 and M1 (peninsular India only)...............................wardii (Moore, 1884)

  • – Forewing black apical area usually with four or five submarginal whitish spots in cells R2, R5 and M1–M3................................................27

  • 27. Forewing underside with black line that curves down to vein M3, where it expands into a pronounced spot with a straight posterior margin; the line then turns to the wing margin at 90°, so that the outer margin is distinctly step-like in appearance (Figs 19J–P, 20T); hindwing with bluish-grey scales either extensive or absent................................................. 28

  • – Forewing underside with outer margin of black line not step-like at M3, i.e. although somewhat jagged in outline, the posterior margin curves gently round to the wing margin (Fig. 20C–E, G–I, K); hindwing upperside often with area of bluish-grey scales along the anterior edge of the black posterior margin (e.g. Fig. 20G); occasionally wings largely black and with more extensive bluish-grey area on hindwing (e.g. Fig. 20I) (widespread, but not found in Mentawais)........................................paulina (Cramer, 1777)

  • 28. Upperside ground colour somewhat male-like, at least partly bluish-grey (Mentawai Islands only)...............................................mata (Kheil, 1884)

  • – Upperside ground colour either white or yellow, usually without bluish-grey (widespread, but not recorded from Mentawais).... albina (Boisduval, 1836) (rare examples only)

  • 29. Forewing black apical area with three submarginal pale spots in cells R5, M1 and M2 (Sri Lanka only)...............................galene (Felder & Felder, 1865)

  • – Forewing black apical area usually with four (but something only three) submarginal pale spots in cells R5 and M1–M3 (widespread, including Sri Lanka)...................................... albina (Boisduval, 1836) (most examples)

Discussion

Species status and interrelationships

According to Corbet & Pendlebury (1992: 89), the species of Appias are difficult to separate by male genitalia, and they note no differences at all between three Appias (Catophaga) dealt with here: nero, albina and paulina (these comments are probably based on dissections made by George Talbot in the 1930s, working in support of Corbet). However, the investigations of Yata (1981), and the many dissections made in Yata's laboratory during this study, reveal small but apparently constant differences in form between the male genitalia of all 15 species recognized here. Moreover, these differences are sufficient to establish the three clusters of species proposed: the paulina complex, the nero group and the melania series. However, there seems little prospect that these small differences can be interpreted further, e.g. to infer monophyly of any of these subgroups. It is interesting to speculate that the rather high ‘phylogenetic inertia’ evident in the male genitalia of these butterflies might indicate that, in this group, sexual selection is acting strongly on male colour pattern, and only weakly or not at all on male genital morphology [see Song & Bucheli (2009) for an interesting assessment of the phylogenetic value of male genitalia characters relative to nongenital characters].

Given the extensive individual variation, polymorphism and geographical variation of many of the species of Appias (Catophaga), coupled with their intrinsic biological interest, the application of molecular characters to investigate the subgenus would be very desirable. This would permit the status of many populations over which doubt remains to be re-assessed, such as the validity of separating A. mata from A. paulina. Equally important, molecular data will probably prove invaluable in the quest to produce a robust cladistic analysis, necessary to gain further insight into the sequence of colour pattern and hostplant changes that appear to have affected these species during their evolution (see below and File S1).

Distribution

The subgenus Appias (Catophaga) occurs across almost the whole of the Indo-Australian tropics, from Sri Lanka through extreme southern China to the Marianas, the Malay Archipelago to north-eastern Australia, and island groups of the West Pacific, from New Caledonia to Samoa (Figs 23, 24). Subgenus Appias (Catophaga) is not represented in north-western India, Pakistan or, rather curiously, beyond Bougainville in the main Solomons chain, where only one member of the genus occurs, A. (Appias) ada (Stoll, 1781; Tennent, 2002).

The collective distribution of the eight species of the paulina complex covers the same area as Appias (Catophaga) as a whole, with A. albina and A. paulina being by far the most widespread members of the subgenus. The other six species of the paulina complex are much more restricted: A. galene (Sri Lanka; sympatric with albina), A. wardii (Western Ghats; sympatric with albina), A. mata (Nias and Sipora; possibly sympatric with albina, which has been recorded from neighbouring Siberut), A. aurosa (southern and central Sulawesi; sympatric with albina and paulina), A. mariana (Marianas; isolated) and A. athama (western Pacific, from New Caledonia to Samoa; isolated in Samoa and Fiji, but sympatric with paulina in New Caledonia and Vanuatu; Tennent, 2009). Because of earlier taxonomic confusion, in some works it appears as if two species belonging to the paulina complex occur in Fiji (e.g. Evenhuis, 2007), and as many as three in Samoa (e.g. Kami & Miller, 1998: 70), but, as indicated correctly by Tennent (2006: 31), only one taxon occurs at this eastern limit: A. athama manaia.

The nero group of three species is restricted essentially to the Oriental Region, from north-eastern India through Indochina to Hainan, and south through the Philippines and the Malay Archipelago to Bali, Sulawesi and Buru. The three species, so far as known, are parapatric, although the demarcation between A. nero and A. galba in the Indochinese region is poorly understood (Fig. 24). Appias zarinda is an insular species, being virtually endemic to the Sulawesi Region [as demarcated by Vane-Wright & de Jong (2003: fig. 1, 109)]. Its eastward extension to Buru (Central Maluku: Peggie et al., 1995) is a little puzzling because, on biogeographical grounds (Vane-Wright & Peggie, 1994: 228), one might expect Buru to be occupied by A. placidia (melania series). However, as discussed by Vane-Wright (1991), the southern Moluccas, including Buru, may be linked geologically to the Sula and Banggai archipelagos of central-eastern Sulawesi. This string of little islands plausibly formed a series of ‘stepping stones' that apparently allowed certain species of butterflies evolved in Sulawesi to spread greater or lesser distances eastward, into the Moluccas, New Guinea and even further east, from the time when the Asian and Australian plates collided approximately 15 million years ago (Vane-Wright, 1991). If so, A. zarinda could represent a Sulawesi species that has spread along this route only as far as Buru, where it has formed a distinct subspecies (A. z. bouruensis).

The melania series of four allopatric species occurs east of Weber's Line, and is thus restricted to the Australian Region. Appias placidia occurs in both northern and central Maluku, which, based on a ‘nearest-neighbour’ analysis of butterfly distributions, constitute two separate areas of endemism (Vane-Wright & Peggie, 1994). However, as already noted, placidia is surprisingly absent from Buru, where it appears to be replaced by A. zarinda (nero group). Appias clementina is restricted to a group of islands in the western Lesser Sundas, from Alor and Timor to Tanimbar. Appias celestina occurs in the New Guinea Region, including Kai and Aru, and thus could be considered parapatric with respect to clementina. Appias celestina also occurs on Cape York Peninsula (Queensland), whereas the final species in the series, A. melania, is confined to a relatively small area of Queensland lying approximately 400 km further south.

In summary, only the highly polytypic A. paulina and A. albina are widely distributed through much of the Indo-Australian Region, where they mostly overlap with other, discontinuously distributed local members of the paulina complex, and the species of the nero group and melania series. Four of the restricted members of the paulina complex occur in the Oriental Region, and two in the Pacific. The three species of the nero group are distributed essentially parapatrically through a large part of the eastern Oriental Region. The four members of the melania series are distributed allopatrically in part of the Australian Region.

