[Herpetology • 2023] Cordylus momboloensis • A New Species of Green-eyed Cordylus Laurenti, 1768 (Squamata: Cordylidae) from the west-central Highlands of Angola, and the Rediscovery of Cordylus angolensis (Bocage, 1895)

 

Cordylus momboloensis 

Bates, Lobón-Rovira, Stanley, Branch & Vaz Pinto, 2023

DOI: 10.3897/vz.73.e95639

Abstract

Cordylus angolensis (Bocage, 1895) was described 128 years ago on the basis of a single specimen collected at Caconda in the west-central highlands of Angola. Additional specimens referred to this species were collected at ‘Mombolo’ (also in the central highlands) during the Vernay Angola Expedition in 1925. As the holotype was apparently destroyed in the fire of 1978 at the Museu Bocage in Lisbon and no additional specimens have been collected, its taxonomic status and phylogenetic relationships has remained uncertain. The species has eluded all efforts aimed at its re-discovery in the vicinity of the type locality, with a single specimen from near Condé, north of Mombolo—collected in 1970 by Wulf Haacke—the only other specimen of Cordylus known from west-central Angola. Recent field work in the Angolan highlands resulted in the collection of a series of specimens from Taqueta Mountain (west of Caconda), Monte Verde (Sandula, ‘Mombolo’) and Uassamba (Vondo). A phylogenetic analysis, using three mitochondrial and six nuclear genes, indicated the existence of two distinct species-level lineages in the Angolan highlands. These two species are allopatric and morphologically distinct, differing especially in terms of their colour patterns, eye colour and certain scalation characteristics. We therefore confirm that C. angolensis is a valid species and designate a neotype, and describe a new species, Cordylus momboloensis sp. nov.

Keywords: Africa, Cordyliformes, distribution, Sauria, taxonomy

Cordylus angolensis: live neotype (A) and habitat at the collecting site of Taqueta Mountain, Angola (B);
and Cordylus momboloensis sp. nov.: live holotype (C) and habitat at the type locality of Sandula, Angola (D).

Photos: Pedro Vaz Pinto (A, B), and William R. Branch (C, D).

Holotype (PEM R25217) of Cordylus momboloensis sp. nov. from Sandula (Mombolo), Cuanza Sul Province, Angola.
A dorsal and ventral aspects, B lateral (right), dorsal and ventral aspects of the head, C underside of the foot, D femoral region of the legs. Although not evident because of the way the holotype was photographed, its tail is intact.

Photos: Pedro Vaz Pinto.

Cordylus momboloensis sp. nov.

 Mombolo girdled lizard; Lagarto-espinhoso do Mombolo

Diagnosis: 

(Where variation in additional material falls outside the range of type material, this is indicated in parentheses.) A medium to large rupicolous Cordylus with a moderately depressed head and body. Referred to Cordylus (rather than any other species of Cordylidae) by the following combination of characters: head distinct from body, two pairs of large and well developed limbs (body serpentiform, head indistinct from body, and limbs rudimentary in Chamaesaura), scales on back large and keeled (granular in Platysaurus, partly granular in Pseudocordylus and Hemicordylus), non-spinose occipitals [and post-occipitals] (occipitals spinose in Smaug), 23–24 transverse dorsal scale rows (40–43 in Ninurta; 31–46 in Karusasaurus; 15–16 in Ouroborus); loreal absent (present in Smaug, Ninurta, Ouroborus, Karusasaurus, and Namazonurus).

Cordylus momboloensis sp. nov. is distinguishable from other members of its genus by the following combination of characters: (1) back dark brown with a paravertebral series of pale markings; (2) top of head with pale blotches; (3) iris of the eye mostly pale green (blue-green to some eyes), with a brown ring around the pupil; (4) scales of the first transverse row of dorsals similar in appearance to those of the row behind; (5) loreal shield absent; (6) nostril pierced in the posterior part of a large nasal, situated behind the suture of rostral and first supralabial, always well separated from the first supralabial, and usually separated from the preocular; (7) an interrupted row of non-spinose occipitals consisting of 2–3 scales on either side of head; (8) a row of 5–6 non-spinose post-occipitals, the median scales of which are in contact with the posterior parietals and separate the occipitals; (9) Frontonasal separated from the frontal by a pair of prefrontals, each of which exceeds it in size; (10) Anterior pair of parietals usually in contact anteriorly; (11) dorsolateral and lateral scales weakly to moderately spinose; (12) tail spinose, more weakly so distally; (13) dorsal scale rows transversely 22–24; (14) dorsal scale rows longitudinally 18–23; (15) ventral scale rows transversely 22–27; (16) ventral scale rows longitudinally 12–13 [rarely 11 or 14]; (17) subdigital lamellae on 4th toe 11–15; (18) femoral pores per thigh 5–8 in males, 4–6 in females [rarely 0; 7 on one side in a non-type specimen, TM 46476]; (19) differentiated femoral scales [generation glands] per thigh in males 25–37; (20) premaxillary teeth 9.

