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Taxonomic Resolution of the Aprasia Repens Species-Group (Squamata

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Taxonomic Resolution of the Aprasia Repens Species-Group (Squamata RECORDS OF THE WESTERN AUSTRALIAN MUSEUM 30 012–032 (2015) DOI: 10.18195/issn.0312-3162.30(1).2015.012-032 Taxonomic resolution of the Aprasia repens species-group (Squamata: Pygopodidae) from the Geraldton Sandplains: a description of a new species and additional mainland records of A. clairae Brad Maryan1,2, Mark Adams3,4 and Ken P. Aplin2 1 Biologic Environmental Survey, 50B Angove Street, North Perth, Western Australia 6006, Australia. Email: [email protected] 2 Research Associate, Department of Terrestrial Zoology, Western Australian Museum, 49 Kew Street, Welshpool, Western Australia 6106, Australia. 3 Evolutionary Biology Unit, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia. 4 Australian Centre for Evolutionary Biology and Biodiversity, University of Adelaide, Adelaide, South Australia 5000, Australia. ABSTRACT – The Australian pygopodid genus Aprasia is a group of morphologically conservative, worm-like fossorial lizards. Combined allozyme and morphological analyses revealed a previously unrecognised species, named herein as A. wicherina sp. nov., known from a small number of specimens from an area of elevated, ancient sandplains on the central west coast of Western Australia. The new species is a member of the Aprasia repens species-group, but it is genetically divergent and morphologically distinguishable from all other previously described members of this group, including the geographically proximate A. clairae, A. haroldi, A. repens and A. smithi. We also report additional specimens of A. clairae from the central west coastal mainland and we provide new information that supplements our original description of this species. Finally, we present preliminary allozyme evidence for additional candidate species in A. repens, thus highlighting the need for greater geographic sampling of this widespread taxon. Addition of A. wicherina sp. nov. to the Western Australian endemic A. repens species-group brings the known diversity to eight species, with the highest diversity on the Geraldton Sandplains. The discovery of yet another new and potentially rare vertebrate species from southwestern Australia underlines our lack of knowledge of the most developed part of the state. KEYWORDS: worm lizard, taxonomy, Aprasia wicherina sp. nov., conservation, allozyme electrophoresis INTRODUCTION al. (1990). The fi rst outcome of this work was the Aprasia Gray, 1839 is a genus of small, worm-like description of A. clairae from single localities on the Houtman Abrolhos islands and the adjacent mainland fossorial pygopodids, which currently includes 10 near Dongara (Maryan et al. 2013a). A second outcome species in Western Australia. Preliminary investigations was the description of A. litorea from the Lake MacLeod of the endemic A. repens species-group (which includes region, and the synonymising of A. fusca with A. all Western Australian species other than A. inaurita, rostrata (Maryan et al. 2013b). During these studies A. pulchella and A. striolata) suggested that species several other unresolved taxonomic issues were noted diversity had been significantly underestimated, within the A. repens species-group, including the especially along the west coast between Geraldton identity of morphologically distinctive specimens from and the North West Cape (Aplin and Smith 2001: 70). localities at the northern edge of the range of A. repens These previous investigations initiated a combined (Maryan et al. 2013a). Previous treatments of A. repens molecular and morphological appraisal of taxa within have included specimens from as far north as Kalbarri the A. repens species-group as proposed by Storr et (Storr et al. 1990; Wilson and Swan 2013), however the http://www.zoobank.org/urn:lsid:zoobank.org:pub:B402B16D-1966-402B-A9DC-52C38F25B705 TAXONOMIC RESOLUTION OF THE APRASIA REPENS SPECIES GROUP 13 northernmost record is based on a single specimen of procedural details for implementing these analyses are uncertain identity (Aplin and Smith 2001: 70). presented in these studies and discussed in more detail In this contribution, our third on the A. repens elsewhere (Hammer et al. 2007; Adams et al. 2014). species-group; we focus on the taxonomic identity of MORPHOLOGICAL ANALYSIS four distinctive specimens from an area of ancient, elevated sandplains on the central west coast of Western The four individuals of the putative new species Australia. In general aspects of morphology, including were compared to 17 specimens of the morphologically the elongate body and protrusive rostral ‘beak’, these similar species, A. rostrata, from the North West Cape specimens resemble A. rostrata (sensu Maryan et peninsula and offshore