First Record of the Poorly Known Skink Sphenomorphus oligolepis（Boulenger， 1914） （Reptilia： Squamata： Scincidae） from Seram Island， Maluku Province， Indonesia

Sven MECKE， Max KIECKBUSCH， Mark O'SHEAand Hinrich KAISER

1Department of Animal Evolution and Systematics and Zoological Collection Marburg， Faculty of Biology， Philipps-Universit?t Marburg， Karl-von-Frisch-Stra?e 8， 35032 Marburg， Germany

2Faculty of Science and Engineering， University of Wolverhampton， Wulfruna Street， Wolverhampton， WV1 1LY， United Kingdom； and West Midland Safari Park， Bewdley， Worcestershire DY12 1LF， United Kingdom

3Department of Biology， Victor Valley College， 18422 Bear Valley Road， Victorville， California 92395， USA； and Department of Vertebrate Zoology， National Museum of Natural History， Smithsonian Institution， Washington， DC 20013， USA

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First Record of the Poorly Known Skink Sphenomorphus oligolepis（Boulenger， 1914） （Reptilia： Squamata： Scincidae） from Seram Island， Maluku Province， Indonesia

Sven MECKE1*， Max KIECKBUSCH1， Mark O'SHEA2and Hinrich KAISER3

1Department of Animal Evolution and Systematics and Zoological Collection Marburg， Faculty of Biology， Philipps-Universit?t Marburg， Karl-von-Frisch-Stra?e 8， 35032 Marburg， Germany

2Faculty of Science and Engineering， University of Wolverhampton， Wulfruna Street， Wolverhampton， WV1 1LY， United Kingdom； and West Midland Safari Park， Bewdley， Worcestershire DY12 1LF， United Kingdom

3Department of Biology， Victor Valley College， 18422 Bear Valley Road， Victorville， California 92395， USA； and Department of Vertebrate Zoology， National Museum of Natural History， Smithsonian Institution， Washington， DC 20013， USA

Based on four specimens discovered in the collection of The Natural History Museum， London， United Kingdom， we present a new distribution record for the skink Sphenomorphus oligolepis for Seram Island， Maluku Province， Indonesia. This fi nd constitutes the westernmost record for the species and extends its range by over 800 km. The species was heretofore only known from apparently isolated mainland New Guinean populations.

Scincidae， Lygosominae， Sphenomorphus oligolepis， new record， Seram， Maluku Islands， Indonesia，Wallacea

1. Introduction

Sphenomorphus oligolepis （suggested common name：MIMIKA FOREST SKINK） is a member of the S. maindroni group （sensu Greer and Shea， 2004）. It is a poorly known skink with apparently disjunct populations on mainland New Guinea and has experienced a very limited treatment in the scientific literature （e.g.， de Rooij， 1915； Greer， 1973； Greer and Shea， 2004）. The species has been reported from the Mimika River （the type locality； Boulenger， 1914） and the Lorentz River，Papua Province， Indonesia （de Rooij， 1915）， and more recently from several localities in Papua New Guinea （see Greer， 1973： Figure 8）， including Bikim， Matkomrae， and Mendua （Western Province）； Soliabeda （Simbu Province），and Oroi （Gulf Province）. Additional specimens housed in the Museum of Comparative Zoology， Cambridge，Massachusetts， USA （MCZ） and the Bernice P. Bishop Museum， Honolulu， Hawaii， USA （BPBM） were collected in Gulf Province at Kikori （MCZ R-150879）and Weiana （MCZ R-101484）， and in Morobe Province at Aseki （BPBM Herp-17441-48； 19103-09）， with a single voucher collected at Timika， Papua Province，Western New Guinea， Indonesia （BPBM Herp-42441）. The westernmost record of S. oligolepis known to date is the type locality， and the species has never been recorded from localities other than on mainland New Guinea. Here we report a fi rst record of S. oligolepis from Seram Island，Maluku Province， Indonesia （for a distribution map see Figure 1）.

2. Material and Methods

Figure 1 Distribution of Sphenomorphus oligolepis in New Guinea and in the Moluccas （black symbols）. The type locality of the species（Mimika River， West Papua， Indonesia； BMNH 1946.8.3.47-48） is indicated by a star. The triangle denotes the new distribution record for Seram Island， Maluku Province， Indonesia （BMNH 1998.299-302）. Numbers accompanying black circles identify the following known localities for S. oligolepis： （1） Lorentz River （de Rooij， 1915）， （2） Timika， Nayaro Settlement （BPBM Herp-42441）， （3） Matkomrae （MCZ R-130716）， （4） Menuda （MCZ R-130717）， （5） Bikim （MCZ R-130718）， （6） 30 km N， 14 km W Kikori （MCZ R-150879）， （7） Weiana （MCZ R-101484）， （8） Soliabedo （MCZ R-118857）， （9） Oroi （MCZ R-109330-45， 118854-56； WAM R-67631-32）， （10） Aseki （BPBM Herp-17441-48； 19103-09）. Map prepared by Sven Mecke.

