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Systematics of the Snake-Eyed Skinks, Cryptoblepharus Wiegmann (Reptilia: Squamata: Scincidae) – an Australian-Based Review

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Systematics of the Snake-Eyed Skinks, Cryptoblepharus Wiegmann (Reptilia: Squamata: Scincidae) – an Australian-Based Review The Beagle, Records of the Museums and Art Galleries of the Northern Territory, Supplement 3 (2007): 2–98 Systematics of the snake-eyed skinks, Cryptoblepharus Wiegmann (Reptilia: Squamata: Scincidae) – an Australian-based review PAUL HORNER Museum and Art Gallery of the Northern Territory GPO Box 4646, Darwin NT 0801, AUSTRALIA [email protected] ABSTRACT Morphologically conservative, Cryptoblepharus was once treated as a single polytypic species comprising many geo- graphical subspecies and was considered the world’s most broadly distributed lizard. Concentrating on taxa from the Australian region, the study investigates morphological characteristics of the two lineages and operational taxonomic units genetically identified by Horner and Adams (2007). Morphological variation of the genetically identified populations was investigated by multivariate analyses of 21 meristic and 12 mensural variables, and resulted in the identification of 25 Australian taxa. Using comparable analyses, species extralimital to Australia were also investigated, resulting in the recognition of 13 taxa from the southwest Indian Ocean region and 24 from the Indo-Pacific region. Overall, Crypto- blepharus was determined to comprise 62 taxa, consisting of 48 monotypic and six polytypic species. Existing types were able to be assigned to 43 taxa and 9 are described as new. Accounts of each taxon are supplied, although those of taxa extralimital to Australia are based on small sample sizes and are less detailed. Dichotomous keys to the identification of taxa from each geographic region are provided. An hypothesis for the biogeography of the genus suggests that it originated in South-east Asia and achieved its present distribution by a combination of rafting and human mediated transport. KEYWORDS: Reptilia, Scincidae, Cryptoblepharus, new species, Australia, Indo-Pacific, South-west Indian Ocean, taxonomy, mor- phology. CONTENTS INTRODUCTION. 23 MATERIAL AND METHODS. 27 RESULTS OF MORPHOLOGICAL ANALYSES. .33 SYSTEMATICS . .48 Genus Cryptoblepharus Wiegmann, 834. 48 Australian Region Taxa. .52 Key to Australian Cryptoblepharus taxa. 52 Cryptoblepharus adamsi sp. nov. .54 Cryptoblepharus australis (Sternfeld, 98) . .57 Cryptoblepharus buchananii (Gray, 838) . 60 Cryptoblepharus cygnatus sp. nov. .64 Cryptoblepharus daedalos sp. nov. .68 Cryptoblepharus exochus sp. nov. .7 Cryptoblepharus fuhni Covacevich and Ingram, 978. .74 Cryptoblepharus gurrmul sp. nov. 76 Cryptoblepharus juno sp. nov. .79 Cryptoblepharus litoralis (Mertens, 958) . 82 Cryptoblepharus litoralis horneri Wells and Wellington, 985 . .83 Cryptoblepharus litoralis litoralis (Mertens, 958) . .85 Cryptoblepharus litoralis vicinus ssp. nov. .87 Cryptoblepharus megastictus Storr, 976. .88 Cryptoblepharus mertensi sp. nov. 90 Cryptoblepharus metallicus (Boulenger, 887). .93 Cryptoblepharus ochrus sp. nov. .97 Cryptoblepharus pannosus sp. nov. 00 Cryptoblepharus plagiocephalus (Cocteau, 836) . .04 Cryptoblepharus pulcher (Sternfeld, 98). .08 Cryptoblepharus pulcher clarus (Storr, 96). 110 Cryptoblepharus pulcher pulcher (Sternfeld, 98). .111 2 P. Horner Cryptoblepharus ruber Börner and Schüttler, 98 . 114 Cryptoblepharus tytthos sp. nov. .118 Cryptoblepharus ustulatus sp. nov. .2 Cryptoblepharus virgatus (Garman, 90). .24 Cryptoblepharus wulbu sp. nov. .27 Cryptoblepharus zoticus sp. nov.. .29 South-west Indian Ocean Region Taxa . .32 Key to South-west Indian Ocean Cryptoblepharus taxa. .32 Cryptoblepharus africanus (Sternfeld, 98) . .33 Cryptoblepharus ahli Mertens, 928 . .33 Cryptoblepharus aldabrae (Sternfeld, 98). .34 Cryptoblepharus ater (Boettger, 93). 35 Cryptoblepharus bitaeniatus (Boettger, 93) . 35 Cryptoblepharus boutonii (Desjardin, 83). .36 Cryptoblepharus caudatus (Sternfeld, 98). .36 Cryptoblepharus cognatus (Boettger, 88). 37 Cryptoblepharus gloriosus (Stejneger, 893). .38 Cryptoblepharus gloriosus gloriosus (Stejneger, 893). .38 Cryptoblepharus gloriosus mayottensis Mertens, 928. .39 Cryptoblepharus gloriosus mohelicus Mertens, 928. 39 Cryptoblepharus quinquetaeniatus (Günther, 874) . .40 Cryptoblepharus voeltzkowi (Sternfeld, 98) . 40 Indo-Pacific Region Taxa. 4 Key to Indo-PacificCryptoblepharus taxa. .4 Cryptoblepharus baliensis Barbour, 911. 43 Cryptoblepharus baliensis baliensis Barbour, 911 . .43 Cryptoblepharus baliensis sumbawanus Mertens, 928 . 44 Cryptoblepharus burdeni Dunn, 927. .44 Cryptoblepharus cursor Barbour, 911 . .45 Cryptoblepharus cursor cursor Barbour, 911. .45 Cryptoblepharus cursor larsonae ssp. nov. 46 Cryptoblepharus egeriae (Boulenger, 889) . .47 Cryptoblepharus eximius Girard, 857. 