Its status as a new species is supported by monophyly with high levels of support from a suite of three mitochondrial and six nuclear markers (see above); and it differs from C. angolensis, the most similar species genetically and morphologically, by an uncorrelated ND2 p-distance of 9.22% (Table 3).

Lateral views of the head in Angolan Cordylus.
 Cordylus angolensis—A CHL 611, B CHL 615, C MNCN 50649, D CHL 613, E MNCN 50648 (neotype);
Cordylus momboloensis sp. nov. —F MNCN 50760 (paratype), G FKH 0125 (paratype), H FKH 0127 (paratype), I FKH 0128 (paratype), J FKH 0129, K PEM R25217 (holotype), L MNCN 50761 (allotype);
C. namakuiyus—M PEM R25218, N CHL 616, O CHL 609;
C. phonolithos—P CHL 852, Q CHL 1056, R MNCN 50651, S CHL 851, T CHL 1057, U MNCN 50652, V CHL 848;
and C. machadoi—W CHL 701, X CHL 700, Y PEM R19784.

Photos: Pedro Vaz Pinto (A–N, P–V), Timóteo Júlio (O, W–X), and William R. Branch (Y).

   

Live specimens of various species of Cordylus:
 A C. machadoi (PEM R25218 – 6 km north of Humpata, Huíla Province, Angola; photo: William R. Branch); B C. namakuiyus (PEM R18005 – between Namibe and Omahua lodge, Namibe Province, Angola; photo: William R. Branch); C C. phonolithos (CAS 263581, holotype – vicinity of N’Dolondolo, Serra da Neve, Namibe Province, Angola; photo: Luis M. P. Ceríaco); D C. rhodesianus (near Chimoio, Mozambique; photo: David Maguire);
E C. tropidosternum (Kayalekera, Malawi; photo: Luke Verburgt); F C. jonesii (Umbabat Private Nature Reserve, Mpumalanga Province, South Africa; photo: Darren Pietersen); G C. vittifer (NMB R8523 – Farm: Dipka 220, Vrede district, Free State Province, South Africa; photo: Michael F. Bates); H C. cordylus (NMB R8540 – Ha Sehlaba, Lesotho; photo: Michael F. Bates). Representatives of all four Cordylus clades recovered by the current genetic analysis are included.

 

Michael F. Bates, Javier Lobón-Rovira, Edward L. Stanley, William R. Branch and Pedro Vaz Pinto. 2023. A New Species of Green-eyed Cordylus Laurenti, 1768 from the west-central Highlands of Angola, and the Rediscovery of Cordylus angolensis (Bocage, 1895) (Squamata: Cordylidae). Vertebrate Zoology. 73: 599-646. DOI: 10.3897/vz.73.e95639

[Herpetology • 2020] A Taxonomic Revision of the South-eastern Dragon Lizards of the Smaug warreni (Boulenger) Species Complex (Squamata: Cordylidae) in southern Africa, with the Description of A New Species, Smaug swazicus

