DOI: 10.18195/issn.0312-3162.25(1).2008.087-093 J?ecords 01 the Western Australian t'vll1sclIJJ1 25: 1-\7-93 (2001-\).

Pygopus (: ) from mid-Holocene cave deposits, Western and South Australia

2 3 J Jim I. MeadIJ', Marci HolIenshead . , Sandra L Swift ', Christopher J. BelP and Alexander Baynes' '!Jepartment of Ccology, Northern Arizona University, Flagstaff, Arizona 1-\6011 USA; E-mail: james.Mead(onau.edu; "Departmcnt of Biological Sciences, Northern Arizona Universitv, Flagstaff, Arizona 1-\6011 USA. 'l,aboratory of Quaternarv Paleontology, Center for Environmental Sciences and Education, Northern Arizona University, Flagstaff, Arizona 1-\6011 USA. 'Department of Ceological Sciences, The jackson School of Ceoscicnccs, University of Texas at Austin, Austin, Tcxas 71-\712 USA. -'Department of Earth and Planetary Scicncl's, Wesll'rn Australian Museum, Locked Bag 49, Welshpool DC, WA 691-\6 Australia. Present address: Dcpartment of Ceosciences, East Tennessee State University, johnson City, Tennessee 37614 USA. E-mail: mead(uetsu.edu

Abstract - We report fossil remains of pygopodid of known or probable Ilolocenc deposits in Webbs Cave in Western Australia and Wombat Cave in South Australia. Knowledge of patterns of skeletal variation within and across pygopodid groups is insufficiently mature to permit -level resolution. Confident identification of apomorphies to diagnose our specimens is limited by a general paucity of skeletal preparations of pygopodids. Preserved anatomical features on the fossils are interpreted in the context of published literature and putative apomorphies, and permit referral to the Pvgoplls. These specimens represent important additional records of Pvgopus, and contribute to an improved understanding of the evolution of faunal communities in the Holocene of western and southern Australia.

INTRODUCTION northeast (Shea 1993). Al though today they are Several cave deposits in Western Australia were widespread throughout most of the continent, their excavated since the 1950s for late Quaternary fossil record is extremely sparse, with only one mammalian and archaeological remains, and known fossil from the Miocene of Riversleigh, produced a rich palaeontological record (Lundelius northwestern Queensland (Hutchinson 1997). 1960; Dortch 1996, 2(04). Marsupials and rodents Estimates of divergence times and mode of were the focus of many of these studies, and evolution and dispersal for the group are based extinctions and changes in their relative abundances mostly on inference from molecular data sets and through time are well documented. These changes tree topologies (e.g. Jennings et a1. 2(03). serve as climatic proxies or to document Although superficially snake-like in appearance, zoogeographic range variations (Lundelius 1960, pygopods retain a pectoral girdle with no external 1983; Balme et al. 1978; Baynes 1982). Squamate vestiges of front limbs and a pelvic girdle remains from these deposits also are abundant but occasionally with hind limb remnants (Creer 1989; have yet to be systematicallv studied in detail. Here Shea 1993). Based on a sequence of molecular data we report the occurrence of pygopodid (125 rRNA and c-m(5) Donne1lan et a1. (1999) remains recovered from cave deposits at Webbs and support the concept that Pygopodidae and Wombat caves (J-Iampton Tableland). Diplodactylidae are sister taxa. Similarlv, the most Pygopods (flap-footed lizards) are a family of recent taxonomic treatments recognize a gekkonoid lizards endemic to Australia and New monophyletic Pygopodidae as sister taxon to Cuinea (Shea 1993). Over 38 species are currently Diplodactylidae (l\ussell and HilLler 2002) or as recognized within the family, and are classified in sister to the padless Australian carphodactylines seven genera (Apra5ia, Delma, LiaJi5, (Flan et a1. 20(4). 1lmvever, details of cranial OphidioCt,'phaI1l5, Parade1ma, [,Ietholax, and osteology and myology are presently ambiguous in PygopII5; Jennings et a1. 2003; Han et al. 2004; supporting a definitive sister-taxon relationship Maryan et al. 2(07). Pygopods occur throughout between pygopods and other groups of gekkonoids most of Australia except for Tasmania, extreme (e.g. Kluge ]987; Estes et a1. 1988; Ri)11 and Henkel southeastern Australia, and the rainforests of the 2(02). The historical uncertainty of higher-level 88 J.I. Mead, M. Hollenshead, S.L. Swift, CJ. Bell, A. Baynes relationships of pygopods contributed to a variable for the pygopods and diplodactyline Indian (see differing views in Klllge 1987; Bauer Ocean 1990; King and Mengden 1990; Hutchinson 1997).