Hostplant relationships, coloration, polymorphism and speciation

Brief discussions of larval hostplant relationships (with special reference to an apparent shift from Brassicales to Malpighiales) and adult pigmentation and coloration, together with a more extensive discussion of adult polymorphism in these butterflies and its possible relationship to Darwinian colour-pattern transference and speciation, are included in File S1. These more speculative elements do not contribute greatly to the basic systematics. Even so, we consider the polymorphism of these species, especially A. nero, to be of outstanding biological and evolutionary interest. In fact, so much so that they provided a major stimulus to completing the present study.

Suggestions for future work

As already discussed, a comprehensive investigation into the molecular systematics of the species and subspecies of Appias (Catophaga) would be basic for comparative biology on these butterflies, including the evolution of coloration within the group (see discussion in File S1). The availability of a fully resolved, robust cladogram is, ultimately, the sine qua non for this type of evolutionary study. However, getting fresh material for DNA work would be difficult in some cases, e.g. the narrowly distributed and rarely encountered A. aurosa, or even impossible in the case of A. mariana, if it really is extinct. A particularly interesting question that could be resolved by molecular work concerns the relationship between A. zarinda and A. placidia. Our morphological investigations suggest that these two species belong to different groups within the subgenus Appias (Catophaga). However, the fact that A. zarinda extends east to Buru (A. z. bourouensis), where A. placidia does not occur but might have been expected, implies the possibility of a closer phylogenetic relationship and/or very similar ecological requirements for these two taxa. Conceivably the nero group is nested within the melania series, with the possibility that A. placidia represents its sister group.

The formal genetics of any of the species and races exhibiting female polymorphism would be of great interest, especially with respect to the complex situation seen in some populations of A. paulina, which may have as many as six female forms. Perhaps most interesting of all would be to investigate the genetic dominance relationships affecting A. nero palawanica, to ascertain whether or not the orange, male-like female morphs of A. nero on Palawan are dominant to the white, piebald and yellow female forms. This seems probable if the male-coloured morphs, seen in many but not all races of A. nero, are the result of Darwinian transference (see discussion in File S1). The generality of this result could be checked by an investigation into the genetics of A. zarinda on Sulawesi, where the females of this species also are coloured white, yellow or male-like orange. Racial and even species hybrids, if they can be created in the laboratory, might also offer valuable insights. Renewed interest in the pigment chemistry (see File S1) of these butterflies might be particularly revealing in this context, taking care to investigate females as well as males. Any programme of genetic research would be facilitated by or even depend also upon much better information about the hostplants and general ecology of Appias (Catophaga) species – a need that amateur entomologists could play a key role in satisfying (Vane-Wright, 2009).

Finally, a recent paper on the biochemistry of Pieridae has revealed that various members of the family produce proteins (‘pierisins’) toxic to certain human cancer cells. Within Appias, pierisins have been found in A. nero and A. paulina, but not in A. lyncida (Matsumoto et al., 2008). Whether these findings have any systematic or ecological significance for our understanding of Appias butterflies is unknown, but these discoveries add another dimension for future research.

Acknowledgements

In completing this joint project, the beginnings of which stretch back over 30 years, OY has been primarily responsible for the descriptive accounts and key to species, JEC has been primarily responsible for typification, the taxonomic appendices, refining the key to females, and re-curating the entire BMNH Catophaga collection (thereby underpinning much of the work as a whole), whereas RIVW has been primarily responsible for the discussion sections, and assembly of the final manuscript. At the Natural History Museum in London we were greatly assisted by, among others, Phillip Ackery, Campbell Smith and Blanca Huertas with access to material and provision of loans; members of the museum's photography unit generously prepared numerous high-quality digital images. Curators at Oxford, Honolulu and Paris (notably the late Georges Bernardi) kindly gave direct access to their collections, or made important material available on loan. John Tennent (U.K.), Andrew Rawlins (U.K.), Min Wang (South China Agricultural University), Djunijanti Peggie (Research Center for Biology, Indonesian Institute of Sciences) and Alexander Monastyrskii (Russia Vietnam Tropical Research Center) gave access to specimens, and provided information on distributions. In Japan, Osamu Tadauchi (Entomological Laboratory, Faculty of Agriculture, Kyushu University), Yasusuke Nishiyama (Tokyo), Etsuzo Tsukada (Yamanashi), Kazuhiko Morishita (Kanagawa), Morio Wakabayashi (Osaka), Hiroaki Onodera (Saitama), Masaya Yago (Tokyo University), Kyoichiro Ueda (Kitakyushu Museum of Natural History and Human History), and staff at the Museum of Nature and Human Activities (Hyogo) and Osaka Museum of Natural History, greatly assisted us through the provision of valuable materials and other information; Ken-Ichi Odagiri (BLKU), Yasuhiro Ohshima (Tochigi Prefectural Museum), Hiroaki Iwasaki (BLKU) and Kei Hashimoto (BLKU) helped us by taking photographs and assisting with the preparation of genitalia figures. Carol Boggs, Ward Watt, Henri Descimon and Michael Braby guided us with access to specific literature, and Haruo Fukuda (Kagoshima) generously shared unpublished information. Michael Boppré (Freiburg) very kindly compiled the colour plates, for which we are most grateful. The Royal Entomological Society's Charles P. Alexander Bequest Fund generously helped defray the costs of colour reproduction. Alexander Monastyrskii, Fred Nijhout, Peter Cranston and especially Michael Braby and Gerardo Lamas made numerous helpful suggestions for correcting and improving the manuscript, most of which we have gratefully adopted. Finally, OY wishes to thank Toyohei Saigusa and the late Takashi Shirôzu (Fukuoka) for their constant guidance and encouragement during the initial stages of this work.

Appendices

Appendix 1

Synonymic checklist of species and subspecies of Appias (Catophaga)

Those taxa for which we have examined type material are marked with an asterisk.

Appias (Catophaga) albina (Boisduval, 1836)

Pieris albina Boisduval, 1836.

Eight subspecies, ranging from Sri Lanka, Western Ghats, north-eastern India, southern China, South-East Asia and Philippines through Malay Peninsula to Papua New Guinea and northern Australia.

albina swinhoei * (Moore, 1905). Catophaga swinhoei. Type locality: Southern India: ‘Nilgiris’ (lectotype here designated: see Appendix 2).

= norma* Evans, 1924. Appias albina f. norma. Sri Lanka ‘Ceylon’.

Distribution: Southern India and Sri Lanka.

albina darada * (Felder & Felder, 1865). Pieris darada. Type locality: Bangladesh: ‘Silhet’.

= confusa*Fruhstorfer, 1910. Appias albina confusa (junior secondary homonym of Belenois confusa Butler, 1872). Type locality: India, Thailand, Vietnam: ‘Sikkim, Assam, Siam, Tonkin’.

Distribution: India (north-eastern region, Andaman and Nicobar islands), Bangladesh, southern China (Yunnan, Hainan), Myanmar, Vietnam, Laos, Cambodia and Thailand.

albina semperi (Moore, 1905). Catophaga semperi. Type locality: northern Philippines: (no further data).

= kawakamii Matsumura, 1909. Appias kawakamii. Type locality: Taiwan: ‘Kotosho I’ (Lanyu Island).

Distribution: Philippines (Babuyan, Bohol, Cebu, Dumaran, Guimaras, Luzon, Marinduque, Mindanao, Negros) and southern Taiwan (Lanyu Island).

albina agatha (Staudinger, 1889). Tachyris agatha. Type locality: Philippines: [N.] ‘Palawan’.

= pancheia*Fruhstorfer, 1910. Appias albina pancheia. Type locality: Philippines: [N.] ‘Palawan’.