Smaug swazicus Bates​​ & Stanley, 2020

Swazi Dragon Lizard  || DOI: 10.7717/peerj.8526  

 twitter.com/DrScanley 
FloridaMuseum.ufl.edu

Abstract

A recent multilocus molecular phylogeny of the large dragon lizards of the genus Smaug Stanley et al. (2011) recovered a south-eastern clade of two relatively lightly-armoured, geographically-proximate species (Smaug warreni (Boulenger, 1908) and S. barbertonensis (Van Dam, 1921)). Unexpectedly, S. barbertonensis was found to be paraphyletic, with individuals sampled from northern Eswatini (formerly Swaziland) being more closely related to S. warreni than to S. barbertonensis from the type locality of Barberton in Mpumalanga Province, South Africa. Examination of voucher specimens used for the molecular analysis, as well as most other available museum material of the three lineages, indicated that the ‘Eswatini’ lineage—including populations in a small area on the northern Eswatini–Mpumalanga border, and northern KwaZulu–Natal Province in South Africa—was readily distinguishable from S. barbertonensis sensu stricto (and S. warreni) by its unique dorsal, lateral and ventral colour patterns. In order to further assess the taxonomic status of the three populations, a detailed morphological analysis was conducted. Multivariate analyses of scale counts and body dimensions indicated that the ‘Eswatini’ lineage and S. warreni were most similar. In particular, S. barbertonensis differed from the other two lineages by its generally lower numbers of transverse rows of dorsal scales, and a relatively wider head. High resolution Computed Tomography also revealed differences in cranial osteology between specimens from the three lineages. The ‘Eswatini’ lineage is described here as a new species, Smaug swazicus sp. nov., representing the ninth known species of dragon lizard. The new species appears to be near-endemic to Eswatini, with about 90% of its range located there. Our study indicates that S. barbertonensis sensu stricto is therefore a South African endemic restricted to an altitudinal band of about 300 m in the Barberton–Nelspruit–Khandizwe area of eastern Mpumalanga Province, while S. warreni is endemic to the narrow Lebombo Mountain range of South Africa, Eswatini and Mozambique. We present a detailed distribution map for the three species, and a revised diagnostic key to the genus Smaug.

 Smaug swazicus sp. nov. (B) Dorsal colouration of live paratype (NMB R9194). (C) Ventral colouration.

(Photos: E.L. Stanley).

Figure 8: (A) Shady rocky outcrops at Nkomati Viewpoint, Malolotja National Park, Eswatini, typical habitat of Smaug swazicus sp. nov. (B) Dorsal colouration of live paratype of S. swazicus (NMB R9194). (C) Ventral colouration of S. swazicus. (Photo credits: E.L. Stanley).

Systematics

Family Cordylidae Gray, 1838

Smaug swazicus Bates & Stanley sp. nov.

Swazi Dragon Lizard

Diagnosis. (includes ‘additional material’) 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.
....

Figure 5: Smaug swazicus sp. nov. (A) Dorsal, (B) ventral and (C) lateral views of the head of the holotype (NMB R9201). (Drawing credit: E.L. Stanley).

   

Figure 2: Differences in colour pattern in the Smaug warreni species complex.
From left to right: Dorsal views of (A) Smaug barbertonensis (NMB R9196, topotype), (B) S. cf. barbertonensis (TM 78918, allotype of new species) and (C) S. warreni (TM 63567);
ventral views of (D) S. barbertonensis (TM 55789), (E) S. cf. barbertonensis (TM 78918) and (F) S. warreni (TM 78969)

(Photos: M.F. Bates).

   

Etymology. Named 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).

Distribution. Highveld and Middleveld of Eswatini in Hhohho, Manzini and Shiselweni Regions, and adjacent areas in the South African provinces of (eastern) Mpumalanga (in Nkomazi municipality) and (northern) KwaZulu–Natal (in uPhongolo and Abaqulusi municipalies) (Fig. 9) at elevations of 462 to 1,139 m a.s.l.

Natural history. Diurnal and rupicolous, living in deep, horizontal (or gently sloping) crevices in granitic rock along hillsides, usually in the partial shade of trees (Fig. 8A; see also Jacobsen, 1989). According to R.C. Boycott (in litt., 2019), rocky terrain in closed canopy bushveld is the preferred habitat in Eswatini. A specimen in Ithala Game Reserve in KwaZulu–Natal was photographed on a tree trunk (ReptileMAP, VM no. 152451). When grasped by the hind limb, an individual from the type series performed an unusual anti-predator behaviour by repeatedly flexing and extending the inhibited limb caudally, so as to pull the captors’ digits directly onto the very sharp whorl of spines at the base of the tail (E.L. Stanley, 2008, personal observation). 

