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Maryan, Brad, Brennan, Ian G., Adams, Mark, Aplin, Ken P. (2015): Molecular and morphological assessment of Delma australis Kluge (Squamata: Pygopodidae), with a description of a new species from the biodiversity ' hotspot' of southwestern Western Australia. Zootaxa 3946 (3): 301-330, DOI: http://dx.doi.org/10.11646/zootaxa.3946.3.1

Delma Gray, 1831 Type species: Delma fraseri Gray, 1831, by monotypy. Diagnosis. Delma differs from all other pygopodid lizard genera in possessing the following combination of characters: head scales, including the parietals, enlarged and symmetrical; anterior nasal scales nearly always in contact; first pair of lower labials in contact behind mental scale; nostril usually bordered by more than two scales (except in some D. impar); external ear opening large; 20 or fewer midbody scale rows; dorsal and ventral scales smooth; precloacal pores absent; tail about three times as long as body (except in the D. australis species-group)., Published as part of Maryan, Brad, Brennan, Ian G., Adams, Mark & Aplin, Ken P., 2015, Molecular and morphological assessment of Delma australis Kluge (Squamata: Pygopodidae), with a description of a new species from the biodiversity ' hotspot' of southwestern Western Australia, pp. 301-330 in Zootaxa 3946 (3) on page 316, DOI: 10.11646/zootaxa.3946.3.1, http://zenodo.org/record/244607, {"references":["Gray, J. E. (1831) Description of a new genus of ophisaurean animal, discovered by the late James Hunter, Esq., in New Holland. In: Gray, J. E. (Ed.), Zoological Miscellany, Treuttel, Wurtz & Co., London, pp. 14."]}

Delma australis species-group Diagnosis. Based on the molecular analysis of Jennings et al. (2003) and this study we formally propose a D. australis species-group, currently comprising D. australis, D. hebesa sp. nov. and D. torquata. The morphological characteristics of this species-group are relatively small-size (SVL to 93 mm); ventral scales usually not markedly larger than adjacent lateral scales (enlarged in some D. torquata); one pair of supranasal scales; modally 16 or 18 midbody scales (occasionally 17, 19 or 20); loreal scale row usually interrupted by a ventral extension of supraloreal scale that contacts upper labials. For comparisons of ventral body scale size between D. australis and D. fraseri see Kluge (1974: 9)., Published as part of Maryan, Brad, Brennan, Ian G., Adams, Mark & Aplin, Ken P., 2015, Molecular and morphological assessment of Delma australis Kluge (Squamata: Pygopodidae), with a description of a new species from the biodiversity ' hotspot' of southwestern Western Australia, pp. 301-330 in Zootaxa 3946 (3) on page 316, DOI: 10.11646/zootaxa.3946.3.1, http://zenodo.org/record/244607, {"references":["Jennings, W. B., Pianka, E. R. & Donnellan, S. (2003) Systematics of the lizard family Pygopodidae with implications for the diversification of Australian temperate biotas. Systematic Biology, 52, 757 - 780. http: // dx. doi. org / 10.1080 / 10635150390250974","Kluge, A. G. (1974) A taxonomic revision of the lizard family Pygopodidae. Miscellaneous Publications of the Museum of Zoology, University of Michigan No. 152, 1 - 72."]}

Delma hebesa sp. nov. Heath Delma (Figs. 9, 10, 11) Holotype: WAM R 144237, male, Bandalup Hill, Ravensthorpe Range (33 �� 40 ' 29 "S 120 �� 23 ' 54 "E), Western Australia, Australia, collected by R. Teale & G. Harold, 14 October 2000. Fixed in 10 % formalin, stored in 70 % ethanol, liver sample stored in ��� 80 ��C ultrafreezer at WAM. Paratypes: All from Western Australia. WAM R 129674, male, 3.8 km W of Kundip, (33 �� 41 'S 120 ��09'E); WAM R 131902, female, Hellfire Bay, Cape Le Grand National Park, (34 ��00' 15 "S 122 �� 10 ' 20 "E); WAM R 132154, female, Duke of Orleans Bay, Wharton Beach, (33 �� 56 'S 122 �� 33 'E); WAM R 144238, male, same details as holotype; WAM R 154234, male, Kundip, (33 �� 40 ' 26 "S 120 �� 11 ' 45 "E); WAM R 156978, male, Canal Rocks, (33 �� 39 ' 46 "S 115 ��00' 45 "E). Other material examined: A full list of material examined is provided at the end of the paper. Diagnosis. A small species of Delma (SVL to 79 mm) with: ventral scales not markedly larger than adjacent lateral scales; one pair of supranasals; modally 18 midbody scales; modally 10 hindlimb scales in both sexes; 73���92 ventral scales (males average 76.8, females 85.5); six upper labials with fourth typically below eye; loreal scale row typically interrupted by a ventral extension of supraloreal scale that contacts upper labials; essentially unpatterned head, sometimes with weak dark variegations on sides of head and indistinct narrow bars or smudges on labial scales, nape and forebody. Diagnostic differences between