During a taxonomic investigation of skinks in the collection of The Natural History Museum， London，United Kingdom （BMNH）， two of the authors （HK and SM） discovered four specimens of a scincid lizard from Seram Island， Maluku Province， Indonesia， labeled“Sphenomorphus sp. A” （BMNH 1998.299-303； Figure 2）. The four specimens had been caught by Paul Edgar and Ronald Lilley in pitfall traps in a lowland rainforest（elevation ca.50 m） near Solea， northwestern Seram，in late August and early September 1987， during a herpetofaunal survey of the island. Climate data for the collection locality and survey methods were summarized in detail by Edgar and Lilley （1993）.

Comparative measurements and scale counts （Table 1） of “Sphenomorphus sp. A” and other museum specimens examined were performed according to the following protocol. Measurements were taken on the right side of the body to the nearest 0.1 mm using digital calipers. Eidonomic characters （abbreviations provided in parentheses） used include snout-vent length （SVL），measured from tip of snout to cloaca； tail length （TailL），measured from cloaca to tip of tail； arm length （ArmL），measured from axilla to tip of longest fi nger； leg length（LegL）， measured from point of posterior body insertion to tip of longest toe； head length （HeadL）， measured from tip of snout to anterior edge of ear opening， head width（HeadW）， measured at widest point of head； number of scales rows at midbody （MBSR）， number of nuchal scales （NS）， number of paravertebral scales （PVS），

counted in one row beginning with the fi rst nuchal scale to the first scale fully anterior to the rear edge of the hindlimbs； number of supralabials （SupraLab）； number of supracilaries （SupraCil）， and the number of enlarged lamellae under the 4thtoe （4TLam）， counted as all scales wider than the plantar scales distal to the cleft between the 3rdand 4thdigits. We also calculated the following ratios： ArmL/SVL， LegL/SVL， HeadL/SVL. Greer （1973）found female specimens of S. oligolepis to be gravid at a minimal SVL of 43.0 mm. Hence， we assume that the four unsexed specimens from Seram （minimal SVL 48.0 mm） are adults.

3. Results and Discussion

The four specimens could be easily identifi ed as members of the Sphenomorphus maindroni group （sensu Greer and Shea， 2004； 22 species recognized） by the presence of a post-supraocular scale. While the highest species diversity of the S. maindroni group is found in New Guinea，members of this assemblage also occur in the Bismarck Archipelago and the Solomon Islands， the southern part of the Philippines， Palau， and some of the Moluccan Islands（Greer and Shea， 2004）.

Table 1 Morphometric （in mm） and meristic data of the four specimens of Sphenomorphus oligolepis from Seram Island， Maluku Province，Indonesia （BMNH 1998.299-302）， and of the two syntypes of this species （BMNH 1946.8.3.47-48）. Only characters that allow comparison with data in the relevant literature are shown. When meristic characters occurring bilaterally where different on both sides of the body， this is indicated by the letters ‘R' （right） and ‘L' （left）. Otherwise the respective character is represented by a single value （right side of the body）. When tails were found to be partly regenerated， this is indicated by a superscript ‘R' after TailL. Numbers in square brackets show sample sizes （individuals） or cases， if the superscript ‘C' is used. Numbers in parentheses refer to mean values or， when underlined， modal values.

Identification of the four Sphenomorphus specimens from Seram as S. oligolepis was confi rmed eidonomically，based on the descriptions in Boulenger （1914）， de Rooij（1919）， Greer （1973）， and the diagnostic characters presented by Greer and Shea （2004）， who also provided a key to the members of the S. maindroni group. We also examined the syntypes of S. oligolepis （BMNH 1946.8.3.47-48； Figure 3） for direct comparison.

In overall eidonomy （size， body proportions， scalation，general aspects of coloration）， the specimens from Seram conform to the descriptions of Sphenomorphus oligolepis presented in the relevant literature （Boulenger， 1914；de Rooij， 1915； Greer， 1973； Greer and Shea 2004）. Morphometric and meristic data for the specimens（Table 1） show that they fall well within the range of S. oligolepis， although the data available in the literature are quite limited. The Seram specimens are also diagnosable as S. oligolepis using the identifi cation key of Greer and Shea （2004）. In addition， the last supralabial scale in the Seram specimens is divided， as is typical for S. oligolepis（Glenn Shea， in litt.）. Moreover， eidonomic data of the Sphenomorphus specimens from Seram also conform to those of the syntypes of S. oligolepis （Table 1）1Data on the number of PVS in S. oligolepis， although available for most other S. maindroni group members， are not provided in the relevant literature. Although our examination of the Seram specimens yielded PVS counts different from those of the type specimens of S. oligolepis（63-69 in the Seram specimens， and 57 and 58 in the type series of S. oligolepis）， Glenn Shea examined 38 specimens of S. oligolepis and obtained a PVS range of 55-73 （Glenn Shea， unpubl. data）， indicating that this character is much more variable than in the type series.. We therefore conclude that the Seram specimens provisionally labeled “Sphenomorphus sp. A” are members of this species， which is hereby recorded for the fi rst time as part of the Seram herpetofauna. This record for S. oligolepis is the westernmost record for the species， and the fi rst non-New Guinean； it is the fi rst from the biogeographic region known as Wallacea.