    

Michael F. Bates​​ and Edward L. Stanley​. 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. DOI: 10.7717/peerj.8526  

 twitter.com/DrScanley/status/1242816810728259584

Here be dragons: Analysis reveals new species in Smaug lizard group
FloridaMuseum.ufl.edu/science/smaug-swazicus-new-dragon-lizard/
  sciencecover.com/3619-2-here-be-dragons-analysis-reveals-new-species-in-smaug-lizard-group via @ScienceCover

         

          

[Herpetology • 2019] Cordylus phonolithos • A New Species of Girdled Lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, southwestern Angola

Cordylus phonolithos 

 Marques, Ceríaco, Stanley, Bandeira, Agarwal & Bauer, 2019

Photo by Luis Ceríaco  facebook.com/LuisCeriaco

DOI: 10.11646/zootaxa.4668.4.4

Cordylus phonolithos 

 Marques, Ceríaco, Stanley, Bandeira, Agarwal & Bauer, 2019

N’Dolondolo Girdled Lizard • Lagarto Espinhoso de N’Dolondolo  || DOI: 10.11646/zootaxa.4668.4.4

photos: Luis Ceríaco facebook.com/LuisCeriaco

Abstract

A new species of rupicolous girdled lizard is described from Serra da Neve Inselberg, Namibe Province, southwestern Angola. We sequenced two mitochondrial gene regions (16S and ND2) for the new species and compared these data with those previously published for other Cordylus species. The new species is genetically divergent from the closely related Cordylus machadoi and C. namakuiyus and morphologically distinguished by head scale arrangement, coloration, and osteological characters. Serra da Neve is the second highest peak in Angola and one of the most understudied areas of the country. The discovery of this new species highlights the importance of this inselberg for regional endemism and emphasizes the high conservation importance of the area.

Keywords: Reptilia, Cordylus phonolithos sp. nov., molecular phylogeny, computed tomography, endemism, taxonomy, Africa

Cordylus phonolithos sp. nov. in life ([Holotype] CAS 263581). N’Dolondolo, Serra da Neve, southwestern Angola.

Photo by Luis Ceríaco

Cordylus phonolithos sp. nov. in life ([Paratype] INBAC: AMB 10272). N’Dolondolo, Serra da Neve, southwestern Angola.

Photo by Luis Ceríaco

Cordylus phonolithos sp. nov. 

C. namakuiyus [part]: Stanley et al. (2016: 209)

Etymology. The specific epithet “phonolithos” is a noun in apposition from the Greek “phono” = sound + “lithos” = rock, which means “sound stone”. In the local Mucobal language the type locality name, “N’Dolondolo” means literally “rock that sounds like a bell” or “bell” and stems from the presence of a large and famous phonolite stone at the locality. Phonolites are rare igneous volcanic stones of intermediate composition between felsic and maphic, with aphanitic to porphyritic texture that produce a very distinctive metallic sound when hit, similar to the ringing of a metallic bell. We suggest “N’Dolondolo Girdled Lizard” and “Lagarto Espinhoso de N’Dolondolo” as the English and Portuguese common names, respectively, for this species.

Mariana P. Marques, Luis M. P. Ceríaco, Edward L. Stanley, Suzana Bandeira, Ishan Agarwal and Aaron M. Bauer. 2019. A New Species of Girdled Lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, southwestern Angola. Zootaxa. 4668(4); 503–524. DOI: 10.11646/zootaxa.4668.4.4 

facebook.com/groups/1045499302182009/permalink/2477243312340927/

facebook.com/groups/HerpingTheGlobe/permalink/2334608893458976/

 Edward L. Stanley, Luis M. P. Ceríaco, Suzana Bandeira, Hilaria Valerio, Michael F. Bates and William R. Branch. 2016. A Review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the Description of A New Species from the Pro-Namib Desert. Zootaxa. 4061(3): 201–226.  DOI: 10.11646/zootaxa.4061.3.1

  

[Herpetology • 2016] Cordylus namakuiyus • A Review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the Description of A New Species from the Pro-Namib Desert

