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Smaug barbertonensis (VAN DAM, 1921)

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Higher TaxaCordylidae (Cordylinae), Scincoidea, Sauria, Squamata (lizards)
Common NamesE: Barberton Dragon Lizard, Barberton Girdled Lizard 
SynonymZonurus barbertonensis VAN DAM 1921
Zonurus barbertonensis barbertonensis — FITZSIMONS 1933 (by implication)
Cordylus warreni barbertonensis — FITZSIMONS 1943
Cordylus warreni barbertonensis — BOYCOTT 1992
Smaug warreni barbertonensis — STANLEY et al. 2011
Smaug warreni barbertonensis — BATES et al. 2014: 211
Smaug warreni barbertonensis — REISSIG 2014
Smaug barbertonensis — STANLEY & BATES 2014 
DistributionSouth Africa (Barberton District) and Swaziland (middleveld)

Type locality: Barberton [Mpumalanga Province], South Africa.  
TypesHolotype: DNMNH (= TM) 4273 
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 dark brown to black with 4–5 bands (usually interrupted) between fore- and hindlimbs, each band consisting of pale, sometimes dark-edged, markings;
(2) pale spot or blotch on nape behind occipitals; (3) flanks dark with narrow, pale, vertical markings; (4) belly mostly dark brown or black; (5) throat almost entirely dark brown or black with only a few pale areas; (6) usually six enlarged, moderately to non-spinose, occipital scales, middle pair the smallest, outer occipitals usually shorter than the adjacent inner ones; (7) dorsolateral and lateral scales moderately spinose; (8) tail moderately spikey; (9) dorsal scale rows transversely 28–34; (10) dorsal scale rows longitudinally 20–24; (11) ventral scale rows transversely 25–28; (12) ventral scale rows longitudinally 14 (rarely 16); (13) femoral pores per thigh 8–12; subdigital lamellae on 4th toe 15–19.
It differs from other species of Smaug as described above in the diagnosis of S. swazicus sp. nov. (but maximum SVL in S. barbertonensis is 140 mm, and femoral pores in males are 8–11 per thigh).
Most similar to S. swazicus sp. nov. and S. warreni, but easily distinguishable by its colour pattern (see comparisons in diagnosis of S. swazicus sp. nov. above); by usually having more elongate and spinose scales at the edges of the ear openings (shorter and non-spinose in S. swazicus sp. nov. and S. warreni); and quadrates lacking 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 shorter than the adjacent inner ones (of about equal length in S. warreni); head relatively wider than the other two species (head width/head length = 80–92% vs. 73–84% in adults); lower numbers of transverse dorsal scale rows (29‒32 in 81%) compared to S. swazicus sp. nov. (32‒37 in 86%) and S. warreni (32‒38 in 92%); and lower numbers of longitudinal dorsal scale rows (20‒24, mean 21.7) compared to S. warreni (22‒28, mean 23.6) [from Bates & Stanley 2020: 26]. 
CommentSee also S. warreni.

Distribution: see map in STANLEY & BATES 2014: Figure 2. 
EtymologyNamed after the type locality. 
  • 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.
  • Boycott, R.C. 1992. An Annotated Checklist of the Amphibians and Reptiles of Swaziland. The Conservation Trust of Swaziland - get paper here
  • 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
  • FitzSimons, V.F. 1943. The lizards of South Africa. Transvaal Museum Memoir No.1 (Pretoria), 528 pp.
  • 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
  • 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
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