D. hebesa sp. nov. and D. australis are listed under the foregoing species account. Delma hebesa sp. nov. differs from D. torquata from southeastern Queensland in having a larger adult size (SVL to 79 mm versus to 63 mm), three precloacal scales (versus two), the fourth upper labial typically below the eye (versus typically the third below the eye), modally 18 midbody scale rows (versus 16) and only dark variegations (if present) on head and neck (versus broad dark bands). Delma hebesa sp. nov. differs from all other Australian species (except D. australis, D. torquata and D. concinna) in having ventral scales not markedly larger than adjacent lateral scales (versus markedly larger). Delma hebesa sp. nov. differs from D. fraseri with which it occurs in sympatry (see below) in having a smaller adult size (SVL to 79 mm versus to 140 mm, Bush et al. 2007), one pair of supranasals (versus two pairs), modally 18 midbody scale rows (versus 16), ventral scales not markedly larger than adjacent lateral scales (versus markedly larger) and only dark variegations (if present) on head and neck (versus broad dark bands, often faded in adults). Description of holotype (Fig. 9). Measurements (in mm) and meristic values: SVL 57, HD 3.4, HL 6.9, HW 4.5, HLL 3.3, ML 4.9, RL 0.9, RW 1.3, SL 2.6, EW 1.1, HLS 10, MSR 18, VE 75. Head short and narrowing gradually forward of eyes, of equal width to body posteriorly; obvious tympanic aperture, indicated by round opening posterior to corner of mouth, opening is more narrow on right side; snout moderately long and rounded in dorsal and lateral profiles; body moderately robust of equal width and round in cross-section; hindlimbs visible as well-developed elongate, rounded flaps adpressed to body at lateral extremes of vent; tail relatively short, tapering very gradually distally to a pointed tip. Head scales smooth, non-imbricate and heterogeneous; rostral rounded anteriorly, wider than long, with obtuse apex projecting between supranasals; one pair of supranasals in broad contact, angled backwards posteromedially behind rostral and in short contact with first upper labial; nostril positioned on posterior junction of supranasal with first upper labial and postnasal; one postnasal, wider than high, angled posteroventrally and in short contact with second upper labial; prefrontals symmetrical and in broad contact; supraloreal much higher than wide, and in broad contact with second upper labial; four loreals, the anterior most slightly larger; five supraciliaries, first and fourth the smallest, second and fifth the largest; two supraoculars, first slightly larger than second; two frontals, the anterior most slightly wider and larger; two parietals, one on left side is slightly larger and longer; occipital present; two upper temporals; six upper labials, fourth the widest and positioned below eye, third the smallest, on left side a small scale intersects suture between second and third upper labial; five lower labials, second the largest and widest, fifth the smallest; mental wider than long with suture starting half-way along first upper labial. General form of head and details of scalation illustrated in Fig. 9 A, B. Body scales smooth, non-imbricate, homogeneous, and arranged in parallel longitudinal rows; ventral scales only very marginally wider than the adjacent lateral body scales; three precloacal scales. In preservative, due to the leaching of colours in ethanol after 14 years, the holotype (Fig. 9) has lost the grey colouration on the dorsal surface which is now light brown. All other aspects of dark bars or smudges on lower labials and obscure variegations on body remained. Colouration in life. The following description of colouration in life is based on Figs. 10 A, B, 11 A and field observations of D. hebesa sp. nov. from the Esperance Plains. Top of head bluish to light grey and unpatterned or with weak dark variegations. Lower labials whitish with blackish bars or smudges, centred on sutures of mental and anterior two lower labial scales, corner of mouth, grading to obscure variegations or weak bars around ear opening and on lateral scales of forebody. There is only very weak ventral extension of black bars or smudges on to chin and throat. Dorsal body surface bluish to light grey anteriorly, gradually merging to light brown body than to light grey tail. Lower flanks are slightly pinkish to reddish brown. Dorsal surface is uniform, except for some indication of obscure dark spots or variegations, particularly on some sutures of body scales. Ventral surface under head and along body whitish, with blackish variegations, becoming less pigmented under tail. The longest tail measured was 140 mm (208 % of SVL) on WAM R 131902. For comparisons of both D. hebesa