Cordylus namakuiyus  

Stanley, Ceríaco, Bandeira, Valerio, Bates & Branch, 2016

Kaokoveld Girdled Lizard || DOI: 10.11646/zootaxa.4061.3.1 

Abstract

The girdled lizard genus Cordylus is represented in Angola by two species, Cordylus angolensis and C. machadoi, separated from their nearest congeners by over 700 km. Here we describe a new species, Cordylus namakuiyus sp. nov., endemic to the arid lowlands west of the southern Angolan escarpment. Phylogenetic analysis using three mitochondrial and eight nuclear genes shows that the low-elevation forms and the proximate, high-elevation species C. machadoi are genetically divergent and reciprocally monophyletic, and together form the earliest diverging lineage of the northern Cordylus clade. Morphological data, collected using computed tomography and traditional techniques (scalation and morphology), identify consistent phenotypic differences between these high- and low-elevation species and allows for a detailed description of the osteology and osteodermal arrangements of the new species. A series of 50 specimens, collected during the 1925 Vernay expedition to southwestern Angola and housed at the American Museum of Natural History, are assigned to the new species, although the identity of Cordylus from northern Namibia remains ambiguous and requires further investigation.

Keywords: Reptilia, computed tomography, girdled lizard, Kaokoveld, molecular phylogeny, ontogeny, osteoderms, osteology

Etymology. The specific epithet “namakuiyus” is the masculine latinised form of namakuiya, which means “thorny” in the Herero language, referring to the sharp spines on the limbs and tail of this species. Suggested common name: Kaokoveld Girdled Lizard.

Distribution. Currently, the new species is known only from the south and central parts of Namibe Province, Angola, southwest of the Leba escarpment, at elevations of 215–509 m a.s.l.


Habitat and natural history notes. This species is found in gently sloping crevices of granite outcrops in the arid Kaokoveld. When approached, specimens retreat into their fissure as far as possible, wedging themselves head-first and protecting their head and flanks with their spiny tails. As with all other cordylines, Cordylus namakuiyus sp. nov. is viviparous; the holotype contained two large fetuses (SVL 49.5% of the holotype). The fetuses appear to be approaching full-term, suggesting a spring/early summer parturition period. Although these fetuses lack any body osteoderms, the slightly larger neonates (SVL 55.6–56.4% of the holotype) possess significant dermal ossification, suggesting that the young rapidly accumulate osteodermal armour following parturition. One of the Pico Azevedo specimens was found occupying the same rock crack as an adult Chondrodactylus pulitzerae (Schmidt), and PEM R18005 from Iona National Park inhabited the same crack as an adult Chondrodactylus fitzsimonsi (Loveridge).

 Edward L. Stanley, Luis M. P. Ceríaco, Suzana Bandeira, Hilaria Valerio, Michael F. Bates and William R. Branch. 2016. A Review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the Description of A New Species from the Pro-Namib Desert. Zootaxa. 4061(3): 201–226.  DOI: 10.11646/zootaxa.4061.3.1

https://indicator.ru/article/2016/12/24/novye-vidy-i-gde-oni-obitayut/

 radiomundial.com.ve/article/descubren-nueva-especie-de-lagarto-en-sur-de-angola

 calacademy.org/press/releases/133-new-species-described-by-the-california-academy-of-sciences-in-2016

[Herpetology • 2014] Here be Dragons: A Phylogenetic and Biogeographical Study of the Smaug warreni species complex (Squamata: Cordylidae) in southern Africa

Figure 2. Distribution ranges of taxa in the Smaug warreni species complex, taken from Jacobsen (1989) and Branch (1998) (S. breyeri, blue; S. vandami, red; S. w. depressus, purple; S. w. warreni, green; northern S. w. barbertonensis clade, yellow; southern S. w. barbertonensis clade, orange). Numbers in coloured circles represent localities of the specimens used in the molecular analyses of the Smaug warreni species complex as listed in Table 1. Dashed lines represent the boundaries of key mountain chains.

DOI: 10.1111/zoj.12187 

Taxonomy of the Smaug warreni species complex remains contentious despite known morphological differences and geographical separation of the various taxa. This study uses an 11-gene dataset to recover phylogenetic relationships between the seven nominal members of the S. warreni complex. Eight well-supported clades were returned, with S. warreni barbertonensis found to be paraphyletic. A time-calibrated analysis of molecular data indicates that all eight clades in the S. warreni complex separated in the late Miocene, much earlier than the date suggested by the existing hypothesis of vicariance through the ingression of Kalahari sands. Ecological niche modelling indicates that although all clades are allopatric, a slight decrease in temperature could potentially render them sympatric, supporting an hypothesis of range expansion through climatic change.

