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Smaug warreni (BOULENGER, 1908)

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Higher TaxaCordylidae (Cordylinae), Scincoidea, Sauria, Squamata (lizards)
Common NamesE: Lebombo Dragon Lizard, Warren’s Girdled Lizard 
SynonymZonurus warreni BOULENGER 1908
Cordylus warreni warreni — FITZSIMONS 1943
Cordylus warreni warreni — BOYCOTT 1992
Smaug warreni warreni — STANLEY et al. 2011
Cordylus warreni — PYRON & BURBRINK 2013
Smaug warreni — STANLEY & BATES 2014
Smaug warreni warreni — BATES et al. 2014: 210 
DistributionNE Republic of South Africa (from Ubombo Mountains in Zululand trough Transvaal Drakensberg, Soutpansberg), Swaziland, Botswana, Mozambique (up to Gorongoza Mountains)  
TypesHolotype: BMNH 1946.8.8.1 (Note:Boulenger's description states two individuals both male but only one registered and only one in the jar) 
DiagnosisDiagnosis. Distinguished from all other cordylids by its unique combination of dorsal, lateral and ventral colour patterns (see descriptions and figures). A medium to large species of Smaug distinguishable by the following combination of characters: (1) back usually sandy brown with 5–6 bands (usually interrupted) between fore- and hindlimbs, each band consisting of pale, dark-edged markings; (2) pale band on nape behind occipitals; (3) flanks pale with brown markings; (4) belly with brown (often pale) markings on most scales; (5) throat usually mostly pale with scattered small brown spots; (6) usually six enlarged, moderately to non-spinose, occipital scales, middle pair the smallest, outer occipitals usually equal in length to the adjacent inner ones; an additional small median occipital is often present; (7) dorsolateral and lateral scales weakly spinose; (8) tail moderately spikey; (9) dorsal scale rows transversely 31–41; (10) dorsal scale rows longitudinally 22–28; (11) ventral scale rows transversely 23–27; (12) ventral scale rows longitudinally 14 (rarely 12 or 13); (13) femoral pores per thigh 7–13; subdigital lamellae on 4th toe 15–20. It differs from other species of Smaug as described above in the diagnosis of S. swazicus sp. nov. (but maximum SVL in S. warreni is 141 mm, and femoral pores in males are 7–13). Most similar to S. swazicus sp. nov. and S. barbertonensis but easily distinguished by its colour pattern (see comparisons in diagnosis of S. swazicus sp. nov. above); by usually having shorter and blunter scales at the edges of the ear openings compared to S. barbertonensis; and quadrates without a pronounced ridge and concave region at the lateral edge of the adductor musculus mandibulae posterior origin (with a pronounced ridge and concave region in S. swazicus sp. nov.). Also differs as follows: outer occipitals and scales adjacent to them of about equal length (outer occipitals usually shorter than the adjacent inner ones in the other two species); head narrower than S. barbertonensis (head width/head length = 73–83% vs. 80–92% in adults); generally higher numbers of transverse dorsal scale rows (32‒38 in 92% of specimens) than S. barbertonensis (29‒32 in 81%); and greater numbers of longitudinal dorsal scale rows (22‒28, mean 23.6) than S. barbertonensis (20‒24, mean 21.7) [from Bates & Stanley 2020: 29] 
CommentDistribution: see map in STANLEY & BATES 2014: Figure 2. Not listed by AUERBACH 1987.

Subspecies: Cordylus warreni mossambicus, breyeri, regius, and vandami have been elevated to full species status and to a new genus, Smaug. Niels Jacobsen, in is unpublished 1989 thesis, wrote that the diagnostic characters of laevigatus were clinal with repect to eastern depressus, and that it should be synonymized [Ed Stanley, pers. comm., 31 Oct 2011]. 
EtymologyNamed after Mr. E. Warren. 
  • Auerbach, R.D. 1987. The Amphibians and Reptiles of Botswana. Mokwepa Consultants, Botswana, 295 pp.
  • Bates MF, Stanley EL. 2020. A taxonomic revision of the south-eastern dragon lizards of the Smaug warreni (Boulenger) species complex in southern Africa, with the description of a new species (Squamata: Cordylidae). PeerJ 8:e8526 - get paper here
  • Bates, M.F.; Branch, W.R., Bauer, A.M.; Burger, M., Marais, J.; Alexander, G.J. & de Villliers, M.S. (eds.) 2014. Atlas and Red List of the Reptiles of South Africa, Lesotho, and Swaziland. Suricata 1. South African National Biodiversity Institute, Pretoria, 512 pp.