sp. nov. and D. australis in life see Thompson & Thompson (2006: 44, ���grey morph��� and ���brown morph���), Bush et al. (2007: 135) and Henkel (2010: 141). Variation. Table 5 presents the means, standard deviations and ranges of the characters counted and measured for each sex of D. hebesa sp. nov. Table 8 presents the individual measurements and meristic counts for the type series of D. hebesa sp. nov. As noted in D. australis, the variation in the contact of the supraloreal with the upper labials and interrupting the loreal scale row or not, is similarly recorded in D. hebesa sp. nov. as the following conditions: the supraloreal contacts second upper labial on both sides (as in holotype) in WAM R 129674 and WAM R 154234; it contacts second and third upper labial on both sides in WAM R 131902 and WAM R 144238, and is separated from upper labials by either one large loreal and an elongate postnasal or two smaller loreals in WAM R 156978. One specimen (WAM R 154234) has five upper labials unilaterally, with the third under eye. All the type material has 18 midbody scales and 2���4 loreals. Variation in colouration and pattern. Most s pecimens are similarly coloured to the holotype in life and in preservative. Occasional preserved individuals (e.g. WAM R 131902) have more pronounced dark bars on the lower labials and lateral scales of forebody, while others (e.g. WAM R 154234) are a very uniform dark grey on head, body and tail. In life, hatchlings of D. hebesa sp. nov. have a similar colour and head pattern to adults (B. Bush, pers. comm.). Etymology. The specific name hebesa is derived from the Latin adjective hebes, meaning dull, alluding to the matt body texture, without much shine, of this species. Distribution and sympatry. Delma hebesa sp. nov. is widespread on the Esperance Plains bioregion and patchily distributed on the Warren and southern Jarrah Forest bioregions (Thackway & Cresswell 1995) in southwestern Western Australia (Fig. 8). Records extend east to the vicinity of Thomas River and Cape Arid, west to Canal Rocks and near Busselton, and inland to Stirling Range National Park, Ongerup, Ravensthorpe Range, Scaddan and Mount Burdett. The geographic distributions of D. hebesa sp. nov. and D. australis appear to be parapatric (Fig. 8). Currently the two species are known to occur within 80 km of each other in the east: WAM R 91740 of D. hebesa sp. nov. from Cape Arid versus WAM R 36229 of D. australis from Pine Hill and within 130 km in the west: WAM R 42637 of D. hebesa sp. nov. from Ongerup versus WAM R 12604 of D. australis from Wagin. Specimens from these parapatric localities do not show any indication of morphological intermediacy. The only recorded instance of sympatry involving D. hebesa sp. nov. is with D. fraseri (Bush 1981, 1984, as D. australis). Habitat. Delma hebesa sp. nov. occupies the proteaceous scrub and mallee heath on south coast sandplains (Beard 1990; Comer et al. 2001; Fig. 12). This habitat preference is exemplified of 65 records of D. hebesa sp. nov. from 18 sites, in which 79 % were recorded from mallee heath (Sanders et al. 2012, as D. australis). These diverse vegetation communities provide ample cover for D. hebesa sp. nov., where most specimens, including the type series, have been pit-trapped and raked from leaf litter, spoil-heaps, and mats of dead vegetation and inside abandoned stick-ant (Iridomyrmex conifer Forel) nests. It has also been found under logs, mallee roots, rocks and rubbish, especially pieces of corrugated iron in disturbed areas adjacent to uncleared heath (Bush 1981). It also occupies the granitic heath where it is occasionally found under exfoliated granite slabs (B. Maryan, pers. obs.). There are no habitat data associated with specimens from the lower southwestern corner of Western Australia (WAM R 129003, WAM R 156978, WAM R 172507). Remarks. Bush (1981: 21), Wilson & Knowles (1988: 246) and Bush et al. (2007: 135 a) illustrated D. hebesa sp. nov. as D. australis. Additionally, Wilson & Knowles (1988: 96) and Wilson & Swan (2013: 144) refer to D. australis individuals from the far southwest of range as bluish grey, a colouration feature consistent with D. hebesa sp. nov. Historically, D. australis has been documented as not occurring in the humid deep southwest of Western Australia (Kluge 1974; Wilson & Knowles 1988; Storr et al. 1990). However, there are a few records of D. hebesa sp. nov. in this area where they appear to be scarce when compared to the Esperance Plains. This apparent scarcity is documented by How et al. (1987), who recorded ��� D. australis ��� from only Two Peoples Bay in a survey of herpetofauna between Busselton and Albany. Biological surveys conducted on the Esperance Plains to date indicate