Keywords: biogeography; geography; palaeoecology; southern Africa; species delimitation; taxonomic revision

Figure 1. Variation in dorsal coloration in the Smaug warreni species complex in South Africa. From left to right: S. breyeri (NMB R9150, Geelhoutkop – type locality), S. vandami (NMB R9188, 4 km north of Lammerkop), S. w. depressus (NMB R9157, 10 km north-east of Louis Trichardt, Soutpansberg), S. w. barbertonensis (NMB R9192, Barberton – type locality), S. w. warreni (NMB R9292, Mount Mananga, Lebombo Range).

Photograph by M. F. Bates.   DOI: 10.1111/zoj.12187

Figure 7. Phenotypic variation in Smaug depressus: A, S. depressus (‘laevigatus’ morph) from the type locality of Entabeni; B, S. depressus (typical spotted morph) from 10 km north-east of Louis Trichardt, Limpopo Province, South Africa.

Photographs by E. L. Stanley.  DOI: 10.1111/zoj.12187

Discussion

Phylogenetic relationships and taxonomic implications

The eight major clades – representing seven currently recognized taxa and an un-named species currently assigned to S. w. barbertonensis – recovered by our analysis (Fig. 3) display high mitochondrial divergence (5.6–8.7%), possess non-identical ecological niches and exhibit diagnostic morphological characters. This confluence of diagnoses supports the working hypothesis that each represents an independently evolving metapopulation and can be treated as a valid species under the General Lineage Species Concept of de Queiroz (1998). No samples from the southern range of S. barbertonensis (see Bourquin, 2004; Bates et al., 2014) were included in this molecular analysis and further investigation is therefore required before we can establish whether S. barbertonensis from Swaziland and KwaZulu-Natal forms a monophyletic group or is rendered paraphyletic by S. warreni. However, as S. warreni barbertonensis is rendered paraphyletic by S. w. warreni and the populations of S. w. barbertonensis from Swaziland and KwaZulu-Natal are genetically and morphologically distinct from S. w. barbertonensis from the type locality (Barberton), these southern populations will be assigned to a new species (Bates & Stanley, in prep.). Furthermore, S. w. depressus and S. w. barbertonensis populations from Barberton northwards should be elevated to full species status to reflect the findings of this study and render S. warreni a monophyletic species. We therefore recognize the following species: Smaug warreni (Boulenger, 1908), Smaug breyeri (Van Dam, 1921), Smaug barbertonensis (Van Dam, 1921) comb. nov., Smaug depressus (FitzSimons, 1930) comb. nov., Smaug vandami (FitzSimons, 1930), Smaug mossambicus (FitzSimons, 1958) and Smaug regius (Broadley, 1962).

The three clades of S. breyeri returned by the analysis correspond to the southern (sites 1–3), western (sites 4 and 5) and north-eastern (site 6) sampling sites for this species (Fig. 2). Jacobsen (1989) recorded an isolated population of S. vandami from the Blouberg/Makgabeng mountains (vicinity of site 6, Fig. 2), and referred to specimens of both S. breyeri (museum voucher TM 58416) and S. vandami (TM 73979) from ‘farm New York’ in the eastern Waterberg mountains. He assigned species status to S. vandami on the basis of this apparent sympatry. A subadult S. breyeri (NMB R9134) collected 10 km south of Lephalale (site 1 in Fig. 2) – close to Jacobsen's record of sympatry – was similar in coloration to typical S. vandami (Fig. 1), and the Makgabeng individual (MBUR 00320) used in this study was initially identified as S. vandami by the collector based on colour pattern. The molecular analysis recovers both of the latter samples in the S. breyeri clade, and their identity was confirmed by subsequent morphological examination of voucher specimens. The two specimens (farm New York) on which Jacobsen (1989) based his comments regarding sympatry of ‘S. w. breyeri’ and S. vandami (the latter taxon elevated to a full species on the basis of his assessment) have also been examined and both are here considered referable to S. breyeri. Therefore, the above-mentioned Blouberg/Makgabeng and farm New York records, rather than representing isolated populations of S. vandami (see map in Bates et al., 2014), are associated with the known range of S. breyeri.