Distribution: Philippines (Mindanao, northern Palawan).

albina albina (Boisduval, 1836). Pieris albina. Type locality: Indonesia, Central Maluku: ‘Ambon’.

= neombo*Boisduval, 1836. Pieris neombo. Type locality: Indonesia, Central Maluku: [Ambon] ‘Brésil, environs de Bahia et Fernamboue’ (error).

= cynisca*Wallace, 1867. Tachyris cynisca. Type locality: Indonesia, Central Maluku, Buru: ‘Bouru’. Syn.n.

Distribution: Malay Peninsula, Indonesia (Sumatra, West Java, Kalimantan, Sulawesi, Moluccas, Irian Jaya), Sabah, Brunei, Sarawak, Philippines (Balabac, Bongao, central and southern Palawan, Sanga Sanga, Sibutu, Tawitawi), Papua New Guinea (including Kiriwina Island), Australia (Darwin, Cobourg Peninsula, a few localities in Northern Territories, and Rimbija, Moa, Thursday and Prince of Wales islands).

albina ambigua * Grose-Smith, 1895. Appias ambigua. Type locality: Indonesia, South Maluku, Wetar. ‘Wetter’ (lectotype here designated: see Appendix 2).

= micromalayana*Fruhstorfer, 1909. Appias albina micromalayana. Type locality: Indonesia: ‘East Java’ (lectotype here designated: see Appendix 2). Syn.n.

Distribution: Indonesia [East Java, Lesser Sunda Islands (not Sumbawa) east to Tanimbar], Christmas Island and East Timor.

albina infuscata * Fruhstorfer, 1910. Appias albina infuscata. Type locality: Indonesia: ‘Sumbawa’.

Distribution: Indonesia (Sumbawa).

albina pulverobasalis Joicey & Noakes, 1915. Type locality: Indonesia, Biak, Bosnik.

Distribution: Indonesia (Irian Jaya: Biak).

Infrasubspecific and unavailable names: alcesta*Talbot, 1939; citrinaFruhstorfer, 1910; citronella* Fruhstorfer, 1897; flava Röber, 1891; latimarginata Matsumura, 1919; matsumurai Sonan, 1930; principalis*Fruhstorfer, 1910; punctata*Fruhstorfer, 1910; saweloides*Fruhstorfer, 1910; semiflava*Fruhstorfer, 1910; subochracea*Fruhstorfer, 1910; umbratilis Fruhstorfer, 1897; virilis*Fruhstorfer, 1910.

Appias (Catophaga) athama (Blanchard, 1848)

Pieris athama Blanchard, 1848.

Until recently this taxon has been treated as a subspecies of Appias albina or A. paulina. With separation, its division into subspecies requires re-investigation; three are provisionally recognized here. Females vary from deep yellow on New Caledonia to cream-coloured in Samoa. Appias athama occurs in New Caledonia, Vanuatu, Tonga, Western Samoa, American Samoa and Fiji.

athama psyche * (Felder & Felder, 1865). Pieris psyche. Type locality: New Caledonia: ‘Nova Caledonia’.

= *argentiferaJoicey & Talbot, 1928. Type locality: New Caledonia ‘Loyalty Is'; (lectotype here designated: see Appendix 2).

Distribution: New Caledonia (including Isle of Pines); status of Lifu population uncertain.

athama athama * (Blanchard, 1848). Pieris athama. Type locality: Fiji, Ovalau: ‘Balaou’. (Type material in MNHN).

= athama*Lucas, 1852. Pieris athama. Type locality: Fiji, Ovalau: ‘Balaou (Nouvelle-Guinée)’ (junior objective synonym and primary homonym of Pieris athama Blanchard).

= jacquinotii*Lucas, 1852. Pieris jacquinotii. Type locality: Fiji, Ovalau: ‘Balaou (Nouvelle- Guinée)’.

= wallacei* (Butler, 1898). Catophaga wallacei. Type locality: Vanuatu, Malekula: ‘New Hebrides (Mellicollo)’ (lectotype here designated: see Appendix 2)

Distribution: Vanuatu (Mota Lava, Gaua, Espiritu Santo, Ambae, Malakula, Paama, Epi, Efaté, Erromango, Aniwa, Futuna, Aneityum), Fiji (Ongea, Viti Levu, Vanua Valava, Totoya, Lau, Walsala, Yasawa, Naviti, Ovalau).

athama manaia * (Hopkins, 1927). Catophaga jacquinotii manaia. Type locality: Western Samoa: ‘Upolu I.’

Distribution: Tonga (Vava'u), Western Samoa (Savai'I, Upolu), American Samoa (Tutuila).

Appias (Catophaga) aurosa* Yata & Vane-Wright sp.n.

Type locality: Indonesia, Sulawesi: Makassar.

Distribution: Known from approximately 20 males collected in southern and central Sulawesi (Indonesia).

Infrasubspecific and unavailable names: aurosa*Fruhstorfer, 1899.

Appias (Catophaga) celestina (Boisduval, 1832)

Pieris celestina Boisduval, 1832.

Five subspecies, ranging from Aru and Kai through New Guinea to the Bismarck Archipelago, Bougainville and Cape York.

celestina galepsus * Fruhstorfer, 1910. Appias celestina galepsus. Type locality: Indonesia: ‘Key Is'.

Distribution: Indonesia (Kep. Kai) only.

celestina barea * Fruhstorfer, 1910. Appias celestina barea. Type locality: Indonesia: ‘Aru’.

Distribution: Indonesia (Kep. Aru) only.

celestina celestina (Boisduval, 1832). Pieris celestina. Type locality: Indonesia, Irian Jaya: ‘Dorei’.

= sekarensis (Ribbe, 1886). Tachyris celestina var. sekarensis. Type locality: Indonesia, Irian Jaya: ‘Sekar, West coast of New Guinea’.

= galerus*Fruhstorfer, 1910. Appias celestina galerus. Type locality: Papua New Guinea: ‘German New Guinea’.

= tamibia Strand, 1912. Appias celestina var. tamibia. Type locality: not located: ‘Tami area’.

Distribution: Indonesia (Aru, Waigeo, Irian Jaya), Papua New Guinea (mainland), Australia (Cape York; probably vagrants from New Guinea).

celestina eumelis (Boisduval, 1832). Pieris eumelis. Type locality: Papua New Guinea: ‘Port-Praslin (Nouvelle-Hollande)’ (=New Ireland).

= delicata* Butler, 1882. Appias delicata. Type locality: Papua New Guinea: Stated to be from ‘New Britain’, but type specimen is labelled Duke of York I.

Distribution: Papua New Guinea: New Ireland, New Britain, Duke of York.

celestina orientalis * Rothschild, 1915. Appias celestina orientalis. Type locality: Papua New Guinea: ‘Bougainville, Arawa’.

Distribution: Papua New Guinea: Bougainville, Normanby (Normanby may be subsp. celestina).

Infrasubspecific and unavailable names: aurifera*Fruhstorfer, 1910; flava* Ribbe, 1886; lileia*Fruhstorfer, 1910, limia*Fruhstorfer, 1910; typicumTalbot, 1932 (refers to the nominotypical female form; even if accepted as a name, it is clearly infrasubspecific).

Appias (Catophaga) clementina (Felder, 1860)

Pieris clementina C. Felder, 1860. Type locality: Indonesia: ‘Ambon’ (presumed error, although there is a female in BMNH labelled ‘Amboina Fruhstorfer’)

= adelpha (Röber, 1891). Tachyris adelpha. Type locality: Indonesia: ‘Alor’.

