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Smaug swazicus BATES & STANLEY, 2020

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Higher TaxaCordylidae (Cordylinae), Scincoidea, Sauria, Squamata (lizards)
Subspecies 
Common NamesE: Swazi Dragon Lizard
G: Swazi-Drachenechse 
SynonymSmaug swazicus BATES & STANLEY 2020
Cordylus warreni barbertonensis — FITZSIMONS 1943: 426 (part)
Cordylus warreni barbertonensis — BRANCH 1988: 164 (part)
Cordylus warreni barbertonensis — BRANCH 1998: 195 (part)
Cordylus warreni barbertonensis — JACOBSEN 1989 (part)
Cordylus warreni barbertonensis — ADOLPHS 1996: 15 (part)
Cordylus warreni barbertonensis — BOURQUIN 2004: 96
Cordylus warreni barbertonensis — ADOLPHS 2006: 22 (part).
Smaug warreni barbertonensis — STANLEY et al. 2011: 64 (part)
Smaug warreni barbertonensis — BATES et al. 2014: 211 (part)
Smaug warreni barbertonensis — REISSIG 2014: 190ff (part).
Smaug sp. — STANLEY & BATES, 2014: 905
Smaug barbertonensis — MOUTON et al. 2018: 460, fig. 2g
Smaug cf. barbertonenis — MOUTON et al. 2018: 464 
DistributionRepublic of South Africa (Mpumalanga), KwaZulu–Natal, elevations of 462 - 1,139 m.

Type locality: 320 m SSE of car park, Maguga Dam, Hhohho Region, Eswatini (26°04′57′′S, 31°15′59′′E; 2631AB; 635 m a.s.l.),  
Reproduction 
TypesHolotype. NMB R9201 (Figs. 5–7; sample from this specimen was used in molecular analysis by Stanley & Bates, 2014), adult male (differentiated glandular femoral scales present; mid-ventral incision present) from collected by E.L. Stanley & J.M. da Silva, 31 October 2008.
Paratypes. (note copied coordinates all lack degree symbols due to formatting in original paper). Allotype: TM 78918 (Figs. 2B and 2E), adult female (no differentiated femoral scales) from Nkomati Gorge, Malolotja Nature Reserve, Hhohho Region, Eswatini (2603′15′′S, 3108′06′′E; 2631AA; 640 m a.s.l.), collected by R.C. Boycott, 29 August 1993. Ten more paratypes: TM 83000, adult male, 1 km NW of Maguga Dam, Hhohho Region, Eswatini (2604′04′′S, 3114′55′′E; 2631AA; 618 m a.s.l.), R.C. Boycott, 25 March 1997; TM 83532, adult male, 5 km SE of Bhunya, Manzini Region, Eswatini (2632′16′′S, 3102′54′′E; 2631CA; 960 m a.s.l.), R.C. Boycott, 28 June 2000; TM 42531, adult female, Mbutini Hills, 23 km N of Sepofaneni, Manzini Region, Eswatini (2631′34′′S, 3135′45′′E; 2631DA), W.D. Haacke, 3 September 1972; TM 51376, adult male, 15 km NW of Gilgal on route to Manzini, Manzini Region, Eswatini (2631DA), W.D. Haacke, 3 September 1972; TM 78931, juvenile, Nkomati Gorge, Malolotja Nature Reserve, Hhohho Region, Eswatini (2603′14′′S, 3108′02′′E; 2631AA; 669 m a.s.l.), R.C. Boycott, 14 September 1993; TM 78921, juvenile, Nkomati Valley, Hhohho Region, Eswatini (2603′12′′S, 3114′24′′E; 2631AA; 580 m a.s.l.), J. Linden, 31 October 1992; NMB R9194, adult male, Nkomati Viewpoint, Malolotja Nature Reserve, Hhohho Region, Eswatini (2604′29′′S, 3107′32′′E; 2631AA; 1,033 m a.s.l.), E.L. Stanley & J.M. da Silva, 31 October 2008 (Fig. 8B); NMB R9195, adult male, Nkomati Viewpoint, Malolotja Nature Reserve, Hhohho Region, Eswatini (2604′55′′S, 3108′03′′E; 2631AA; 1,139 m a.s.l.), E.L. Stanley & J.M. da Silva, 31 October 2008; NMB R9202 (mid-ventral incision present; sample from this specimen was used in molecular analysis by Stanley & Bates, 2014), adult male from 230 m SSE of car park, Maguga Dam, Hhohho Region, Eswatini (2604′54′′S, 3115′58′′E; 2631AB; 640 m a.s.l.), collected by E.L. Stanley & J.M. da Silva, 31 October 2008; TM 73290, adult female, Nzulase, Mpumalanga Province, South Africa (2551′S, 3138′E; 2531DC), N.H.G. Jacobsen, 29 March 1983. 
