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A Review Ofthe Namaqua Gecko, Pachydactylus Namaquensis (Reptilia: Gekkonidae) from Southern Africa, with the Description of Two New Species

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A Review Ofthe Namaqua Gecko, Pachydactylus Namaquensis (Reptilia: Gekkonidae) from Southern Africa, with the Description of Two New Species S. Afr. J. Zool. 1996,31(2) 53--69 A review ofthe Namaqua gecko, Pachydactylus namaquensis (Reptilia: Gekkonidae) from southern Africa, with the description of two new species William R. Branch' Department of Herpetology, Port Elizabeth Museum, P.O. Box 13147, Humewood, 6013 South Africa Aaron M, Bauer Biology Department, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA David A. Good Museum of Natural Science, 119 Foster Hall, Louisiana State University, Baton Rouge, Louisiana 70803, USA Received 19 September 1995; accepted 22 January 1996 An analysis of morphological and allozyme variation in the Namaqua gecko, Pachydactylus namaquensis from southern Africa is presented. Three separately evolving lineages, well defined by morphology and allozyme var­ iation, are identified. The isolated southern population, occurring on the southern escarpment and Cape Fold Mountains surrounding the western Litlle Karoo, is named P. kladaroderrna sp. nov., and is characterized by a slit-like ear opening, low number of granules bordering the mental (3--6) and mental and adjacent infralabials (5- 13), the frequent (79%) occurence of the supralabial entering the nostril, and its drab brown base colouration. A northern population, occurring in southern Namibia and the Richtersveld is named P. haackei sp. nov" and is characterized by its more rounded or squared ear opening; high number of granules bordering the mental and adjacent infralabials (11-19), the general exclusion of the supralabial from the nostril (only 3.7% entry), and brighter, lighter colouration. It is further differentiated from P. kladaroderrna on the basis of fixed differences at 11 allozyme loci. Both new species differ from P. namaquensis, which is mainly restricted to Litlle Namaqua­ land, but is sympatric with P. haackei in the Lower Orange River region, by their more heterogenous dorsal scalation, smaller cloacal spurs, lack of spine-like tubercles on the lateral surfaces of the tail, and more fragile skin. The type locality of P. namaquensis is restricted to 'the vicinity of Springbok, Litlle Namaqualand, Northern Cape Province, South Africa'. "'To whom correspondence should be addressed . The) lizard fauna of southern Africa is one of the most diverse although the recognition of such groups in the absence of a 9 and0 speciose in the world (Bauer 1993) and yet it remains phylogenetic hypothesis may be imprudent. A non-phyloge­ 0 2 relatively poorly known. In an analysis of terrestrial verte­ netic alpha·tevel taxonomic revision of Pachydactylus sensu d e stricto is currently in progress (McLachlan, pers. comm.), but bratet diversity and endemism in southern Africa, Crowe (1990)a found that the southern and western parts of the sub­ the only formal 'groups' so far defined in the literature are the d ( continent constitute the area of highest regional endemicity. P. weberi and P. serval groups of McLachlan & Spence r e (1966) and the complex of Broadley (1977; see Heh reported that lizards exhibit moderate to high levels of P. capensis s i localizedl endemism throughout southern AfTica, with up to also Jacobsen 1989). More recently an analysis of morpho­ b 43%u specific endemism in the western arid zones of Namibia logical variation in the Pachydactylus punctatus complex P and the Cape. The western arid regions are an epicentre of (Bauer & Branch 1995) has revealed significant differences e speciationh (Poynton & Broadley 1978), particularly for between popUlations above and below the western escarp­ t geckos.y Of the 76 gecko species in southern Africa, 35 are in ment in Namibia, and has led to the recognition of two spe­ b the genus or in genera that make cies. Bauer & Branch (1995) also suggested that subtle d Pachydactylus Pachydacty­ e Iust sensu stricto paraphyletic (Branch 1988; Bauer 1990; differences in the lowland western species, P. scherzi, might n Bauera & Branch 1995). Members of the lineage occur reveal further species if larger samples became available and! r g throughout the subcontinent, generally are abundant, have or molecular methods could be employed. e restrictedc ranges, and show a great range of microhabitat spe­ At the end of the last century, Sclater (1898) described a n cializations.e Eight representatives of this group have been c large new gecko 'obtained in Namaqualand', naming it after i l found in sympatry in the Karoo National Park, South Africa its locality and placing it in the genus Elasmodactylus, r (Branche & Braack 1989), and II are broadly sympatric in the recently erected by Boulenger (1894) for tuberculosus from d arean between Kamanjab