= agar*Fruhstorfer, 1910. Appias clementina agar. Type locality: Indonesia: ‘Dammer, Babber, Tenimber’.

Distribution: This small Indonesian species is confined to a group of islands in the eastern Lesser Sunda Islands (Alor, Damar, Babber, Tenimber). It does not appear to be divisible into subspecies.

Appias (Catophaga) galba (Wallace, 1867 stat. rev.)

Tachyris galba * Wallace, 1867. Type locality: India: ‘N. India’.

= nebo Grose-Smith & Kirby, 1894. Appias nebo. Type locality: Upper Burmah, Chindwin River.

= hainanensis* Fruhstorfer, 1902. Tachyris nero hainanensis. Type locality: China: ‘Hainan, Five Finger Mountains'.

Distribution: Until now treated as a subspecies of A. nero, this butterfly occurs in northern India, Myanmar and south-eastern China (Hainan), south to Vietnam, Laos and northern Thailand.

Appias (Catophaga) galene* (Felder & Felder, 1865)

Pieris galene * (Felder & Felder, 1865). Type locality: Sri Lanka: ‘Ceylon, Trincomali’.

= lankapura* Moore, 1879. Catophaga lankapura. Type locality: Sri Lanka: ‘Ceylon’.

= venusta* Moore, 1881. Catophaga venusta. Type locality: Sri Lanka: ‘Vavonia Vivankulam, N. of Trincomali’.

Distribution: Restricted to Sri Lanka, where it occurs widely except in the extreme north.

Infrasubspecific and unavailable names: fasciata*Fruhstorfer, 1910.

Appias (Catophaga) mariana Yata & Chainey sp.n.

Type locality: U.S.A.: ‘Guam’

Distribution: This newly described species, known from three islands in the Marianas (Guam, Rota, Saipan), may be extinct.

Appias (Catophaga) mata (Kheil, 1884)

Pieris mata Kheil, 1884

This rare Indonesian butterfly is restricted to the Mentawi Islands, off the south-west coast of Sumatra, where it occurs as two distinct subspecies.

mata mata (Kheil, 1884). Tachyris mata. Type locality: Indonesia, Mentawi Islands: ‘Nias’.

Distribution: Known only from Nias.

mata caeca * Corbet, 1941. Appias paulina caeca. Type locality: Indonesia, Mentawi Islands: ‘Sipora’.

Distribution: Known only from Sipora.

Appias (Catophaga) melania (Fabricius, 1775)

Papilio melania * Fabricius, 1775. Type locality: Australia: ‘New Holland’ (Queensland, Cooktown).

= asteria Miskin, 1889. Tachyris asteria. Type locality: Australia: ‘N. Queensland, Port Douglas' (type material in South Australian Museum, Adelaide).

= cerussa Fruhstorfer, 1904. Tachyris cerussa. Type locality: Australia: ‘Queensland’ (whereabouts of type material unknown).

Distribution: Australian endemic with its breeding range restricted to north-east coastal region of Queensland, where it occurs in upland rainforests from Cooktown south to the Bluewater State Forest.

Appias (Catophaga) nero (Fabricius, 1793)

Papilio nero * Fabricius, 1793

Fifteen recognized subspecies, some very distinct, ranging from the Malay Peninsula to Bali, southern Sulawesi region, Borneo and the Philippines.

nero nero * (Fabricius, 1793). Papilio nero. Type locality: (Indonesia, Java): ‘Asia’ (neotype here designated, see Appendix 2).

= thyria Godart, 1819. Pieris thyria. Indonesia: ‘Java’.

= figulina* Butler, 1867. Pieris figulina. Type locality: Malaysia: ‘Singapore’ (lectotype here designated, see Appendix 2).

Distribution: Malay Pensinsula, Sumatra, Java, Bali.

nero ramosa * (Fruhstorfer, 1898). Catophaga nero ramosa. Type locality: Indonesia: ‘Nias’.

Distribution: Indonesia (Nias).

nero pulonus * (Fruhstorfer, 1906). Tachyris nero pulonus. Type locality: Indonesia: ‘Batu’.

Distribution: Indonesia (Batu).

nero baweanicus * (Fruhstorfer, 1905). Tachyris nero baweanicus. Type locality: Indonesia: ‘Bawean’.

Distribution: Indonesia (Bawean).

nero neronis * (Fruhstorfer, 1903). Tachyris nero neronis. Type locality: Indonesia: ‘Kangean’.

Distribution: Indonesia (Kangean).

nero acuminata (Snellen, 1890). Pieris nero var. acuminata. Type locality: Indonesia: ‘Tanah-Djampea’.

Distribution: Indonesia (Tanahjampea).

nero chelidon * (Fruhstorfer, 1905). Tachyris nero chelidon. Type locality: Sabah: ‘North Borneo’.

Distribution: Indonesia (Kalimantan), Sabah, Brunei, Sarawak.

nero flavius * Grose-Smith, 1892. Appias flavius. Type locality: Philippines: ‘Taganac I., NE Borneo’.

Distribution: Philippines: Turtle Islands (off the coast of Borneo, near Sandakan).

nero palawanica (Staudinger, 1889). Tachyris nero var. palawanica. Type locality: Philippines: ‘Palawan’.

Distribution: Philippines (Balabac, Palawan).

nero corazonae Schröder & Treadaway, 1989. Appias nero corazonae. Type locality: Philippines: ‘Sulu Archipelago’.

Distribution: Philippines (Bongao, Sanga Sanga, Sibutu).

nero fleminius Fruhstorfer, 1911. Appias nero fleminius. Type locality: Philippines: ‘Mindoro’.

Distribution: Philippines (Mindoro).

nero domitia * (Felder & Felder, 1862). Pieris domitia. Type locality: Philippines: ‘Luzon’.

= asterope Felder & Felder, 1862. Pieris asterope. Type locality: Philippines: ‘Luzon’ (junior primary homonym of Pieris asterope Godart, 1819).

= korridona*Grose-Smith 1894. Appias korridona. Type locality: Indonesia, Irian Jaya: ‘Korrido’ (presumed error). Syn.n.

= yamazakii Sonan, 1936. Appias nero yamazakii. Type locality: Taiwan: ‘Formosa’.

Distribution: Philippines (Luzon, Marinduque, Masbate), Taiwan.

nero soranus Fruhstorfer, 1910. Appias nero soranus. Type locality: Philippines: ‘Cebu’.

Distribution: Philippines (Cebu, Negros, Panay, Sibuyan).

nero zamboanga (Felder & Felder, 1862). Pieris zamboanga. Type locality: Philippines: ‘Mindanao, Celebes' (type locality here restricted to Mindanao. Lectotype designation required, but no type material located).

= mindanensis* Butler, 1883. Appias mindanensis. Type locality: Philippines: ‘Mindanao, Pasananca valley’.

= boholensis Okano & Okano, 1989. Appias nero boholensis. Type locality: Philippines: ‘Visayan Islands'.

Distribution: Philippines (Bohol, Dinagat, Leyte, Mindanao, Panaon, Samar).

nero tibericus * Fruhstorfer, 1910. Appias nero tibericus. Type locality: Philippines: ‘Bazilan’.

Distribution: Philippines (Basilan).

nero subsp. near zamboanga (Vane-Wright & de Jong, 2003: 109)

Distribution: Indonesia, Kep. Talaud (Talaud, Salebabu).

Infrasubspecific and unavailable names: coelitus*Fruhstorfer, 1899; helvola*Fruhstorfer, 1910; sufflava*Fruhstorfer, 1910.

Appias (Catophaga) paulina (Cramer, 1777)

Papilio paulina Cramer, 1777.