D. hebesa sp. nov. to be widespread and locally abundant in areas of suitable habitat (Chapman & Dell 1975; Bush 1985; Chapman & Newbey 1995; Sanders et al. 2012, as D. australis). Bush (1983, 1984, as D. australis) provides additional information on reproduction in captivity and field observations of winter aggregations of D. hebesa sp. nov. from the Esperance Plains. The Esperance Plains bioregion is a biogeographically significant area rich in endemic plants, rare ecosystems, and vulnerable and specially protected fauna (Comer et al. 2001). Approximately 87 % of the Esperance Plains bioregion has been cleared and developed for intensive agriculture. However, much of the remaining vegetation is afforded statutory protection, including many nature reserves and national parks (Cape Arid, Cape Le Grand, Stokes, Fitzgerald River and Stirling Range National Parks) where D. hebesa sp. nov. is known to occur., Published as part of Maryan, Brad, Brennan, Ian G., Adams, Mark & Aplin, Ken P., 2015, Molecular and morphological assessment of Delma australis Kluge (Squamata: Pygopodidae), with a description of a new species from the biodiversity ' hotspot' of southwestern Western Australia, pp. 301-330 in Zootaxa 3946 (3) on pages 321-325, DOI: 10.11646/zootaxa.3946.3.1, http://zenodo.org/record/244607, {"references":["Bush, B., Maryan, B., Browne-Cooper & Robinson, D. (2007) Reptiles and Frogs in the bush: Southwestern Australia. University of Western Australia Press, Perth, 302 pp.","Thompson, S. A. & Thompson, G. G. (2006) Reptiles of the Western Australian Goldfields. Goldfields Environmental Management Group, Kalgoorlie-Boulder, 140 pp.","Henkel, F. W. (2010) Terralog Vol. 10: Geckos of Australia. Edition Chimaira, Frankfurt am Main, Germany, 160 pp.","Thackway, R. & Cresswell, I. D. (1995) An Interim Biogeographic Regionalisation for Australia: A Framework for Setting Priorities in the National Reserves System Cooperative Program Version 4.0. Australian Nature Conservation Agency, Reserves Systems Unit, Canberra.","Bush, B. (1981) Reptiles of the Kalgoorlie-Esperance Region. B. Bush, Perth, 46 pp.","Bush, B. (1984) Seasonal aggregation behaviour in a mixed population of legless lizards, Delma australis and D. fraseri. Herpetofauna, 16, 1 - 6.","Beard, J. S. (1990) Plant Life of Western Australia. Kangaroo Press, Sydney, 317 pp.","Comer, S., Gilfillan, S., Barrett, S., Grant, M., Tiedemann, K. & Anderson, L. (2001) Esperance 2 (ESP 2 - Recherche subregion). A Biodiversity Audit of Western Australia's 53 Biogeographical Subregions in 2002. Department of Conservation and Land Management, Perth. [unkown pagination]","Sanders, A., Chapman, A., Teale, R. J. & Harold, G. (2012) Vertebrate fauna of the Fitzgerald Biosphere Reserve, Western Australia. The Western Australian Naturalist, 28, 141 - 253.","Wilson, S. K. & Knowles, D. G. (1988) Australia's Reptiles. A Photographic Reference to the Terrestrial Reptiles of Australia. Collins, Sydney, 447 pp.","Wilson, S. & Swan, G. (2013) A Complete Guide to Reptiles of Australia. Fourth Edition. New Holland Publishers, Sydney, 592 pp.","Kluge, A. G. (1974) A taxonomic revision of the lizard family Pygopodidae. Miscellaneous Publications of the Museum of Zoology, University of Michigan No. 152, 1 - 72.","Storr, G. M., Smith, L. A. & Johnstone, R. E. (1990) Lizards of Western Australia. III. Geckos & Pygopods. Western Australian Museum, Perth, 141 pp.","How, R. A., Dell, J. & Humphreys, W. F. (1987) The ground vertebrate fauna of coastal areas between Busselton and Albany, Western Australia. Records of the Western Australian Museum, 13, 553 - 574.","Chapman, A. & Dell, J. (1975) Reptiles, Amphibians and Fishes. In: A biological survey of Cape Le Grand National Park. Records of the Western Australian Museum, 1 (Supplement), pp. 34 - 38.","Bush, B. (1985) Some reptiles and frogs recorded in Stokes National Park. The Western Australian Naturalist, 16, 52.","Chapman, A. & Newbey, K. R. (1995) A biological survey of the Fitzgerald area, Western Australia. CALM Science, 3 (Supplement), 1 - 258.","Bush, B. (1983) A record of reproduction in captive Delma australis and D. fraseri (Lacertilia: Pygopodidae). Herpetofauna, 15, 11 - 12."]}

