SPHENOPHRYNE - American Museum of Natural History

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4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 253 ABSTRACT Parker (1934) distinguished Sphenophryne from other genera of the Genyophryninae (Sphenophryninae in his work) by its pectoral girdle having a more nearly complete complement of bony and cartilaginous elements. Otherwise, the species he placed there were a diverse lot, including ones adapted for climbing, for living in or on leaf litter, or for a cryptic, even burrowing existence. The characteristics of the pectoral girdle are primitive compared to those of other genyophrynine genera, which have reduced or lost one or more of the elements. Thus, the common possession of a primitive girdle is inadequate as the sole character defining the genus. In the present work I review species with the supposedly diagnostic character of Sphenophryne and recognize four genera: Sphenophryne (monotypic), and three removed from synonymy—Austrochaperina (23 species), Liophryne (6 species), and Oxydactyla (5 species). The characters I use to define the genera are for the most part closely tied to the habits of the species and thus are subject to the criticism that homoplasy rather than synapomorphy is demonstrated. Regardless, the probability that the new arrangement identifies four monophyletic lines is considerably greater than that Sphenophryne, as it has been constituted, is monophyletic. Sphenophryne, Liophryne, and Oxydactyla are confined to New Guinea. Austrochaperina has one species on New Britain, four endemic to northern Australia, one shared between Australia and New Guinea, and 18 on New Guinea and adjacent islands. The genus and species Microbatrachus pusillus, based on a hatchling from the Aru Islands, are considered unidentifiable. Of the 35 species recognized, 17 are described as new and one is removed from synonymy. In his classic monograph of the Microhylidae, Parker (1934) recognized the genus Sphenophryne with 10 species found in Australia and New Guinea and considered three genera as synonymous to it. 1 In more than six decades since, the only taxonomic modification at the generic level was the synonymization of Microbatrachus (which Parker recognized) with Sphenophryne (Tyler, 1978). The number of included species has risen from 10 to 18, mostly through discovery of new species (listed in Zweifel, 1985a, with an additional species in Zweifel, 1985b). Parker’s diagnostic feature of the genus within the subfamily Sphenophryninae (Genyophryninae of current authors) was that the pectoral girdle had a more nearly complete set of ventral elements than did the 1 Parker (1934: 159) also reported a ‘‘?Sphenophryne sp.’’ based on embryos ‘‘collected from the pitcher of a Nepenthes at Kuching [Sarawak],’’ far from the known range of any genyophrynine microhylid. It is likely that this extralimital record is based on a rhacophorid, possibly Philautus, which oviposits in Nepenthes and whose larvae may metamorphose prior to hatching; see Dring (1979: 207–208) and Inger and Stuebing (1997: 163). INTRODUCTION other genera he recognized 2 : coracoids, clavicles that extend from the scapula to near the midline, procoracoid cartilages, and a cartilaginous sternum. The set of features described for the pectoral girdle is unquestionably primitive compared to the more reduced states in all other genyophrynine genera and thus cannot diagnose a monophyletic clade. Species currently assigned to Sphenophryne on the sole basis of girdle structure are a diverse lot morphologically and ecologically, and they include scansorial, riparian, and terrestrial forms. The inevitable impression is that they may not be a monophyletic group. My principal objective has been to examine the suite of genyophrynine species with the primitive pectoral girdle, including several undescribed and synonymized taxa, in order to see if phylogenetic sense can be made and a defensible arrangement can be proposed that does not depend on shared primitive characters. Additional objectives were to identify and describe previously unrecognized species, to assess the status of names in synonymy, and 2 Oreophryne, Aphantophryne. Microbatrachus, Cophixalus, and
2000 ZWEIFEL: PARTITION OF SPHENOPHRYNE 5 to describe as far as possible the distribution and ecology of the species. In a previous paper (Zweifel, 1985b) I reviewed the five Australian species of Sphenophryne in detail. Except for one of these that occurs also in New Guinea, they are not treated individually in this account but are considered where pertinent. ACKNOWLEDGMENTS Over the many years since initiating this project I have become increasingly in debt to many institutions and to colleagues without whose assistance I could have accomplished little. A list of institutions whose specimens I examined (or if not examined, as cited in the text) follows, with abbreviations and names of responsible parties. AMNH American Museum of Natural History, New York; C. Myers, L. Ford AMS Australian Museum, Sydney; H. Cogger, A. Greer, R. Sadlier BMNH British Museum (Natural History), London; B. Clarke BPBM Bishop Museum, Honolulu; Allen Allison, Carla Kishinami FMNH Field Museum of Natural History, Chicago; R. Inger MCZ Museum of Comparative Zoology, Harvard University, Cambridge; José Rosado, E. Williams, J. Cadle MNH National Museum of Hungary, Budapest MSNG Museo Civico di Storia Naturale di Genova, Genoa; E. Tortonese, L. Capocaccia-Orsini MTKD Staatliches Museum für Tierkunde, Dresden MZB Zoological Museum, Bogor, Indonesia; S. Somadikarta, Siti Nuramaliati Prijonok, Curator Mumpuni NMBA Basel Natural History Museum, Basel PNGM National Museum and Art Gallery, Port Moresby, Papua New Guinea; J. Menzies, I. H. Bigilale QM Queensland Museum, Brisbane, Australia RMNH Rijksmuseum van Natuurlijke Historie, Leiden; R. Hoogmoed, L. Brongersma SAMA South Australian Museum, Adelaide; Adrienne Edwards UPNG University of Papua New Guinea, Port Moresby; J. Menzies YPM Yale Peabody Museum, New Haven; Fred Sibley ZMA Zoological Museum, University of Amsterdam, Amsterdam; S. Daan ZMUC Zoological Museum, University of Copenhagen, Copenhagen; Arne Schitz ZSM Zoologisches Sammlung des Bayerischen Staates, Munich; Frank Glaw Fieldwork that permitted me to gain acquaintance with living individuals of certain species studied was supported by the American Museum of Natural History, the National Geographic Society (1968), the National Science Foundation (1964 and Alpha Helix Expedition, 1969), and the Sabin Conservation Fund (1987). Individual thanks are in order to Linda Ford, for her efficient and prompt replies to my many requests for information and specimens from the AMNH; David Dickey, who produced some of the audiospectrograms; Allen Allison, James Menzies, and Stephen Richards, all of whom provided specimens, photographs, and tape recordings; Carol Townsend for the preparation of cleared and stained specimens; Joan Whelan for SEM photography; Wade Sherbrooke, Director of the AMNH Southwestern Research Station, for providing laboratory space where I carried out much of my work with specimens; and the British Library of Wildlife Sounds and Ian Redmond for the loan of tape recordings. Charles Myers’ careful critique of the manuscript improved it significantly in a variety of ways, as did comments from two anonymous referees. Fred Parker of Townsville, Queensland, Australia, deserves special mention. Not only did I have access to the multitudes of specimens he collected and generously deposited in several museums, but I was privileged to accompany him on patrol in the Eastern Highlands of Papua New Guinea in 1964, and again to share time in the field with him in 1968, 1969, and 1987. As always, my wife, Frances W. Zweifel, applied her artistic talents to produce many original renderings for this paper in addition
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