Nearly two dozen recognized races make up this wide-ranging species, found from north-east India to Vanuatu. As commented above, some of these taxa may eventually prove to be distinct species.

paulina adamsoni * (Moore, 1905). Catophaga adamsoni. Type locality: Myanmar: ‘Burma, Upper Tenasserim, Thoungying Valley; Dounat Range; Muong Gnow, Shan States; Rangoon; Arrakan; Hlaingbeoo Arrakan’.

= pseudoleis*Fruhstorfer, 1910. Appias melania pseudoleis. Type locality: Thailand: ‘Siam’.

Distribution: North-east Indian region to south-western China, Myanmar, Vietnam, Laos, Cambodia and Thailand.

paulina griseoides * Moulton, 1923. Appias paulina griseoides. Type locality: Vietnam: ‘Pulo Condore (south-east coast of Cochin-China)’.

Distribution: Known only from Pulo Condore in the Con Son Islands, off the coast of southern Vietnam.

paulina subsp. (A. Rawlins, personal communication, 2009).

Distribution: Indonesia (Natuna Islands).

paulina minato * (Fruhstorfer, 1899). Catophaga paulina minato. Type locality: Japan: ‘Ishigaki’.

= iwasakii Matsumura, 1919. Appias melania var. iwasakii. Type locality: Japan, Okinawa: ‘Yayeyama’.

Distribution: Taiwan and Japan (Yaeyama Is north to Akusekijima).

paulina galathea (Felder, 1862). Pieris galathea. Type locality: India, Andaman Is: ‘Sambelong’.

= roepstorffiiMoore, 1884. Catophaga roepstorffii. Type locality: India: ‘Nicobar Is.’

Distribution: Confined to the Andaman and Nicobar Islands.

paulina distanti * (Moore, 1905). Catophaga distanti (as ‘nom. n.’ for leis of Distant, 1885; de Nicéville & Martin, 1895). Type locality here restricted to Malaysia ‘Malay Pen.’, but no type material from this locality is available for lectotype designation.

Distribution: Confined to the Malay Peninsula, including Langkawi Islands.

paulina grisea * Moulton, 1923. Appias paulina grisea. Type locality: Malaysia: ‘Pulo Aur; Pulo Tioman’.

Distribution: This subspecies is confined to the small group of islands that includes Aur and Tioman, off the east coast of the Malay Peninsula.

paulina pione * Fruhstorfer, 1910. Appias melania pione. Type locality: Indonesia: Sumatra.

Distribution: Sumatra only.

paulina paulina * (Cramer, 1777). Papilio paulina. Type locality ‘Java’ (lectotype here designated, see Appendix 2):

= leis Geyer, 1832. Catophaga leis. Type locality: Indonesia: Java.

= idaLucas, 1852. Pieris ida. Type locality: Indonesia: Java.

= subtusluteaRoepke, 1935. Appias leis f. subtuslutea. Type locality: Indonesia: Java.

= raksasa Kalis, 1941. Appias raksasa. Type locality: Indonesia: Bali.

= alope*Wallace, 1867. Tachryis alope. Type locality: here restricted to Java, but no type material from this locality available for lectotype designation.

Distribution: Indonesia: Java, Bali and Bawean.

paulina agave (Felder & Felder, 1862). Pieris agave. Type locality: Philippines: ‘Luzon’.

Distribution: Philippines: Dinagat, Homonhon, Leyte, Luzon, Marinduque, Mindanao, Negros, Panaon and Samar.

paulina athena * Fruhstorfer, 1903. Appias melania athena. Type locality: Sabah: ‘Mt. Kinabalu, N. Borneo’.

Distribution: Known from the whole of Borneo, and Bongao, Sanga Sanga, Sibutu and Tawitawi in the Philippines.

paulina nikomedia * Fruhstorfer, 1910. Appias paulina nikomedia. Type locality: Philippines: ‘Basilan’.

Distribution: Confined to the island of Basilan, Philippines.

paulina plaetoria * Fruhstorfer, 1910. Appias melania plaetoria. Type locality: Philippines: Balabac.

Distribution: Confined to the island of Balabac, Philippines.

paulina sithonia Fruhstorfer, 1911. Appias melania sithonia. Type locality: Philippines: Mindoro.

Distribution: Confined to the island of Mindoro, Philippines.

paulina terentilia * Fruhstorfer, 1910. Appias melania terentilia. Type locality: Philippines: ‘Palawan’.

Distribution: Palawan and Calamian, in the Philippines.

paulina thyre Fruhstorfer, 1911. Appias melania thyre. Type locality: Banguey I

Distribution: Banguey I, off Sabah, Malaysia.

paulina albata (Hopffer, 1874). Tachyris albata. Type locality: Indonesia, Sulawesi: ‘Celebes’.

= dohertyi* Rothschild, 1892. Appias dohertyi. Type locality: Indonesia, Sulawesi: ‘southern Celebes'.

= kalisi Röber, 1940. Pandemos (Saletara) melania form kalisi. Type locality: Indonesia, Sulawesi: ‘Süd-Celebes (Bantimurung) und Mittel-Celebes (Tanah Metah)’.

Distribution: Indonesia: Sulawesi, Buton, Muna, Kabaena, Kep. Banggai (Peleng), Kep. Sula (Mangole, Taliabu).

paulina pietersi Kalis, 1933. Appias melania pietersi. Type locality: Indonesia: Kangean.

Distribution: Indonesian island of Kangean only.

paulina sawela * (Fruhstorfer, 1896). Tachyris sawela. Type locality: Indonesia: ‘Lombok’.

Distribution: Indonesian islands of Lombok and Flores.

paulina tambora * (Fruhstorfer, 1903). Catophaga melania tambora. Type locality: Indonesia: ‘Tambora, Sumbawa’.

Distribution: Indonesian island of Sumbawa only.

paulina emilia * (Fruhstorfer, 1903). Catophaga melania emilia. Type locality: Indonesia: ‘Sumba’.

Distribution: Indonesian island of Sumba only.

paulina eurosundana * Grose-Smith, 1895. Appias eurosundana. Type locality: Indonesia, Timor: ‘Oinainisa (Timor)’. Lectotype here designated (see Appendix 2).

Distribution: Indonesian Timor and East Timor.

paulina paula (Röber, 1891). Tachyris paula. Type locality: Indonesia: ‘Wetter’.

Distribution: Indonesian islands of Wetar and Roma.

paulina subsp.

= ‘cynisca’ auctt. (e.g. Peggie et al., 1995) nec Wallace.

Distribution: Indonesian island of Buru (Central Maluku) only.

paulina antoniae * Fruhstorfer, 1910. Appias melania antoniae. Type locality: Indonesia, Central Maluku: ‘East Ceram, Saparoea’.

Distribution: Central Maluku (Indonesia): Seram, Ambon, Kelang, Haruku and Saparua.

paulina zoe (Snellen van Vollenhoven, 1865). Pieris zoe. Type locality: Indonesia, North Maluku: ‘Batjan’.

Distribution: North Maluku (Indonesia): Obi, Halmahera, Bacan, Morotai and Kasiruta.

paulina saina * Grose-Smith, 1894. Appias saina. Type locality: Indonesia, Irian Jaya: ‘Humboldt Bay, Dutch New Guinea’. Lectotype here designated (see Appendix 2).

= sosias* Fruhstorfer, 1913. Appias melania sosias. Type locality: Papua New Guinea: ‘Yule I.’