  • Beolens, Bo; Michael Watkins, and Michael Grayson 2011. The Eponym Dictionary of Reptiles. Johns Hopkins University Press, Baltimore, USA - get paper here
  • Berghof, H.-P. 2006. Erfahrungen bei der Pflege und Vermehrung von Cordylus warreni depressus (FITZSIMONS 1930). Sauria 28 (2): 5-13 - get paper here
  • BERGHOF, H.-P.: 2011. Bemerkungen zur Geschlechtsreife von Cordyliden am Beispiel von Ouroborus cataphractus, Smaug mossambicus und Smaug warreni depressus. Sauria 33 (4): 41-46 - get paper here
  • Berghof, H.P. 2007. Cordylus mossambicus FITZSIMONS 1958 - Ein ausdauernder und interessanter Pflegling. Reptilia (Münster) 12 (65): 32-39 - get paper here
  • Boulenger, G. A. 1908. On a collection of fresh-water fishes, batrachians, and reptiles from Natal and Zululand, with description of new species. Ann Natal Gov. Mus. (Pietermaritzburg) 1(3): 219-235
  • Boycott, R.C. 1992. An Annotated Checklist of the Amphibians and Reptiles of Swaziland. The Conservation Trust of Swaziland - get paper here
  • Branch, W. R. 1998. Field Guide to the Snakes and Other Reptiles of Southern Africa. 3rd ed. Fully Revised and Updated to Include 83 New Species. Ralph Curtis Books (Sanibel Island, Florida), 399 pp.
  • Branch, William R. 1993. A Photographic Guide to Snakes and Other Reptiles of Southern Africa. Cape Town: Struik Publishers, 144 S.
  • Broadley, D.G. 1962. Cordylus warreni, a new addition to the southern Rhodesian list, with some remarks on other rupicolous lizards. Journal of the Herpetological Association of Rhodesia, 17/18: 5-8. - get paper here
  • Broadley, D.G. 1962. On some reptile collections from the North-Western and North-Eastern Districts of Southern Rhodesia 1958-1961, with descriptions of four new lizards. Occ. Pap. Nat. Mus. South. Rhodesia 26 (B): 787-843
  • Dam, G.P.F. van 1921. Descriptions of new species of Zonurus, and notes on the species of Zonurus occurring in the Transvaal. Annals Transvaal Mus. 7 (4): 239-243 - get paper here
  • Dathe, Falk 1988. AT Terrarientierlexikon, Cordylus warreni depressus (Fitzsimons, 1930), Zoutpansberg-Gürtelschweif (Familie: Cordylidae, Gürtel- und Schildechsen). Aquarien und Terrarien (Leipzig), 35 (10): 359.
  • FitzSimons, V. F. 1930. Descriptions of new South African Reptilia and Batrachia, with distribution records of allied species in the Transvaal Museum collection. Annals Transvaal Mus. 14: 20-48. - get paper here
  • FitzSimons, VIVIAN 1933. Description of five new lizards from the Transvaal and Southern Rhodesia. Annals Transvaal Mus. 15 (2): 273-280. - get paper here
  • FitzSimons, VIVIAN 1958. A new Cordylus from Gorongoza, Mocambique. Ann. Natal Mus. (Pietermaritzburg) 14 (2): 351-353
  • Kirchhof, S., M. Krämer, J. Linden & K. Richter 2010. The reptile species assemblage of the Soutpansberg (Limpopo Province, South Africa) and its characteristics. Salamandra 46 (3): 147-166 - get paper here
  • Krabbe-Paulduro, Ursula & Paulduro, Ernst 1989. Cordylus warreni (Boulenger). Sauria 11 (4) Suppl.: 153-160. - get paper here
  • Mouton, le Fras; Alexander Flemming; Michael Bates and Chris Broeckhoven 2018. The relationship between generation gland morphology and armour in Dragon Lizards (Smaug): a reassessment of ancestral states for the Cordylidae. Amphibia-Reptilia 39 (4): 445–456 - get paper here
  • MOUTON, P. LE FRAS N.; DAHNÉ A. JANSE VAN RENSBURG and JOHANNES H. VAN WYK 2010. Epidermal glands in cordylid lizards, with special reference to generation glands. Zoological Journal of the Linnean Society 158: 312–324. - get paper here
  • Pyron, R. Alexander; Frank T. Burbrink 2013. Early origin of viviparity and multiple reversions to oviparity in squamate reptiles. Ecology Letters 17 (1): 13–21 (published online 2013, in print 2014), DOI: 10.1111/ele.12168 - get paper here
  • Reissig, J. 2014. Girdled Lizards and Their Relatives. Edition Chimaira, Frankfurt, ISBN-10: 3899734378
  • Stanley, E. L. and Bates, M. F. 2014. Here be dragons: a phylogenetic and biogeographical study of the Smaug warreni species complex (Squamata: Cordylidae) in southern Africa. Zoological Journal of the Linnean Society 172 (4): 892–909; doi: 10.1111/zoj.12187 - get paper here
  • Stanley, Edward L.; Aaron M. Bauer; Todd R. Jackman, William R. Branch, P. Le Fras N. Mouton 2011. Between a rock and a hard polytomy: rapid radiation in the rupicolous girdled lizard (Squamata: Cordylidae). Molecular Phylogenetics and Evolution, 58(1): 53-70. - get paper here
  • Switak, Karl H. 1995. Girdle-tailed lizards of the genus Cordylus. Reptiles 3 (6): 8-24
  • Switak, Karl H. 1995. Die Gürtelschweife der Gattung Cordylus (2). TI-Magazin, Nr. 122 (März 1995), 39-42.v
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