Delma australis Kluge, 1974 Marble-faced Delma (Figs. 6, 7, 11) Holotype: WAM R 27359, male, Port Lincoln (34 �� 44 'S 135 �� 52 'E), South Australia, Australia, collected by G.M. Storr, 19 October 1966. Paratype: WAM R 24528, female, 37 km ENE of Wirrula (32 �� 22 'S 134 �� 54 'E), South Australia, Australia. Other material examined: A full list of material examined is provided at the end of the paper. Revised diagnosis. A small species of Delma (SVL to 93 mm) with: ventral scales not markedly larger than adjacent lateral scales; one pair of supranasals; typically 18 midbody scales; 68���92 ventral scales (males average 76.3, females 83.5); six upper labials typically with fourth below eye; loreal scale row typically interrupted by a ventral extension of supraloreal scale that contacts upper labials; modally 5���7 hindlimb scales in both sexes; strong dark variegations on upper surface of head; narrow dark bars on side of head (extending onto labial scales), nape and forebody. This revised diagnosis is essentially unchanged from those provided by previous authors (Kluge 1974; Storr et al. 1990; Shea 1991), despite the exclusion herein of D. hebesa sp. nov. Delma australis differs from the closely related D. torquata of southeastern Queensland in: larger adult size (SVL to 93 mm versus to 63 mm); three precloacal scales (versus two); the fourth upper labial scale typically below the eye (versus typically the third below the eye); modally 18 midbody scale rows (versus 16); and dark variegations or narrow bars (if present) on head, neck and forebody (versus broad dark bands). It differs from D. hebesa sp. nov. in: hindlimb scale counts in both sexes modally 5���7 (versus > 9); body colour brownish on head and tail (versus greyish on head and tail); head, nape and lateral scales of forebody with strong dark variegations or narrow barring (versus weak variegations); dark barring on head typically extends ventrally onto the chin and throat (versus indistinct dark bars or smudges present on the lower labials); and dark pigment on rostral and lower labials not aligned with sutures (versus dark smudges positioned over sutures between rostral and lower labials). Description of holotype (Fig. 6). Measurements (in mm) and meristic values, as determined during this study: SVL 65, HD 3.5, HL 7.5, HW 5.0, HLL 3.2, ML 4.7, RL 1.0, RW 1.7, SL 2.8, EW 1.3, HLS 10, MSR 19, VE 71. Head short and blunt, narrowing very gradually forward of eyes, of equal width to body posteriorly; obvious tympanic aperture, indicated by round opening directly posterior to corner of mouth; snout blunt and rounded in dorsal profile, rounded in lateral profile; body moderately robust of equal width and round in cross-section; hindlimbs visible as well-developed elongate, rounded flaps adpressed to body at lateral extremes of vent; tail relatively short, tapering very gradually distally to a pointed tip. Head scales smooth, non-imbricate and heterogeneous; large rostral blunt anteriorly, wider than long, with obtuse apex projecting between supranasals; one pair of supranasals in broad contact, angled backwards posteromedially behind rostral and in short contact with first upper labial anterior to nostril; nostril positioned on posterior junction of supranasal with first upper labial and postnasal; one postnasal, much wider than high and narrower posteriorly, slightly angled posteroventrally and in broad contact with second upper labial; prefrontals symmetrical, in broad contact; one supraloreal, much higher than wide and in broad contact with second upper labial; four loreals, the anterior most much larger; five supraciliaries, first and fourth the smallest, second the largest; two supraoculars, first slightly larger and wider than second; two frontals, the posterior most slightly wider and larger; two parietals, one on left side fused (partial suture evident) with upper temporal; occipital present; two upper temporals; six upper labials, fourth the widest and positioned below eye, third the smallest and fifth the highest; five lower labials, second the largest and widest, fifth the smallest; mental wider than long with suture starting half-way along the first upper labial. General form of head and details of scalation illustrated in Fig. 6 A, B. Body scales smooth, non-imbricate, homogeneous, and arranged in parallel longitudinal rows; ventral scales only very marginally wider than the adjacent lateral body scales; three precloacal scales. After more than 40 years in preservative (Fig. 6), the holotype is light brown on the dorsal surface with a slightly darker head bearing dense black variegations. Lower labials whitish with distinct black bars positioned centrally on first to third lower labial scales. Black bars continue on to corner of mouth, around ear opening and on the lateral scales of forebody. A faint dark bar is also present within mental scale. Dark bars extend ventrally onto chin and throat. Ventral surface under head and along body is whitish. Variation in measurements and scalation. Table 5 presents the means, standard deviations and ranges of the characters counted and measured for each of the three geographic populations of D. australis, as defined in Material and Methods. Data are presented separately for each sex. Most head scales