15 - CAVE DEPOSITS

20 - Webbs Cave Western -;!" Webbs Cave (6N 132; Western Australia; Figure Australia -_~~_....-.- 1) is situated on the Hampton Tableland south of South Australia the Nullarbor Plain at 31°46'S, 127°48'E, 25"- approximately 9.5 km north of MlIndrabilla homestead. The entrance is an open, shallow sinkhole with abundant skeletal remains under 30 ,,..... overhangs. Passages go to small chambers with bones and speleothems; these chambers are \ \ \ \ ultimately terminated by roof collapse (Lundelius 35 110 115 120' 125" 1963; E. L. Lundelius field notes, May 1964; Vertebrate Paleontology Laboratory, Texas Natural Figure 1 Map of Western Australia and adjacent Science Center, The University of Texas at Austin). regions showing the locations of Webbs and Sixteen species of fossil mammals were recorded Wombat caves from which mid-Holocene from the cave by Baynes (1987). Pygopus sp. lizard remains were collected. On top of a large collapse-boulder in the entrance Heavy line represents the coastline position during the full glacial low-stand sea level sinkhole are many years of bone and sediment (adapted from Dortch 1996). accumulation, and it is here that E. L. Lundelius recovered a Sarcophilus (Tasmanian devil) premolar in 1955 and conducted excavations in basal edges of the debris cones are buried by 1964. Bulk samples of bones and sediments were reddish-brown silt of unknown age. The reddish- removed from five depth intervals. These were brown silt continues to a depth of about 92 cm originally recorded in inches as 0-3, 3-6, 6-9, 9-12, (originally determined as 36 inches). Lasiorhinus and 12-15 inches below surface but are here treated latifrons (southern hairy-nosed wombat) dug as their metric equivalents of 0-7.5, 7.5-15.0, 15.0- extensively into this silt layer. Abruptly below the 23.0, 23.0-30.5, 30.5-38.0 cm, respectively. All reddish-brown silt is a brighter red sand and clay sieved residue, charcoal, and bones are curated at unit (that shows no sign of wombat burrowing or the Texas Memorial Museum, Vertebrate bioturbation) containing limestone boulders and Paleontology Laboratory, The University of Texas skeletal remains (E. L. Lundelius field notes, May at Austin (TMM). 1964). Lizard remains, including one pygopod The principal agents of bone accumulation in the specimen, were recovered from 92-106 cm below Nullarbor and Hampton Tableland caves were Tyto surface, in the lower red sand and clay unit. The alba, T. novaehollandiae (masked owl) and Falm pygopod specimen from this lower unit is clearly cenchroides (Australian kestrel; Baynes 1987). not modern based on stratigraphic placement. The Lizard remains were common throughout the actual age is not known but is presumed to be from Webbs Cave sedimentary deposit. Two pygopod the Holocene, if not late Pleistocene. Clearly more dentaries were recovered from the 15.0-23.0 cm and work is needed in this cave. All fossils recovered by 30.5-38.0 cm intervals. Charcoal from the 15.0-23.0 E. L. Lundelius in 1964 are curated in TMM. cm interval produced an AMS radiocarbon intercept age of Cal BP 4,160 (Beta-214963), and Pygopod remains represents a mid-Holocene record. With an Relatively few publications specifically address assumption of a constant deposition rate, the the osteology of pygopodid lizards. General aspects skeletal remains in the undated 30.5-38.0 cm of skeletal morphology were discussed by Camp interval would be approximately 5,200 years old. (1923; Lialis; Pygopus), Stokely (1947; ), Parker (1956; Aprasia), Underwood (1957; Aprasia, Wombat Cave Delma, Lialis, Pygopus) and Moffat (1973; Aprasia, Wombat Cave (5N 264) is located approximately