Systematic Entomology (2010), DOI: 10.1111/j.1365-3113.2010.00535.x The Golden and Mariana albatrosses, new species of pierid butterflies, with a review of subgenus Appias (Catophaga) (Lepidoptera) OSAMU YATA1, JOHN E. CHAINEY2 andRICHARD I. VANE-WRIGHT2,3 1Biosystematics Laboratory, Kyushu University, Fukuoka, Japan and 2Department of Entomology, The Natural History Museum, London, U.K. and 3Durrell Institute of Conservation and Ecology (DICE), School of Anthropology and Conservation, University of Kent, Canterbury, U.K. Abstract. This paper presents an overview of the subgenus Appias (Catophaga) Hubner¨ (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 list of all currently accepted subspecies to facilitate the future revisionary taxonomy that will probably be needed once This work does not constitute a fully comprehensive revi- extensive molecular investigations have been made into the sion. Its purpose is three-fold: to provide an overview of the numerous populations of these insects. species of subgenus Appias (Catophaga)Hubner,¨ including The butterflies assigned currently to the genus Appias the description of two new members of the group, and a key Hubner¨ comprise more than 40 species of ‘whites’ (Pieridae). to adult males and females; to review known larval hostplant Although found largely in the old world tropics, the genus is relationships, distribution and, especially, the spectacular poly- also represented in the Americas. Appias is placed, together morphism of certain species, with a view to stimulating further with Saletara Distant (three or four species: Parsons, 1998; research into their ecology, genetics, molecular systematics Vane-Wright & de Jong, 2003), Udaiana Distant (one species) and evolutionary biology; and to provide a fully synonymic and Aoa de Niceville´ (one species), in the Appiadina Kusne- zov, a subtribe of the Pierinae: Pierini (Braby et al., 2006). Correspondence: Osamu Yata, Kyushu University Museum, Although the species of Appias were divided by Klots Hakozaki 6-10-1, Higashi-ku, Fukuoka 812-8581, Japan. E-mail: (1933: 208) into four subgenera, Yata (1981) proposed seven [email protected] subdivisions, five of which were recognized as subgenera (Appias s.s., Catophaga Hubner,¨ Phrissura Butler, Hiposcritia Unpublished for the purposes of zoological nomenclature (Art. 8.2, Geyer, Glutophrissa Butler, plus the lyncida and sylvia species ICZN) groups). Braby et al. (2006: 263) suggested that the collective © 2010 The Authors Systematic Entomology © 2010 The Royal Entomological Society 1 2 O. Yata et al. genus Appias, as accepted currently, is almost certainly a Wing venation terminology paraphyletic group (notably with respect to the exclusion of Saletara: Yata, 1981: 392). The Comstock–Needham wing-vein and cell nomenclature Among these divisions, the Indo-Australian subgenus adopted in the descriptions is based on Nielsen & Common Catophaga Hubner¨ is remarkable for the inclusion of several (1991) and Smith & Vane-Wright (2001). This terminology, brightly coloured species in which the adult males, instead together with the numerical system employed by Yata (1981) of the typical white or yellow of most Appias, are brilliant and many other lepidopterists (e.g. Corbet & Pendlebury, flame orange, chocolate brown or powder blue. The females 1992), is illustrated in Fig. 1. of these butterflies are variously coloured, either white, yel- low, polymorphic white or yellow, polymorphic white, yellow Taxonomic results or male-like, or, in some cases, simply male-like with respect to their dominant colour. Catophaga is also notable because Appias Hubner¨ some, if not all, of its species seem capable of switching larval hosts between Capparaceae (a family of the Brassi- Appias Hubner,¨ 1819: 91. Type species by selection of cales, now often placed within the Brassicaceae) and certain Butler (1870a: 49): Papilio zelmira Stoll, 1780. [P. zelmira 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 Entomolog- ical Laboratory, Faculty of Agriculture, Kyushu Univer- sity (AGKU), Museum´ 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 Fig. 1. Wing venation of Appias (Catophaga) paulina, showing both the Comstock–Needham terminology and the numerical system (small any compression by glass slide and cover slip. For better ciphers) for the long veins. The short cross-veins closing the discal contrast, some preparations were stained with Chlorazol Black. are notated according to common lepidopterological practice: upper, Terminology for male genitalia is based on Shirozu’sˆ (1960: middle and lower discocelluar veins (udc, mdc, ldc). The cells are 1–10) extensive account, except that we use the term phallus notated using the Comstock–Needham system only. Dotted lines in instead of the more frequent ‘aedeagus’, as endorsed by the discal cells indicate ‘folds’ (probable courses of proximal parts of Kristensen (2003). Terminology for female genitalia mainly veins M1 –M3), and in CuA2 the lost vein CuB (which supposedly follows van Son (1949), with some additions from Kusnezov appears during early development but is later resorbed). Based in part (1915) and Yamauchi & Yata (2004). on Smith & Vane-Wright (2001: 513, fig. 7). © 2010 The Authors Systematic Entomology © 2010 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/j.1365-3113.2010.00535.x Pierid butterflies of the subgenus Appias (Catophaga) 3 is considered to represent the same species group taxon as the these all comprise monophyletic units. The paulina complex, older nominal species, Papilio libythea Fabricius, 1775; Appias which comprises eight closely related species in which the (Appias) libythea occurs widely in the Oriental Region. Butler males are white, yellow or bluish, is represented throughout (1870a: 49) gave priority, as first reviser, to Appias Hubner¨ the entire Indo-Australian region (Fig. 23). The nero group, over Catophaga Hubner.]¨ 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 Subgenus Appias (Catophaga)Hubner¨ Buru (Central Maluku), and is entirely parapatric with respect to the third group, the melania series (four allopatric species in Catophaga Hubner,¨ 1819: 93. Type species by selection of which the males are brown or bluish), confined to the Papuan Scudder (1875: 136): Papilio paulina Cramer, 1777. subregion, including Australia (Fig. 24). Trigonia Geyer, 1837: 21, 35. Type species by selection of paulina complex: galene, wardii, albina, aurosa sp.n., Scudder (1875: 286): Papilio nero Fabricius, 1793. (Invalid athama, paulina, mariana sp.n., mata name: junior homonym of Trigonia Brugiere,` 1789.) nero group: galba, nero, zarinda Tachyris Wallace, 1867: 361. Type species by selection of melania series: placidia, clementina, celestina, melania Scudder (1875: 274): Papilio nero Fabricius, 1793. The paulina complex Diagnosis Diagnosis. Male: upperside ground colour usually