in D. australis display minimal intraspecific variation. However, the condition of the supraloreal (described as a prefrontal by Shea 1991) extending ventrally to contact the upper labials, thereby interrupting the row of small loreals, is variable in this species. The most common condition is contact with the upper labials, as noted by Storr et al. (1990: 115), Shea (1991: 115) and shown in Figure 6 B. Our examination of 97 specimens of D. australis from Western and South Australia recorded the supraloreal (sometimes the postnasal and prefrontal) contacting the upper labials in over 70 % of specimens. The opposing condition of no contact between the supraloreal and upper labial scales was mostly caused by the presence of either one large loreal (sometimes with elongate postnasal), or two smaller loreals extending continuously from the postnasal to the subocular upper labial. We observed 0���5 loreals in this species, with one loreal being the typical condition, as reported by Storr et al. (1990). Most specimens had six upper labials with the 4 th below the eye but we saw occasional individuals with five (3 rd below the eye) or seven (5 th below the eye), mostly unilaterally. Over 70 % of specimens examined had the typical 18 midbody scales. Most others had 20 midbody scales, with occasional counts of 17 and 19, corresponding to the range given by Storr et al. (1990). One specimen (SAMA R 36487) has extensively fused head scales, with the supraloreal and prefrontal fused on the left side only and only four upper labials on the right side. The longest tail measured was 162 mm (245 % of SVL) on WAM R 112667. Kluge (1974: 78) and Storr et al. (1990: 115) illustrate the general form of the head and details of scalation of D. australis. Variation in colouration and pattern. Shea (1991: 72���73) documented geographic variation in the intensity of the head pattern in this species in South Australia. He distinguished and mapped a ���patterned��� form from the southern half and northwest of the state (Fig. 7 A; Cogger 2014: 388) and an ���unpatterned��� form from the western Lake Eyre drainage (Fig. 7 B). The holotype and 15 paratypes of D. australis from the Eyre Peninsula (south of the Gawler Ranges) in South Australia (Kluge 1974) are considered representative of the ���patterned��� form (Shea 1991: Fig. 1) in having strong dark variegations on the head, although the pattern is reduced, particularly laterally, in a few specimens. The ���patterned��� form is contiguous and similar to populations in northwestern Victoria and southwestern New South Wales (Swan et al. 2004: 49; Swan & Watharow 2005: 26; Wilson & Swan 2013: 145). Kluge (1974: 78) illustrates a preserved adult male specimen (SAMA R 10375) of the ���patterned��� form from near Kokatha in South Australia. In addition, strongly patterned individuals of D. australis have been illustrated by Ehmann (1992: 87) and Henkel (2010: 141). Western Australian populations of D. australis also display geographic patterning in colouration. Strongly patterned individuals with dark variegations and narrow bars on the lateral scales of forebody occur throughout the eastern Coolgardie Goldfields (Fig. 7 C) and Mallee bioregions (Fig. 7 D) (Thackway & Cresswell 1995). Particularly strongly patterned individuals are found on the Houtman Abrolhos (Fig. 7 E), while the population found to the immediate north around Shark Bay has very reduced patterning, with some individuals almost having dark brown uniform heads (Fig. 7 F), similar in many respects to the unpatterned form from South Australia. By contrast, a unique representative of the group (WAM R 132470) from the North West Cape has a rich brick-red body with an intensely black head flecked with small pale spots (Fig. 7 G). Distribution and sympatry. Delma australis is widespread throughout the subhumid to arid areas of southern Australia, from northwestern Victoria, and southwestern New South Wales, through most of South Australia and adjacent southern Northern Territory to southern and central west Western Australia (Wilson & Knowles 1988; Shea 1991; Swan et al. 2004; Wilson & Swan 2013; Fig. 8). In Western Australia, it extends north to Shark Bay (base of Peron Peninsula), Meedo Station, Weld Range, Paynes Find, Windarling Hill and Buningonia Spring, south through the Avon Wheatbelt, Mallee and Coolgardie Goldfields bioregions, and east to Cocklebiddy. There is a disjunct population on the North West Cape, represented by a single specimen from Shothole Canyon in the Cape Range (Fig. 8). Other possible outlier populations in the mid-west of Western Australia are Walyering Hill, Oakajee and near Kalbarri. Insular populations occur on Rat and Middle Islands in the Houtman Abrolhos. Recorded instances