Sphenomorphus oligolepis is readily distinguishable from S. undulatus， the only other species of the S. maindroni group known from Seram （de Rooij， 1915；Dunn， 1927； Edgar and Lilley， 1993） by separated prefrontal scales （vs. prefrontals in medial contact in S. undulatus）， a single infralabial in contact with the postmental （vs. two infralabials in contact with thepostmental in S. undulatus）， and a much lower number of 4TLam （9-12 vs.17-23 in S. undulatus） （see Greer and Shea， 2004： Table 2 and Key to Species； pers. obs.）.

Figure 2 Sphenomorphus oligolepis （BMNH 1998.299-302） from Seram Island， Maluku Province， Indonesia. （A） Specimens in dorsal view.（B） Specimens in ventral view. Scale = 10 mm. Photos by Thomas Beitz.

Figure 3 Syntypes of Sphenomorphus oligolepis （BMNH 1946.8.3.47-48） in （A） dorsal and （B） ventral view. Photos by Mark O'Shea.

The species （listed as “Sphenomorphus sp. A”） was reported to be diurnal and fossorial by Edgar and Lilley（1993）. According to these authors， on Seram it was found in lowland rainforest （50 m） and in forest above 700 m，but no voucher specimens were obtained from the higher locality. In a lowland rainforest near Solea， northwestern Seram， S. oligolepis occurs in syntopy with three species of geckos， a dibamid， five skinks， two blindsnakes， one colubrid snake， and one elapid snake species （Edgar and Lilley， 1993： Table 4）.

Sphenomorphus oligolepis appears to be a widely，though not necessarily continuously， distributed species in southern New Guinea （distribution extends ～1200 km from west to east； Figure 1）， where it has been found in lowland rainforests and freshwater swamp forests（elevations 0-550 m）， but also at higher elevations in the lower montane southeastern Papuan rainforests（elevations up to 1250 m）. It might be expected that the species also occurs in the lowland rainforests of the‘neck' of the Vogelkop Peninsula （West Papua Province，Indonesia）， and further range extensions in the western part of New Guinea can be expected. These would fi ll the largest known distribution gap for S. oligolepis （linear distance of ＞ 800 km； Figure 1）.

It should be noted that many mainland New Guinean lizard taxa have rather discontinuous distribution patterns，often with larger gaps between isolated populations （see distribution maps provided by Allison and Kraus， 2011）. Obvious distribution gaps might be the result of a true spatial separation of single species （intraspecifi c allopatry）or represent potential interspecific barriers betweensimilar looking， though different taxa （interspecific allopatry in an undiscovered biodiversity）. However，distribution gaps might rather reflect an undersampling bias.

Some of these taxa with spatially separated populations are also found on islands west of New Guinea， including Seram. Examples may be the gecko Cyrtodactylus papuensis， and the skinks Eugongylus rufescens，Sphenomorphus muelleri， Sphenomorphus undulatus， and Tiliqua gigas （Brongersma， 1953； de Rooij， 1915； Dunn，1927； Shea， 2000）.

The absence of records of Sphenomorphus oligolepis between the type locality and Solea， Seram （including the ‘neck' of the Vogelkop Peninsula and eastern Seram）may be explained by an undersampling bias resulting from （1） under-collection in areas potentially difficult to access； and （2） the semifossorial habit of this taxon，which makes it difficult to find individuals （especially by expeditions not primarily focusing on herpetofauna species and if no pitfall traps were used）. Voucher specimens were thus almost exclusively collected by experienced herpetologists （Fred Parker， Allen Allison）and predominantly during more recent expeditions to Papua New Guinea.

The presence of Sphenomorphus oligolepis in Seram increases to three the number of Sphenomorphus skinks known from this island and， together with recent species descriptions from the region （e.g.， Harvey et al.， 2000；Oliver et al.， 2009； Vogel and van Rooijen， 2008； Weijola and Sweet， 2010； Ziegler et al.， 2007）， demonstrates how little is known about the herpetofauna of the Moluccas（Maluku and North Maluku Provinces）.

Acknowledgements We are indebted to Patrick Campbell （BMNH） for his hospitality and the loan of specimens. We thank José Rosado and Joseph Martinez（MCZ）， and Pumehana Imada and Allen Allison （BPBM）for access to specimens photographed by one of us（MOS）. We also thank Ronald Lilley （The Indonesian Nature Foundation） for providing some of the cited literature， Glenn Shea （The University of Sydney） for providing unpublished data of Sphenomorphus oligolepis，and Thomas Beitz for the photographs of the S. oligolepis specimens from Seram （Figure 2）. We are very grateful to George R. Zug （USNM） and an anonymous reviewer for their helpful comments on earlier versions of the manuscript. Travel funds to visit the MCZ were provided by an Ernst Mayr Grant from Harvard University to MOS.

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*

Sven MECKE， from Philipps-Universit?t Marburg， Germany， with his research focusing on the taxonomy，systematics， and biodiversity of Indo-Australian amphibians and reptiles.

E-mail： meckes@staff.uni-marburg.de

17 July 2015 Accepted： 17 February 2016