Distribution: New Guinea (Indonesia and Papua New Guinea).

paulina falcidia * Fruhstorfer, 1910. Appias melania falcidia. Type locality: Indonesia: ‘Biak’.

Distribution: Island of Biak only, in Irian Bay (Indonesia).

paulina ega * (Boisduval, 1836). Pieris ega. Type locality: Australia: ‘Nouvelle-Hollande’.

= caledonica Felder, 1862. Pieris melania var. caledonica. Type locality: New Caledonia.

Distribution: Australia (northern and eastern regions, Groote Eylandt, Darnley, Thursday, Moa, Sue, Lord Howe), New Caledonia (Isle of Pines, Lifu, Ouvée, Maré), Vanuatu (Espiritu Santo, Efaté, Tanna, Futuna, Aneityum).

Infrasubspecific and unavailable names: aegina*Fruhstorfer, 1910; flaminia*Fruhstorfer, 1910;

furia * Fruhstorfer, 1910; horatia*Fruhstorfer, 1910; iria*Fruhstorfer, 1910; limbata*Fruhstorfer, 1910; marginata*Fruhstorfer, 1910; melanides*Fruhstorfer, 1910; obscurior*Fruhstorfer, 1910; ochracea Moulton, 1914; shima Sonan, 1930; uranides*Fruhstorfer, 1910.

Appias (Catophaga) placidia (Stoll, 1790)

Papilio placidia Stoll, 1790. Type locality: Indonesia, Central Maluku: ‘Ambon’.

= maculata* Staudinger, 1884. Tachyris placidia var. maculata. Type locality: Indonesia, North Maluku: ‘Bacan’.

Distribution: This distinctive Indonesian species is known from North (Bacan, Halmahera, Obi) and Central Maluku (Ambon, Seram) only. As suggested by D’Abrera (1971), the separation of the northern Maluku populations as subspecies maculata appears, on exophenotypic grounds, untenable.

Appias (Catophaga) wardii (Moore, 1884)

Catophaga wardii * Moore, 1884. Type locality: India: ‘Coonoor, Nilgiris'.

= arida*Talbot, 1939. Appias wardi [sic] arida. Type locality: India: ‘Nilgiri Hills'.

Distribution: Restricted to the Western Ghats of peninsular India, where it occurs in scattered localities throughout much of the range south of latitude 20°N.

Appias (Catophaga) zarinda (Boisduval, 1836)

Pieris zarinda Boisduval, 1836.

An Indonesian species restricted to the Sulawesi region and the adjacent island of Buru (Central Maluku). Four recognized subspecies, one of which must be considered doubtful.

zarinda zarinda (Boisduval, 1836). Pieris zarinda. Type locality: Indonesia: ‘Java’ [recte Sulawesi].

= fatime Snellen van Vollenhoven, 1866. Pieris fatime. Type locality: Indonesia: ‘Celebes’.

Distribution: Sulawesi and some adjacent islands to the south and east: Kabaena, Kep. Tukangbesi, Kep. Banggai (Peleng).

zarinda sulana * (Fruhstorfer, 1899). Tachyris nero sulana. Type locality: Indonesia, Maluku: ‘Sula, Mangoli’.

Distribution: Known only from Mangole in Kep. Sula, eastern Sulawesi region; doubtfully distinct from A. zarinda zarinda.

zarinda phestus * (Westwood, 1888). Tachyris phestus. Type locality: Indonesia, Sulawesi: ‘Talissa I, N. Celebes' (type material in UM Oxford; seen by RIVW).

Distribution: Kep. Talisei and Kep. Sangihe, in northern Sulawesi region.

zarinda bouruensis (Wallace, 1867). Tachyris bouruensis. Type locality: Indonesia, Central

Maluku: ‘Buru’ [no type specimen located. Butler (1898) states ‘the type should be in Hewitson's collection, but was probably not in good enough condition to induce him to retain it'].

Distribution: Buru island (western Central Maluku) only.

Taxa excluded from Appias subgenus Catophaga

Appias (Hiposcritia) urania * (Wallace, 1867) (described in Tachyris)

= zondervani Toxopeus, 1950

Appias (Appias) libythea rouxii (Boisduval)

= yaksha*Fruhstorfer, 1910: 157

Appias (Appias) ada (Stoll, 1781)

Appendix 2

Nine lectotype designations, one neotype designation, and two type-locality restrictions for 12 nominal species of the genus Appias.

Catophaga swinhoei Moore, 1905: 11 [Appias albina swinhoei]

Moore described this species from southern India: ‘♂, Ahmedabad and Bisnuggar in Guzerat (Swinhoe) in December and Poona, November to January & February; ♂ North Kanara (Davidson), ♂ Orissa (coll. de Nicéville); ♂, Poona, January(Swinhoe); ♂♀, North Kanara (S. N. Ward); ♂♀, Nilgiris (Hampson) and Travancore. Madras, July-August (Watson) (larva – Elamane, Madras, November 25 (Elliott) (pupated and emerged); Malabar; W. Ghats (S. N. Ward)’ and included the following: Appias ares Swinhoe, 1885 (part), A. paulina of Taylor (1888); neombo of Hampson (1888), lankapura of Watson (1890) and neombo of Davidson & Aitken (1896). Talbot (1939) stated ‘swinhoei (Moore), usually associated with this species [albina], was founded upon a male of libythea (Fabr.) belonging to the dry form ares Swinhoe’. The type material in BMNH clearly includes both Appias albina and Appias libythea. To stabilize usage of the name, we designate one of the specimens figured by Moore (1905) as the lectotype of Catophaga swinhoei Moore.

Lectotype ♂, INDIA, Nilgiris, Moore purchase BM 1903-361, Moore fig., 1905, 556, 1e, BMNH(E) 229139.

Paralectotypes identified as A. albina: 3 ♀♀, INDIA, Nilgiris, Moore purchase BM 1903-361, BMNH(E) 229139-40, 229144-5; 229140, Moore fig., 1905, 556, 1g,h.; 1♀, INDIA, Malabar, BMNH(E) 229141, Moore purchase BM 1903-361, Moore fig., 1905, 556, 1d;; 1♂, 1♀, INDIA, Madras, 10.viii.1889, Watson BMNH(E) 229135-6, Watson BM 1892-43; 1♂,3♀♀, INDIA, Moore purchase BM 1903-361, BMNH(E) 229137 (♂), 229142-3, 229146 (♀), 229137 Moore fig., 1905, 556, 1a, 229142 Moore fig., 1905, 556, 1i,j, 229143 Moore fig., 1905, 556, 1f.; 1♀, INDIA, Travancore, BMNH(E) 229138, Moore purchase BM 1903-361, Moore fig., 1905, 556, 1b; 1♀, INDIA, Travancore, Mynall, 2500’, i.1897, BMNH(E) 229147, Moore purchase BM 1903-361; 1♂, same data except 1200’, March, BMNH(E) 229148; 1♂, INDIA, North Kanara, 149963, Swinhoe coll. BM 1926-239.

Paralectotypes identified as Appias libythea (Fab.): 1♂, INDIA, Poona, BMNH(E) 142244, also a syntype of Appias libythea ares Swinhoe, 1883 (with label ‘co-type ares Swin., type swinhoei♂ Lep.Ind. vii. p11’). This is presumably the specimen referred to by Talbot (see above); 1♂, INDIA, Madras, 12.viii.1889, Watson BMNH(E) 229229, Watson BM 1892-43; 1♂, INDIA, Gujarat, Ahmedabad, xii.1886, BMNH(E) 229225, Moore purchase BM 1903-361, Moore 1905 fig. 556, 1a (locality cited as ‘Poona’); 1♂, INDIA, Poona, i.1888, BMNH(E) 229226, Moore purchase BM 1903-361, with hand-written label ‘neombo apud Swinhoe’; 1♂, INDIA, Gujarat, Bisnuggar [?=Visnagar], xii.1886, BMNH(E) 229227, Crowley bequest BM 1901-78, with hand-written label ‘Appias ares ♂ Swinhoe’; paralectotype? ♂, INDIA, ‘S. India’, S. N. Ward, BMNH(E) 229228, Moore purchase BM 1903-361.