DiagnosisDiagnosis. Distinguished from all other cordylids (Cordylidae) by its unique combination of dorsal, lateral and ventral colour patterns (see descriptions and figures). Referable to the genus Smaug on the basis of its large size and robust body, enlarged and spinose dorsal and caudal scales, enlarged occipital scales, and frontonasal in contact with the rostral, separating the nasal scales. A medium to large species of Smaug distinguishable by the following combination of characters: (1) back dark brown usually with 5–6 pale bands (usually interrupted) between fore- and hindlimbs, each band consisting of pale, sometimes dark-edged, markings; (2) pale band on nape behind occipitals; (3) flanks with large pale spots or blotches; (4) belly pale with a dark median longitudinal band bordered on either side by broad, dark, bands; (5) throat pale with extensive bold brown mottling (sometimes forming transverse bands; often much of throat is dark); (6) 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 31–41; (10) dorsal scale rows longitudinally 20–26; (11) ventral scale rows transversely 23–29; (12) ventral scale rows longitudinally 14 (rarely 12); (13) femoral pores per thigh 10–13; subdigital lamellae on 4th toe 16–19. Its status as a new species is also supported by monophyly with high levels of support from three mitochondrial and eight nuclear markers (see Stanley & Bates, 2014; using samples from NMB R9201–2). It differs from the terrestrial S. giganteus by its smaller adult size (maximum SVL 145 mm vs. 198 mm), and possession of six moderate sized and weakly spinose occipitals, vs. four (occasionally five) large and distinctly spinose occipitals. Differs from other species of Smaug as follows: from S. vandami by having six (versus usually four) occipitals; from S. depressus by having only 10‒13 (vs. 16‒24) femoral pores per thigh in males; and from S. breyeri by having much less rugose head shields. It differs from S. giganteus, S. breyeri and S. vandami by having less spinose occipitals and tail spines, and two-layer (rather than multi-layer) generation glands. Differs from S. mossambicus and S. regius by having the first supralabial with moderate or no (vs. distinct) upward prolongation and lacking obvious sexual dichromatism (only males of the latter two species have bright yellow to orange flanks). Most similar to S. barbertonensis and S. warreni, but easily distinguishable by its colour pattern (as described above) compared to S. barbertonensis (back dark brown with 4–5 pale bands between the limbs, pale spot or blotch on nape behind occipitals; flanks dark with narrow pale vertical markings; venter mostly dark brown or black) and S. warreni (back usually pale brown with 5–6 pale dark-edged bands between the limbs, pale band on nape behind occipitals; flanks pale with brown markings; venter with brown markings on most scales) (Figs. 2, 6 and 8, and others below); by usually having short, blunt, non-spinose scales at the edges of the ear openings (usually elongate and spinose in S. barbertonensis); and quadrates with a pronounced ridge and concave region at the lateral edge of the adductor musculus mandibulae posterior origin (no pronounced ridge or concave region in the other two species). Also differs as follows: outer occipitals usually shorter than the adjacent inner ones (of about equal length in S. warreni); head narrower than S. barbertonensis (head width/head length = 76–84% vs. 80–92% in adults); generally higher numbers of transverse dorsal scale rows (32‒37 in 86% of specimens) than S. barbertonensis (29‒32 in 81% of S. barbertonensis). 
CommentHabitat: rupicolous, living in deep, horizontal (or gently sloping) crevices in granitic rock along hillsides, usually in the partial shade of trees. 
EtymologyNamed for the Kingdom of Eswatini, the country where most of the species’ range is located. Both ‘eSwatini’ and ‘Swaziland’ derive from the word iSwazi, after the name of an early chief, Mswati II (c. 1820–1868). 
References
  • Adolphs, K. 1996. Bibliographie der Gürtelechsen und Schildechsen. Squamata Verlag, Sankt Augustin 1996: 1-255
  • Adolphs, K. 2006. Bibliotheca Cordyliformium. Squamata Verlag, Sankt Augustin, 303 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.
  • Bourquin, O. 2009. Reptiles (Reptilia) in KwaZulu-Natal: I - diversity and distribution. Durban Museum Novitates 29: 57-103
  • 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,W.R. 1988. Field Guide to the snakes and other reptiles of southern Africa. Struik Publishers, Cape Town, 328 pp.
  • FitzSimons, V.F. 1943. The lizards of South Africa. Transvaal Museum Memoir No.1 (Pretoria), 528 pp.
  • Jacobsen, N. H. G. 1989. A herpetological survey of the Transvaal. Unpubl. thesis, Univ. Natal, South Africa
  • 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
  • Reissig, J. 2020. Ein neuer Drache erwacht aus dem Schatten: die Swazi-Drachenechse. Elaphe 2020 (4): 64-65
  • 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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