48 Cryptoblepharus furvus sp. nov. .48 Cryptoblepharus intermedius (de Jong, 926). .50 Cryptoblepharus keiensis (Roux, 90). .5 Cryptoblepharus leschenault (Cocteau, 832). .52 Cryptoblepharus nigropunctatus (Hallowell, 860). 52 Cryptoblepharus novaeguineae Mertens, 928 . .53 Cryptoblepharus novocaledonicus Mertens, 928 . .54 Cryptoblepharus novohebridicus Mertens, 928. .55 Cryptoblepharus poecilopleurus (Wiegmann, 834). .55 Cryptoblepharus poecilopleurus paschalis Garman, 908 . 56 Cryptoblepharus poecilopleurus poecilopleurus (Wiegmann, 834) . 56 Cryptoblepharus renschi Mertens, 928. .57 Cryptoblepharus richardsi sp. nov.. .57 Cryptoblepharus rutilus (Peters, 879). 60 Cryptoblepharus schlegelianus Mertens, 928. 60 Cryptoblepharus xenikos sp. nov. 6 Cryptoblepharus yulensis sp. nov.. 63 Cryptoblepharus sp. .65 DISCUSSION. .65 CONCLUSION . .69 ACKNOWLEDGMENTS. .69 REFERENCES . .70 Appendix . Results for Discriminant Function Analyses. .77 Appendix 2. Results for taxon comparisons by ANOVA. .79 Appendix 3. Comparative regional summary of morphological characters . 83 Appendix 4. Non-type material examined . .95 22 Systematics of the snake-eyed skinks INTRODUCTION are based on colour pattern and geographic distribution (29 are allopatric, insular species), with the few traditional In a companion paper Horner and Adams (2007) in- scalation characteristics mentioned being attributable to vestigated the genetic status of Australian populations of virtually any member of the genus. Literature relevant to the scincid genus Cryptoblepharus, ultimately recognising Cryptoblepharus is limited, and that available is mostly that species diversity in that region was greatly understated. restricted to aspects of taxonomy and biogeography, with This paper follows and complements Horner and Adams’s knowledge of reproduction, ecology and evolutionary his- (2007) work, investigating the morphological characteristics tory markedly lacking. of genetically identified and other presumptive taxa from Seven species of Cryptoblepharus are currently recog- Australia and other regions, assessing their taxonomic status nised as Australian taxa (Cogger 2000; Cogger et al. 983a; and describing or re-describing all known species. Stanger et al. 998; Ehmann 992; Greer 989; Wilson Cryptoblepharus is the most geographically widespread and Knowles 988; Wilson and Swan 2003): C. carnabyi taxon in the family Scincidae, the most species-rich, mor- Storr, 976; C. egeriae (Boulenger, 889); C. fuhni Co- phologically diverse and geographically widespread lizard vacevich and Ingram, 978; C. litoralis (Mertens, 958); group. Achieving greatest diversity in the Australian/New C. megastictus Storr, 976; C. plagiocephalus (Cocteau, Guinea and South-east Asian regions (Greer 970), Scin- 836) and C. virgatus (Garman, 90). All but one are cidae is found in most tropical and temperate regions of mainland species, with C. egeriae restricted to the Australian the world and comprises about 200 species in 27 genera Territory of Christmas Island in the Indian Ocean. As cur- (Uetz et al. 2000). rently recognised, C. fuhni, C. litoralis and C. megastictus Exhibiting a marked degree of morphological conser- have limited distributions in northern Australia, C. virgatus vatism, Cryptoblepharus are small (<55 mm snout-vent occupies much of eastern and far southern Australia, and length), heliotropic, arboreal or saxicoline skinks that C. carnabyi and C. plagiocephalus have broad continental range through three broad, geographic regions, the Ethio- distributions. pian-Malagasy (south-west Indian Ocean), Indo-Pacific Of Cryptoblepharus extralimital to Australia, Mertens and Australian. Historically Cryptoblepharus has been (93) recognised 4 taxa from the south-west Indian Ocean treated as monotypic, with the various forms considered to region: C. boutonii africanus (Sternfeld, 98); C. b. ahli be geographical races of a single species (Mertens 93), Mertens, 928a; C. b. aldabrae (Sternfeld, 98); C. b. ater a concept now considered oversimplified (Auffenberg (Boettger, 93); C. b. bitaeniatus (Boettger, 93); C. b. 1980) and many subspecies have been elevated to specific boutoni (Desjardin, 83); C. b. caudatus (Sternfeld, 98); status (e.g. Brygoo 986; Storr 976; Zug 99). Although C. b. cognatus (Boettger, 88); C. b. degrijsi Mertens, acknowledged as problematic (Dunn 927; Storr 976; 928a; C. b. gloriosus (Stejneger, 893); C. b. mayottensis Haacke 977; Crombie and Steadman 986) there has been Mertens, 928a; C. b. mohelicus Mertens, 928a; C. b. no recent attempt to systematically revise Cryptoblepha- quinquetaeniatus (Günther, 874), and C. b. voeltzkowi rus taxonomy. That 76 years has elapsed since Mertens’s (Sternfeld, 98). Geographical distribution of these is cen- monographic work on the genus is perhaps related to a tred on the Mozambique Channel, with two taxa occurring universal perception that the group is taxonomically com- on the
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