The Golden and Mariana albatrosses, new species of pierid butterflies, with a review of subgenus Appias ( Catophaga ) (Lepidoptera)
2010; Wiley; Volume: 35; Issue: 4 Linguagem: Inglês
10.1111/j.1365-3113.2010.00535.x
ISSN1365-3113
AutoresOsamu Yata, John Chainey, Richard I. Vane‐Wright,
Tópico(s)Biological Control of Invasive Species
ResumoSystematic EntomologyVolume 35, Issue 4 p. 764-800 Free Access The Golden and Mariana albatrosses, new species of pierid butterflies, with a review of subgenus Appias (Catophaga) (Lepidoptera) OSAMU YATA, Corresponding Author OSAMU YATA Biosystematics Laboratory, Kyushu University, Fukuoka, JapanOsamu Yata, Kyushu University Museum, Hakozaki 6-10-1, Higashi-ku, Fukuoka 812-8581, Japan. E-mail: [email protected]Search for more papers by this authorJOHN E. CHAINEY, JOHN E. CHAINEY Department of Entomology, The Natural History Museum, London, U.K.Search for more papers by this authorRICHARD I. VANE-WRIGHT, RICHARD I. VANE-WRIGHT Department of Entomology, The Natural History Museum, London, U.K. Durrell Institute of Conservation and Ecology (DICE), School of Anthropology and Conservation, University of Kent, Canterbury, U.K.Search for more papers by this author OSAMU YATA, Corresponding Author OSAMU YATA Biosystematics Laboratory, Kyushu University, Fukuoka, JapanOsamu Yata, Kyushu University Museum, Hakozaki 6-10-1, Higashi-ku, Fukuoka 812-8581, Japan. E-mail: [email protected]Search for more papers by this authorJOHN E. CHAINEY, JOHN E. CHAINEY Department of Entomology, The Natural History Museum, London, U.K.Search for more papers by this authorRICHARD I. VANE-WRIGHT, RICHARD I. VANE-WRIGHT Department of Entomology, The Natural History Museum, London, U.K. Durrell Institute of Conservation and Ecology (DICE), School of Anthropology and Conservation, University of Kent, Canterbury, U.K.Search for more papers by this author First published: 23 August 2010 https://doi.org/10.1111/j.1365-3113.2010.00535.xCitations: 6AboutSectionsPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinkedInRedditWechat 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 1Open in figure viewerPowerPoint 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 23Open in figure viewerPowerPoint Range map for all currently recognized species of the Appias (Catophaga) paulina complex. Figure 24Open in figure viewerPowerPoint 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 (2, 19) Figure 2Open in figure viewerPowerPoint 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 19Open in figure viewerPowerPoint 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 (3, 19) Figure 3Open in figure viewerPowerPoint 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 (4, 19) Figure 4Open in figure viewerPowerPoint 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 (5, 19, 20) Figure 5Open in figure viewerPowerPoint 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 20Open in figure viewerPowerPoint 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. (19, 20). 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. (19, 20). 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 are
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