of sympatry involving D. australis include D. butleri, D. fraseri, D. grayii Smith, D. nasuta Kluge and D. petersoni Shea (Dell & Chapman 1981; Dell & How 1984; Chapman & Dell 1985; Shea 1991; Mckenzie et al. 1993; B. Maryan & G. Shea, per. obs.). For instance, in the vicinity of Poochera in South Australia, D. australis, D. butleri and D. petersoni have all been recorded (Shea 1991: 82). Four other species of Delma are recorded on the North West Cape. Delma nasuta and D. tincta are known from several localities (Maryan et al. 2007), while D. tealei Maryan, Aplin & Adams has been collected at Shothole Canyon and other localities on the Cape Range. Delma butleri is known from a single specimen collected at the Learmonth Air Weapons Range, immediately south of the Cape Range National Park. Habitat. In northwestern Victoria and southwestern New South Wales D. australis mainly occupies mallee habitats with a spinifex (Triodia) understorey (Swan et al. 2004; Swan & Watharow 2005). This habitat association is repeated in South Australia where the majority of the southern and western populations are from Triodia or mallee habitats, or combination of both (Shea 1991). The western Lake Eyre Basin population lives on gibber plain with Atriplex, on watercourses lined with Eucalyptus, and on low, stony hills with drainage channels and Acacia (Shea 1991; B. Maryan, pers. obs.). In Western Australia, D. australis occupies a variety of habitats growing on different soils, including mallee and/or other Eucalyptus woodlands and Acacia with a spinifex (Triodia and Plectrachne) or shrubland understorey (Storr et al. 1990; Smith et al. 1997; Bush et al. 2007). The habitat at Cape Range on the North West Cape consists of a heavily dissected limestone plateau sparsely vegetated with Triodia, shrubs and low eucalypts; gorges within the range are more heavily vegetated (Storr & Hanlon 1980). These diverse vegetation communities provide ample cover for D. australis, where most specimens have been pit-trapped, found in and under spinifex and sedge tussocks, raked from leaf litter, spoil-heaps, mats of dead vegetation and found under logs, mallee roots, rocks (including coral slabs on the Houtman Abrolhos) and rubbish, especially pieces of corrugated iron in disturbed areas adjacent to uncleared vegetation. When found in sympatry with other Delma species, D. australis tends to occur in moister microhabitats (Shea 1991; Wilson & Swan 2013). Interestingly, nocturnal observations of D. australis on sealed roads or tracks are rare, unlike other, larger-bodied species of Delma. Remarks. Using morphology alone, Kluge (1974) recognized and assessed differences between three geographic samples of male D. australis: a southwestern Western Australia sample (south of 32 �� 30 ���S, west of 120 ��E), an Eyre Peninsula of South Australia sample (south of 32 ��S), and a Victorian sample. He found significant mean differences in number of preorbital, preanal, hindlimb and caudal scales; as well as throat pattern and visceral pigmentation. A multivariate analysis of these six characters also revealed significant differentiation among the three regions. However, Kluge���s character definitions were different to those used in this study, and were based on a very small sample size (n= 3 per locality = 7, 8 and 12 respectively, with the Western Australian sample potentially including both D. australis and D. hebesa sp. nov.). Further studies in the laboratory and field are needed to determine whether or not the geographic sub-populations identified in each of South Australia and Western Australia correspond with the ���northern��� and ���southern��� groups distinguished by allozyme differences in the present study. Large gaps remain in the geographic sampling of all regional populations and the collection of additional specimens at or near potential contact zones would be particularly valuable to establish the status of the various forms., Published as part of Maryan, Brad, Brennan, Ian G., Adams, Mark & Aplin, Ken P., 2015, Molecular and morphological assessment of Delma australis Kluge (Squamata: Pygopodidae), with a description of a new species from the biodiversity ' hotspot' of southwestern Western Australia, pp. 301-330 in Zootaxa 3946 (3) on pages 316-321, DOI: 10.11646/zootaxa.3946.3.1, http://zenodo.org/record/244607, {"references":["Kluge, A. G. (1974) A taxonomic revision of the lizard family Pygopodidae. Miscellaneous Publications of the Museum of Zoology, University of Michigan No. 152, 1 - 72.","Storr, G. M., Smith, L. A. & Johnstone, R. E. (1990) Lizards of Western Australia. III. Geckos & Pygopods. Western Australian Museum, Perth, 141 pp.","Shea, G. M. (1991) Revisionary notes on the genus Delma (Squamata: Pygopodidae) in South Australia and the Northern Territory. Records of the South Australian Museum, 25, 71 - 90.","Cogger, H. G. (2014) Reptiles & Amphibians of Australia. Seventh Edition. CSIRO Publishing, Melbourne, 1033 pp. http: // dx. doi. org / 10.1086 / 678659","Swan, G., Shea, G. & Sadlier, R. (2004) A Field Guide to Reptiles of New South Wales. Second Edition. New Holland Publishers, Sydney, 302 pp.","Swan, M. & Watharow, S. (2005) Snakes, Lizards and Frogs of the Victorian Mallee. CSIRO Publishing, Melbourne, 91 pp.","Wilson, S. & Swan, G. (2013) A Complete Guide to Reptiles of Australia. Fourth Edition. New Holland Publishers, Sydney, 592 pp.","Ehmann, H. (1992) Encyclopedia of Australian Animals. Reptiles. Angus & Robertson, Sydney, 495 pp.","Henkel, F. W. (2010) Terralog Vol. 10: Geckos of Australia. Edition Chimaira, Frankfurt am Main, Germany, 160 pp.","Thackway, R. & Cresswell, I. D. (1995) An Interim Biogeographic Regionalisation for Australia: A Framework for Setting Priorities in the National Reserves System Cooperative Program Version 4.0. Australian Nature Conservation Agency, Reserves Systems Unit, Canberra.","Wilson, S. K. & Knowles, D. G. (1988) Australia's Reptiles. A Photographic Reference to the Terrestrial Reptiles of Australia. Collins, Sydney, 447 pp.","Dell, J. & Chapman, A. (1981) Reptiles and frogs of East Yuna and Bindoo Hill Nature Reserves. In: Biological survey of the Western Australian Wheatbelt. Part 14. Records of the Western Australian Museum, 13 (Supplement), pp. 95 - 102.","Dell, J. & How, R. A. (1984) Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 2: Widgiemooltha - Zanthus Study area. Records of the Western Australian Museum, 18 (Supplement), pp. 57 - 89.","Chapman, A. & Dell, J. (1985) Biology and zoogeography of the amphibians and reptiles of the Western Australian Wheatbelt. Records of the Western Australian Museum, 12, 1 - 46.","Smith, G. T., Leone, J. & Dickman, C. R. (1997) Small terrestrial vertebrate communities in remnant vegetation in the central wheatbelt of Western Australia. The Western Australian Naturalist, 21, 235 - 249.","Bush, B., Maryan, B., Browne-Cooper & Robinson, D. (2007) Reptiles and Frogs in the bush: Southwestern Australia. University of Western Australia Press, Perth, 302 pp.","Storr, G. M. & Hanlon, T. M. S. (1980) Herpetofauna of the Exmouth region, Western Australia. Records of the Western Australian Museum, 8, 423 - 439."]}

International audience; The worm-like snakes (Scolecophidia; approximately 400 nominal extant species) have a conservative morphology and are among the most poorly-known terrestrial vertebrates. Although molecular evidence has helped determine their higher-level relationships, such data have rarely been used to discriminate among species. We generated a molecular data set for the continental Australian blindsnakes (genus Ramphotyphlops) to determine the concordance of molecular and morphological information in the taxonomic recognition of species. Our dataset included 741 specimens morphologically attributed to 27 nominal Ramphotyphlops species. We proposed species hypotheses (SHs) after analysis of sequences from a variable mitochondrial gene (cytochrome b) and examined these SHs with additional evidence from a nuclear gene (prolactin receptor) and geographical data. Although the nuclear marker was not as fast-evolving and discriminating as the mitochondrial marker, there was congruence among the mitochondrial, nuclear, and geographical data, suggesting that the actual number of species is at least two times the current number of recognized, nominal species. Several biogeographical barriers and complex phytogeographical and geological patterns appeared to be involved in the division of some burrowing snake populations and, by consequence, in their diversification and speciation through isolation.

The Am Islands lie near the edge ot the Australian continental shelf in the Arafura Sea, approximately 150 km south of the coast of Papua (formerly Irian Jaya). For at least the first 40,000 years of occupation of Sahul they formed part of a continuous land bridge linking Australia and New Guinea. During this time they would have been a dissected limestone plateau on the exposed Carpentarian Plain. About 14,000 years BP sea level rose and began to encircle the island group, separating it from Australia and by 11,500 years BP it was completely separated from New Guinea. The presence on Am of numerous marsupials, the cassowary and Birds of Paradise attest to this shared history, a fact first recognised by Darwin’s co-discoverer of the theory of evolution by natural selection, the naturalist Alfred Russel Wallace (Wallace 1869)., Series: Advances in Geoecology (34). Chapter 14, pages 279--306.