Appias ambigua Grose-Smith, 1895: 76 [Appias albina ambigua]

Indonesia (♀ only): Described from ‘Two specimens from Wetter; one from Dili and one from Halmaheira’. The specimen from Halmahera cannot be located in BMNH, unless it is a female specimen of Appias paulina labelled ‘Halmaheira’, ‘ex Grose-Smith 1910’. However, this specimen is not a very good fit for the original description. To restrict the name to a particular subspecies, the following specimen is here designated the lectotype of Appias ambigua Moore.

Lectotype ♀, INDONESIA, Wetar, v.1892, W. Doherty, BMNH(E) 135769, Rothschild bequest BM 1939-1; with hand-written label ‘Ambigua Grose-Smith Type ♀’.

Paralectotypes: 1♀, INDONESIA, Wetar, v.1892, W. Doherty, 135769-70, Rothschild bequest BM 1939-1, 1♀, INDONESIA, Dili, v.1892, W. Doherty, ex Grose-Smith coll., 149931, Joicey bequest BM 1934-120.

Appias albina micromalayana Fruhstorfer, 1909: 201 (1910: 154) (= Appias albina ambigua Grose-Smith)

First described in 1909, exclusively from Babber Island, the intended original description did not appear until 1910 (as ssp. nov. and with type locality ‘East Java, Bawean and small Sunda islands'). To ensure that this name refers to the same taxon the following specimen is here designated the lectotype of Appias albina micromalayanaFruhstorfer, 1909, and Appias albina micromalayanaFruhstorfer, 1910.

Lectotype ♂, INDONESIA, Babber, H. Fruhstorfer, BMNH(E) 135826, Fruhstorfer purchase BM:1937-285.

Paralectotypes: 3♂♂, INDONESIA, Lombok, Sapit, 2000’, v.-vi.1896, H.Fruhstorfer, BMNH(E) 135807-9, Fruhstorfer purchase BM:1937-285, 135808 with Fruhstorfer label ‘albina micromalayana Fruhst. ’; 2♂♂, same data, Oberthür purchase BM 1927-3, BMNH(E)149956-7; 1♂, same locality, iv.1896, BMNH(E)135810, Fruhstorfer purchase BM:1937-285; 1♂, same locality, iv.1896, BMNH(E)149936, J. J. Joicey bequest BM 1934–120; 2♂♂, same locality, iv.1896, unregistered; 3♂♂, same data, 135811-3, Rothschild bequest BM 1939-1; 1♂, INDONESIA, Lombok, Sambalun, 4000’, iv.1896, H. Fruhstorfer BMNH(E)135814, Rothschild bequest BM 1939-1; 4♂♂, INDONESIA, Java, Sukabumi, 2000’, 1893, H.Fruhstorfer, BMNH(E) 135815-8; Fruhstorfer purchase BM:1937-285, 135815 with Fruhstorfer label ‘albina micromalayana Fruhstt'; 6♂♂, INDONESIA, Bawean, July-September., H.Fruhstorfer BMNH(E) 135819-24, Fruhstorfer purchase BM:1937-285, with Fruhstorfer label ‘albina micromalayana Fruhst.’; 1♂, same locality, wet season, BMNH(E) 135825, Fruhstorfer purchase BM: 1937-285; ♂, INDONESIA, Babber, H. Fruhstorfer, BMNH (E) 135827, Fruhstorfer purchase BM:1937-285; 2♂♂, INDONESIA, Wetar, H. Fruhstorfer, BMNH(E) 135828-9, Fruhstorfer purchase BM:1937-285; 1♂, INDONESIA, Dammer, H. Fruhstorfer, BMNH(E) 135830, Fruhstorfer purchase BM:1937-285; 1♂, INDONESIA, Kisser, H. Fruhstorfer, BMNH(E) 135831, Fruhstorfer purchase BM:1937-285; 1♂, INDONESIA, Sumba, xii.1896, H. Fruhstorfer, BMNH(E) 135832, Fruhstorfer purchase BM:1937-285; 1♂, same data except no date, 135833; 3♂♂, INDONESIA, Java, 1500’, 1891, H.Fruhstorfer, 1358 34-6, Fruhstorfer purchase BM:1937-285.

Catophaga wallacei Butler, 1898: 399 [= Appias athama athama (Blanchard, 1848)]

Type localities Vanuatu, Malekula: ‘New Hebrides (Mellicollo); New Caledonia’. To restrict the name to a particular subspecies we designate the following specimen as the lectotype of Catophaga wallacei Butler.

Lectotype ♂, VANUATU, Mallicollo I., Woodford, BMNH(E) 142267, Godman & Salvin BM 1896-12, with hand-written label ‘Catophaga wallacei ♂ type Butler’.

Paralectotypes: 1♀, VANUATU, Malekula [as Mellicollo], Woodford, BMNH(E) 135840, Godman & Salvin BM 1896-12, slide no. 4678; 1♀, same data, BMNH(E) 135841; 1♂, same data, 135842.

Further paralectotypes belong with nominate athama: 1♀, NEW CALEDONIA, J. Macgillivray, BMNH(E) 142268, purchase Cuming BM 1859-63, with hand-written label ‘Catophaga wallacei ♀ type Butler’; 1♀, NEW CALEDONIA, BMNH(E) 135839, purchase Cuming BM 1859-63, slide no.4639; 2♀♀, NEW CALEDONIA, BMNH(E) 135843 & 135846, Hewitson bequest, BM 1879-69, 135843 with det. ‘Pieris boisduvaliana Feld 4’, 135846 with labels ‘Pieris boisduvaliana Feld.3’ and ‘Athama Pol Sud’; 1♂, same data, BMNH(E) 135844, slide no. 11180, with det. ‘Pieris albina Boisd. 4’; 1♂, same data, BMNH(E) 135845, slide no. 4638, with det. ‘Pieris albina Boisd. 5’.

Appias paulina argentifera Joicey & Talbot, 1928: 19 [= Appias athama psyche (Felder & Felder)]

Type locality: Loyalty Is. Based on two males (Lifu) and one female. The males represent A. paulina ega; the female is A. athama. To stabilize the name the female is here designated the lectotype of Appias paulina argentifera Joicey & Talbot. Designation of the female provides an available name, should the Loyalty Is. population of A. athama ever be recognized as a valid subspecies.

Lectotype ♀, NEW CALEDONIA, Loyalty Is., ex Grose-Smith coll., BMNH(E) 141741, Joicey bequest BM 1934-120, with printed label ‘A. paulina argentifera J&T 1928 ♀ A.T.’. Identifiable with Appias athama (Blanchard, 1848).

Paralectotype: ♂, NEW CALEDONIA, Loyalty Is., Lifou, Lomar ?, BMNH(E) 142321, Joicey bequest BM 1934-120, with printed label ‘A. paulina argentifera J&T 1928 ♂ H.T.’. Identifiable with Appias paulina ega (Boisduval, 1836). We have not seen the second male referred to in the original description.