islands, and to the nearest al. 2013b); the most northerly distributed member of geographical congeners comprising 7 specimens of the A. repens species-group. However, the southern A. clairae and 29 A. repens from between Perth and sandplains population differs from A. rostrata and Geraldton (Appendix and type lists in Taxonomy all other members of the A. repens species-group on section). All specimens are from the collections of both molecular and morphological criteria. We herein the Western Australian Museum, Perth (WAM). Sex diagnose this population as the eighth species in the of individuals was determined by visual inspection A. repens species-group. Our expanded analyses also of everted hemipenes and postcloacal spurs in males, resulted in the recognition of additional mainland presence of eggs in heavily gravid females or internal populations of the recently described A. clairae (Maryan examination of gonads. Head scale defi nitions follow et al. 2013a), and allow for an expanded account of those used by Storr et al. (1990), and methods of scale the morphology, distribution and habitat associations counting and morphometric measurements follow of this geographically restricted species. Finally, our those used by Maryan et al. (2013a, 2013b), with two results provide further evidence for as yet unresolved additional measures: body width and body depth. taxonomic complexity within A. repens sensu stricto. For the purpose of this study the following linear measurements reported in millimetres (mm) were taken with digital calipers or plastic ruler: snout-vent MATERIAL AND METHODS length measured from tip of snout to vent (SVL), head depth measured from a point immediately behind ALLOZYME ANALYSIS eyes (HD), head length measured from tip of snout Liver samples of those specimens with tissues to posterior margin of frontal scale (HL), head width extracted are stored at -70°C at the Western Australian measured from a point between eyes (HW), rostral Museum, Perth (WAM) or Evolutionary Biology Unit, length measured between anterior and posterior point South Australian Museum, Adelaide (SAMA), including of scale (RL), rostral width measured between lateral tissues from three of the four specimens of the putative extremes of scale (RW), snout length measured from new species (WAM R121129, WAM R146587, WAM tip of snout to anterior margin of eye (SL), body width R173106). These tissues were incorporated into an measured half way on body between lateral surfaces extended allozyme study that also included all previous (BW) and body depth measured half way on body analysed Aprasia tissues (Maryan et al. 2013a, 2013b) between dorsal and ventral surfaces (BD). Three as well as some previously uncharacterised samples of meristic counts were taken: number of midbody scale the A. repens species-group tissues from the central rows counted half way around body (Mbs), number of west coast of Western Australia. The allozyme study ventrals counted from immediately behind mental scale also included exemplars of all other species of Aprasia, to vent including precloacal scale (Vent) and number some of which were not included in previous studies. of vertebrals counted from immediately behind frontal Notable among the latter was the only available tissue scale to above vent (Vert). Specimens preserved in of A. picturata. Details of the 83 specimens used for the a circular or twisted position were straightened on a allozyme study are presented in the Appendix. Figure fl at surface when measured for snout-vent length. We 1 shows the geographic origin of the A. repens species- excluded tail length from our linear measurements and group specimens used in the allozyme study. multivariate analysis of morphological variation, as the Allozyme electrophoresis of liver homogenates was majority of tails in specimens were recently broken or undertaken as described in Richardson et al. (1986) obviously regenerated, as indicated by a clear break in for the same suite of enzymes as previously employed colouration. In any case, x-rays are necessary to reliably (Maryan et al. 2013a, 2013b). As in our earlier studies, distinguish between original and fully regenerated we fi rst used an individual-based analytical procedure tails in pygopodid lizards (G. Shea, pers. comm.) and (stepwise Principal Coordinates Analysis: PCO), before these were not taken during this study. In this study, we subsequently undertaking taxon-based assessments provide only tail length measured from tip of tail to vent of genetic affinities (Neighbour-joining tree). All in the designated holotype of the putative new species 14 B. MARYAN, M. ADAMS AND K.P. APLIN for descriptive purposes only (type lists in Taxonomy deny the possibility of some genetic interaction with one section). or more of the regionally sympatric species, leading to An index of body
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