and the Skeleton Coast National Park Tanzania, and now known from Zaire, Zambia and adjacent u iny northern Nambia (Bauer, Branch & Haacke 1993). In both areas. Subsequently, Elasmodactylus namaquensis was areas,a the majority of species occur in rocky habitats where transferred to the genus Pachydactylus Wiegmann 1834 by w e theyt occupy cracks and surfaces of different types, or they are Methuen & Hewitt (1914). They noted that justification for terrestriaLa the recognition of Elasmodactylus as distinct fTom Pachydac­ G t i.e., the presence of claws on the toes in the former, is e Attempted revisions within Pachydactylus sensu stricto tylus, n havei met with limited success (Hewitt 1927; FitzSimons an inconstant character. Subsequent workers (Haacke 1976; b 1943;a Loveridge 1947; McLachlan & Spence 1966; McLach­ Joger 1980, 1984, 1985; Bauer & Good 1995) have demon­ S lan 1979). Several species groups have been recognized on strated that females of the Pachydactylus-group genera gener­ y b the basis of superficial similarities of habitus and scalation, ally possess claws on digits I and V of the pes. Jager (1985) d e c u d o r p e R 54 S. Afr. J. Zoo!. 1996, 31 (2) argued for the revival of Elasmodactylus for tuberculosus, but in the Port Elizabeth Museum (PEM). This was supplemented this was based on the retention of piesiomorphic characters with additional material and localities obtained from the fol­ such as the presence of preanal pores and a partly divided ros­ lowing museums: J. Ellennan Museum, Stellenbosch (JEM), tral, which he regarded as having transfonned in the puta­ South African Museum, Cape Town (SAM); Transvaal tively monophyletic group (Pachydacty/us proper + Museum, Pretoria (TM), California Academy of Sciences, Chondrodacty/us + Pa/matogecko + C%pus + Kaokogecko San Francisco (CAS), Museum of Vertebrate Zoology, Ber­ + Rhoptropus). With respectto P. namaquensis, Joger (1985) keley (MVZ), United States National Museum of Natural suggested that it was reasonably closely related to P. bibronii History, Washington (USNM), and American Museum of and reported an immunological distance of about 22%, corre­ Natural History (AMNH). Description of scutellation follows sponding to a divergence time of 20 million years. Joger's FitzSimons (1943), and data were obtained at x8 or x40 mag­ (1985) material was supposedly derived from 'Kharkamas, nification with a Carl Zeiss dissecting microscope. Cleared Namibia' (presumably Karkams, Linle Namaqualand, South and double stained specimens and radiographs prepared in a Arrica). Faxitron cabinet X-ray system were used to examine osteo­ Methuen & Hewin (1914) noted the first Namibian speci­ logical features. The following details were recorded. mens to be referred to Pachydactylus namaquensis from the Great Karasberg Mountains, and additional specimens from Sca/alion: Scales entering the nostril (recorded separately for Namaqualand were later recorded by Hewin (1932) and Fitz­ both sides of the head); number of granules separating Simons (1938). In his monographic revision of the subconti­ nasorostrals (if they were in contact behind the rostral, even if nent's lizards, FitzSimons (1943) summarized museum separated posteriorly by a granule(s), they were recorded as records for the species, including a new southern record from being in contact); number of granules bordering mental and the Wineberg, south of Matjiesfontein, that he later reported adjacent inrralabials; the number of supra- and infralabials on in more detail (FitzSimons 1946). Haacke (1965) extended (including small scales at the rictus); dorso-Iateral body the northern limit with a record from Farm Tiras, Bethanie tubercles (objective description of the degree of tuberculation District, Namibia. More recent range extensions referred to is difficult; the size of tubercles in the dorso-Iateral region the species have occurred in the south, including the Nuwe­ relative to the size of surrounding granules, and the average veldberge in the escarpment mountains of the central Karoo number of granules surrounding 10 enlarged tubercles were (Branch & Braack 1989; Branch 1990), the eastern Lange­ recorded); caudal tubercles (presence or absence of enlarged berg Mountains in the Cape Fold Mountains (Boycon 1990), spine-like tubercles on the lateral surfaces of the tail); cloacal and the western Linle Karoo (Burger 1993). The distribution spurs (size and number of enlarged, flanened tubercles on the of Pachydacty/us namaquensis, as understood and illustrated ventro-Iateral surface of the tail base); shape of ear opening . ) in Visser (1984) and Branch (1988), is disjunct, with a gap of (in non-distorted heads); claws (on the fingers and toes); skin 9 0 approximately 370 km separating the southern population tears (including rresh and healed damage). 0 2 rrom the closest record (Garies) of the Namaqualand and d Morphometries: The following measurements were
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