Delma, Lialis, Pletholax, Pygopus). Pectoral and 42 km west of Koonalda on the Hampton Tableland pelvic girdle morphology were discussed by in western-most South Australia (Figure 1). This Fiirbringer (1870; Lialis, Pygopus), Cope (1892; small cavern has a low arched roof and three Pygopus) and Stephenson (1962; Aprasia, Delma, entrances, each with an associated debris cone. The Lialis, Pletholax, Pygopus). General features of the Mid-Holocene Pygopods from Caves 89 skull were noted by Kinghorn (1926) and Bellairs Webbs Cave and Kamal (1981); more extensive coverage of One right dentary (TMM 41209-964; 15.0-23.0 cm particular taxa was provided by Jensen (1901; interval) and one left dentary (TMM 41209-965; Aprasia), Kinghorn (1923; Aprasia), Broom (1935; 30.5-38.0 cm interval) were recovered (Figure 2A, Pygopus), McDowell and Bogert (1954; Aprasia, B). Both are complete and have the same De1ma, Lia1is, Pygopus), Stephenson (1962; Aprasia, osteological features. The groove for the Meckelian Delma, Lia1is, P1etho1ax, Pygopus) and Rieppel cartilage is completely enclosed by bone (a (1984b; Aprasia, P1etho1ax, Pygopus). Detailed universal feature within gekkonoids, but also description of mandibular morphology was present in all dibamids and xantusiids, and variably summarized by Hutchinson (1997). Additional present within gymnophthalmids, scincids, many published osteological data sets include vertebral non-acrodont iguanian groups an.d some morphology (Etheridge 1967; Hoffstetter and Gasc amphisbaenians (Greer 1970; MacLean 1974; Gans 1969), scleral ring formulae (Underwood 1970; 1978; Estes et al. 1988; Etheridge and de Queiroz Aprasia, De1ma, Lialis, and Pygopus), and the 1988; Bell et al. 2003). The angular process of each detailed information on specific anatomical regions dentary is near-complete and positioned well of the skeleton provided by Kluge (1976, 1987), posterior of the coronoid. The intramandibular Rieppel (1984a), and Lee (1998) as part of their septum is not extended posteriorly. The fossils do phylogenetic studies of the group. not belong to Aprasia, Ophidiocephalus, or Our analysis of the pygopodid fossil remains is Pletholax because the anterior contact for the based primarily on published descriptions by splenial is ventral to the four posterior-most tooth Hutchinson (1997) and comparisons made with positions. Loci for up to 15 teeth are present. The modern skeletal material. Identifications are based teeth are robust and each has an apical groove (see on morphological characters observed on modern discussion in Sumida and Murphy 1987 and specimens; an apomorphy-based approach was not Hutchinson 1997). The posterior edge of the tooth possible because sufficiently detailed analyses of row is raised dorsally to meet a high coronoid, a isolated skeletal elements of all pygopodids and character found only in Pygopus. Teeth decrease in their outgroups have yet to be conducted, and size and robustness from mesial to distal position. adequate skeletal collections of Australian These features (possible apomorphies) on the two gekkotans are lacking. Possible apomorphies are dentaries indicate that they belong to Pygopus. identified, and are combined with characters (the apomorphic status of which is not confirmed) that Wombat Cave appear to exclude referral to one or other pygopod A posterior fragment of a right dentary (TMM taxa. This approach permits referral to genus for all 41374-1 Figure 2C) was recovered from the red sand fossils considered here. We made no attempt to and clay unit at 92-106 cm below surface. The identify specimens to the species level. angular process of the dentary is near-complete and