Papilio nero Fabricius, 1793: 153 [Appias nero nero (Fabricius, 1793)]

(Indonesia, Java): ‘Asia. Mus Britann’ (neotype here designated). A female in Banks coll. from ‘Siam’ is not considered to be a type. No possible type specimens have been located in the BMNH collections. Following Butler (1870b), this taxon has long been considered to be based on material from Java. This has recently been questioned (H. Gaonkar, personal communication, 2002), on the grounds that little material from Java reached England during the 18th century, suggesting that the original material probably came from the Malay Peninsula. However, Vane-Wright & Hughes (2005) demonstrated the opposite to be the case. To stabilize the application of nominate nero, which represents a highly polytypic species as currently conceived, the following specimen from Java is designated as the neotype of Papilio nero Fabricius.

Neotype ♂, INDONESIA, W. Java, Preanger, Palboehan Ratoe, B.M. 1922-165.

Pieris figulina Butler, 1867: 399 [= Appias nero nero (Fabricius, 1793)]

Described from a male from Singapore (coll. Roberts) and a female from Borneo (coll. Hewitson). To restrict the name to a particular subspecies, the male from Singapore is here designated the lectotype of Pieris figulina Butler.

Lectotype ♂, SINGAPORE, BMNH(E) 141751, pres. Entomological Club BM 1844-12.

Paralectotype: ♀, Borneo, 141758, Hewitson bequest, BM 1879-69. This specimen represents ssp. chelidon Fruhstorfer.

There is also a non type ♀ (but with red type label and hand-written ‘Pieris Figulina Butler type’) ‘SINGAPORE, BMNH(E) 149693, Maj. H. Roberts, BM 1928-309’. The only female mentioned in the original description is from Borneo.

Catophaga distanti Moore, 1905: 14 [Appias paulina distanti (Moore, 1905)]

Described as ‘nom. n.’ for leis of Distant (1885) and de Nicéville & Martin (1895) with the localities ‘Malay Pen.; Sumatra; Borneo’. Distant (1885) refers to ‘Malay Pen. Prov. Wellesley (coll. Distant); Perak (Biggs – coll. Distant); Malacca, Ayer-panas (Godfrey – coll. Distant); Sumatra (Brit. Mus.). Type locality is here restricted to the Malay Peninsula. A lectotype designation is desirable but, unfortunately, distanti is applied to the subspecies occurring in Peninsular Malaysia, and no syntypes from this area are available. The following three syntypes from elsewhere have been located:

Syntype ♂, ‘Borneo’, BMNH(E) 229168, Hewitson bequest BM 1879-69; syntype ♂, MALAYSIA, Sabah, Labuan, BMNH(E) 229167, BM 1868-30; syntype ♂, INDONESIA, Sumatra, S.Raffles coll., BMNH(E) 229128, Purchase Stevens BM 1854-76, with hand-written label ‘Sumatra 54.76/Catophaga alope Wallace’. This last specimen is also a possible syntype of Tachyris alopeWallace, 1867.

In addition it is necessary to consider two specimens from Java: 1♂, INDONESIA, Java, BMNH(E) 229166, BM 1860-15 & 1♀, INDONESIA, Java, BMNH(E) 229165, pres. Entomological Club BM 1844-12. Distant (1885) lists BMNH material of leis from Borneo, Java and Sumatra. Thus the above specimens would all have been in the museum collection at the time of Moore's (1905) publication, but only those from Borneo and Sumatra are considered to be syntypes of distanti, as Moore (1905) did not include Java under ‘Habitat’.

Papilio paulina Cramer, 1777: 21 [Appias paulina paulina* (Cramer, 1777)]

Described from ‘Côte de Coromandel, à Tranquebar, & dans l’Isle de Java, près de Batavia’, but here restricted to Indonesia: Java. Moore (1905) thought that paulina was from Ceylon (= Sri Lanka) and not from Java, but the lectotype is identical to material from Java and most other authors have treated paulina as such. The Sri Lankan population is now considered to represent a distinct species (A. galene). The following specimen is here designated the lectotype of Papilio paulina, to fix application of this name.

Lectotype ♀, bearing van Lennep label ‘No.60, PAULINA Cr.2. 110. E, F’ and ‘FELDER COLLN’ label; BMNH(E) 229172. This specimen is illustrated by Chainey (2005: 327, fig. 40).

Tachyris alope Wallace, 1867: 372 [= Appias paulina paulina (Cramer, 1777)]

India, Java, Sumatra, Borneo. Type locality here restricted to Java. A lectotype designation is desirable, but the type status of the material so far located is questionable.

Syntype? ♂, INDONESIA, Java, amasene, Dr. Horsf[ield], BMNH(E) 229127, slide no. Rh. 4653, with hand-written label ‘Java 33a/Catophaga alope Wallace’; syntype? ♂, INDONESIA, Java, A. R. Wallace, ex Bates coll., BMNH(E) 229173, pres. Godman Salvin BM 1896-12; syntype? ♂, INDONESIA, Sumatra, S. Raffles coll., BMNH(E) 229128, Purchase Stevens BM 1854-76, with hand-written label ‘Sumatra 54.76/Catophaga alope Wallace’. This last specimen has the best credentials to be a type specimen; however, it is identifiable with ssp. pione. It is also a possible syntype of Catophaga distantiMoore, 1905. Syntype?? (i.e. unlikely) ♂, Borneo, BMNH(E) 229126, Moore purchase, BM 1899-234, with hand-written label ‘P.alope Wall.’;

Appias eurosundana Grose-Smith, 1895: 75 [Appias paulina eurosundanaGrose-Smith, 1895]

Described from males from ‘Oinainisa (Timor) and Sambawa’ and females from ‘Oinainisa (Timor) and one from Wetter’. To restrict the name to a particular subspecies the following specimen is here designated the lectotype of Appias eurosundana Grose-Smith.

Lectotype ♂, INDONESIA, Timor, Oinainisa, xi.-xii.1891, W. Doherty, BMNH(E) 229051, Rothschild bequest, BM 1939-1; with Grose-Smith ♂ type label.

Paralectotypes: 6♂♂, 10♀♀, INDONESIA, Timor, Oinainisa, xi.-xii.1891, W. Doherty, BMNH(E) 229045-61 (except 229051 above), Rothschild bequest, BM 1939-1; 229052 with Grose-Smith ♀ type labels; 1♀, same data, BMNH(E)149995, ex Grose-Smith coll., Joicey bequest BM 1934-120; 3♂♂, same data, BMNH(E) 149978-80, Oberthur purchase BM 1927-3. A further paralectotype ♂, INDONESIA, Sumbawa, ix.1891, W. Doherty, BMNH(E) 229044, Rothschild bequest, BM 1939-1; represents ssp. tambora.

Appias saina Grose-Smith, 1894: 336 (Appias paulina sainaGrose-Smith, 1894)

Described from two males from ‘Humboldt Bay, Dutch New Guinea; ’ and one female from ‘Biak’Joicey & Noakes (1915) noted that ‘Smith’s male is a female’. To restrict the name to a particular subspecies the following specimen is here designated the Lectotype of Appias saina Grose-Smith.

Lectotype ♀, INDONESIA, Humboldt Bay, ix.-x.1893, W. Doherty, BMNH(E) 229041, Rothschild bequest, BM 1939-1; with Grose-Smith type ♂ label.

Paralectotype: ♀, BMNH(E) 229043, with same data.

A further paralectotype male from Biak, W. Doherty, BMNH(E) 229042, Rothschild bequest, BM 1939-1, with Grose-Smith type male label belongs with ssp. falcidia.

Ancillary