A

B

c

Figure 2 Pygopodid dentaries from mid-Holocene cave deposits. A, Pygopus sp., right dentary (TMM 41209-964) from the 15.0-23.0 cm level (Cal BP 4,120) and B, Pygopus sp., left dentary (TMM 41209-965) from the 30.5- 38.0 cm level in Webbs Cave, WA. C, posterior fragment of a Pygoplls sp. right dentary (TMM 41374-1) from the red sand and clay unit at 92-106 cm level in Wombat Cave, SA. Scale equals 1 mm. 90 J.1. Mead, M. Hollenshead, S.L. Swift, CJ. Bell, A. Baynes

extends well posterior to the coronoid. The The distance from the coast mostly determines the intramandibular septum is not extended post- amount of rainfall on the Hampton Tableland and eriorly. The fragment contains tooth loci for seven Nullarbor Plain, which in turn, controls the teeth (excluding referral to Aprasia). The splenial is distribution of vegetation. The three vegetation not present but its articulation area is evident and zones recognized along the region are a mallee shows that it probably separated the prearticular scrub zone located nearest to the coast, arid scrub, from the dentary (thus excluding referral to and a treeless plain found furthest from the coast Aprasia, Ophidiocephalus or Pletholax); this feature (Martin 1973). During the mid-Holocene, vegetation is found in Delma, Pygopus and Paradelma. The patterns of the Nullarbor were broadly similar to anterior position of the splenial contact on the those of today, suggesting that the climate of these dentary is ventral to the four posterior-most tooth times was not substantially different. Prior analysis positions, and thus the specimen cannot represent of the recovered pollen from late Quaternary Aprasia, OphidiocephaIus, or Pletholax, but is Madura Cave sediments (located 65 km south and consistent with that condition found in Delma, west of Webbs Cave) provides a direct vegetation Pygopus and Lialis. Although the posterodorsal community reconstruction for the Nullarbor Plain area is somewhat damaged on TMM 41374-1, it (Martin 1973). We apply the results of that pollen shows the contact placements for both the labial and analysis from Madura Cave to Webbs and Wombat lingual projections of the coronoid to be at different caves because of their close proximity and roughly elevations in the bone (versus the same elevation equal distance from the coastline. found on Delma and Paradelma). Tooth crowns are Today Webbs and Wombat caves are about 35 km simple with blunt-pointed, slightly grooved apices, from the coastline. During glacial maximum, the and are not recurved as found on Lialis. This coastline was approximately 160 km south of its combination of characters suggests that the present position along the Hampton Tableland and specimen is a species of Pygopus. Nullarbor south of these caves. With the rise of sea level after the 18,000-15,000 yr B.P. low-stand, the coastline moved closer to the caves. The period DISCUSSION between 9,000 and 4,000 yr B.P. was one of dramatic The pygopods are a distinctive element of the changes in the vegetation mainly in response to an modern lizard fauna in Australia. A phylogenetic increase in rainfall, and therefore, an increase in the analysis of pygopods based on mitochondrial DNA, mallee (eucalypt Myrtaceae) scrub community. nuclear DNA, and morphological data led Jennings et After that time, the mallee scrub cover declined in a1. (2003) to support earlier studies that indicated that abundance, followed by development of the arid speciation of mesic-adapted biotas in the southeastern scrub to treeless plain seen in the region today and southwestern corners of the continent occurred (Martin 1973). largely within those geographic regions between 23 Pygopus was differentiated and established in and 12 million years ago as opposed to repeated Australia by the Miocene (Hutchinson 1997); dispersals between the regions. However, it is not however, we do not know when Pygopus moved clear whether or not dispersals occurred during cyclic into the region around Webbs and Wombat caves. changes from glacial to interglacial regimes of the Our data document that they were present, if not latest Pliocene and throughout the Pleistocene. Cave established, by approximately 4,200 years ago. Both sites along the greater Hampton Tableland (such as P. lepidopodus and P. nigriceps live today in the Webbs and Wombat caves; see others in Baynes 1987) greater Nullarbor and nearby desert areas in mainly and nearby Nullarbor Plain are positioned to record low shrubland with sandy soils (Storr et a1. 1990; east-west dispersals (if they occurred) through these Wilson and Swan 2003). climatic cycles. The requirement of a detailed chronology for most of these sites has yet to be attained. CONCLUSIONS Pygopus was identified from Webbs and It is interesting that only three dentaries of Wombat caves on the Hampton Tableland dating pygopods were identified from the thousands of to Cal BP 4,160 (and possibly earlier at Wombat bones recovered from both Webbs and Wombat Cave). Species of pygopods living in the region caves. At the generic level, the mid-Holocene fossils today include Aprasia inaurita, Lialis burtonis, are logical discoveries. The recovery of any Pygopus lepidopodus (likely at the caves today) pygopod remains is significant because all remains and P. nigriceps (Storr et al. 1990; Wilson and help reconstruct the timing for the development of Swan 2003). Based on the known living fauna, and the community mosaic that occurs today. Based on the mid-Holocene age of the remains, it is this scant and preliminary record, it appears that at predictable that some species of Pygopus would least Pygopus inhabited the communities outside be recovered from deposits within Webbs and the caves, as they do today, by the middle to late Wombat caves. Holocene. Mid-Holocene Pygopods from Caves 91

It is apparent from our studies that a precise, fossil deposits of WA. We thank Bryan Jennings isotopic-based chronology for each of the caves that (Harvard University) and Eric Pianka (University of contain fossil vertebrates is a prerequisite to truly Texas at Austin) for discussions about pygopods. understand the establishment of modern squamate Ernie Lundelius graciously provided copies of his community mosaics. Although this as yet has not field notes and took the time to discuss his work in been systematically implemented, it is beginning to WA during the 1950s and I960s. We appreciate the be put into practice (see Prideaux et al. 2007a,b). assistance of Lyndon Murray (University of Texas Additional Pleistocene deposits with lizard remains at Austin) for the curation of the specimens into the are needed to assess the development of the current Texas Memorial Museum, Vertebrate Paleontology biotic communities and how the faunas responded Laboratory. Partial funding for our research was to changing climate regimes and other influential received from the College of Engineering and factors. Not all paleontological deposits were Natural Sciences, Northern Arizona University. screened through mesh sieves small enough (less Edits and suggestions to an earlier draft of this than 500 ~lm) to recover isolated bones of small manuscript were received from Michael Lee (South lizards. The use of micro-mesh screens is now a Australian Museum), Mark Hutchinson and Paul consistent practice at select excavations. The ability Doughty; their help is greatly appreciated. to reconstruct the development of the modern lizard communities and the community mosaic changes through time for Western and South REFERENCES Australia largely are dependant on the capability to Balme, j., Merrilees, D. and Porter, j.K. (1978). Late identify accurately the recovered fossils. 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