FitzSimons (1933) proposed that S. depressus from Entabeni (type locality) in the Soutpansberg range was a separate species, namely Zonurus laevigatus. Jacobsen (1989) suggested that this was a case of clinal phenotypic variation, with a gradual reduction in spinocity occurring from west to east. Individuals collected from the Z. laevigatus type locality of Entabeni were indeed less spinose in appearance than typical forms of S. depressus from 10 km north-east of Louis Trichardt (Fig. 7), and strongly resembled females of the two northern species S. mossambicus and S. regius. Spinose as well as relatively smooth-bodied specimens of S. depressus have also been recorded from east of Entabeni (Pienaar, Haacke & Jacobsen, 1983), but such specimens were not available for this study. A fairly deep divergence was seen within S. depressus, but both the typically spiny and smooth-scaled phenotypes were recovered on either side of the divide. Although there is genetic structure within this group, these clades do not correspond to a particular phenotype or distribution pattern. The consistency of this pattern across the separate mitochondrial and nuclear analyses suggests that this is not a result of introgression, and while it could conceivably be a result of incomplete lineage sorting, our limited sampling in this area was unable to reconcile this. Pending further work, we recommend that Z. laevigatus should remain in synonymy with S. depressus. All analyses recovered the specimen collected near the type locality of S. w. perkoensis within the S. vandami clade in the central part of the range. The three deeply divergent lineages of Smaug vandami may well represent independently evolving metapolulations (species). However, as our sampling is not comprehensive in this group, finer-scale phylogeographical analyses should be performed before any taxonomic re-assessment is attempted and Z. v. perkoensis should, for the moment, remain in the synonymy of S. vandami.

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. DOI: 10.1111/zoj.12187

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.  DOI: 10.1016/j.ympev.2010.08.024

[Herpetology • 2015] Platysaurus attenboroughi • A New Species of Spectacularly Coloured Flat Lizard Platysaurus (Squamata: Cordylidae: Platysaurinae) from southern Africa

Platysaurus attenboroughi

  Whiting, Branch, Pepper & Keogh, 2015

Attenborough’s Flat Lizard | Attenborough se Platakkedis || DOI: 11646/zootaxa.3986.2.2 || WhitingLab.com

Abstract

We describe a new species of flat lizard (Platysaurus attenboroughi sp. nov.) from the Richtersveld of the Northern Cape Province of South Africa and the Fish River Canyon region of southern Namibia. This species was formerly confused with P. capensis from the Kamiesberg region of Namaqualand, South Africa. Genetic analysis based on one mtDNA and two nDNA loci found Platysaurus attenboroughi sp. nov. to be genetically divergent from P. capensis and these species can also be differentiated by a number of scalation characters, coloration and their allopatric distributions. To stabilize the taxonomy the type locality of Platysaurus capensis A. Smith 1844 is restricted to the Kamiesberg region, Namaqualand, Northern Cape Province, South Africa.

Keywords: Reptilia, southern Africa, lizard, new species, reptile, Platysaurus attenboroughi sp. nov., Platysaurus capensis

Platysaurus attenboroughi sp. nov.

English: Attenborough’s Flat Lizard

Afrikaans: Attenborough se Platakkedis

Synonymy. Platysaurus capensis (part). FitzSimons, 1935: 535; 1943: 473; Loveridge, 1944: 97; Rose, 1950: 155; 1962: 156; Broadley, 1978: 157; Branch, 1998: 165; Van Wyk & Mouton, 1996: 117; Whiting, 2014: 214.

Distribution. Along the lower Orange River from Goodhouse to the Richtersveld, extending north into Namibia and recorded from the Hunsberg, Huamsib and Ploegberg mountains and the Fish River Canyon (Figure 1).

Habitat and climate. Platysaurus attenboroughi sp. nov. occurs in the arid-subtropical region of the Northern Cape Province of South Africa and southern Namibia and specifically within the Gariep Desert Bioregion (Mucina & Rutherford, 2006). This is an arid area characterized by low and erratic summer rainfall. Summers are typically hot and dry. Like all flat lizards, they are dependent on rock (mostly granite in this area) and take refuge in narrow rock fissures where they can escape suboptimal temperatures and predators. These areas are largely devoid of significant vegetation bar the occasional fig tree (Ficus) or succulent. For more detailed descriptions of climate, vegetation and topography see Mucina & Rutherford (2006).

Natural history and behaviour. In South Africa, it is mainly restricted to the Richtersveld region (Bauer & Branch, 2001; Whiting, 2014). There it is widespread and common in boulder-strewn areas and on broad rock faces, often far from river courses (e.g. Tierhoek). It does not form the high density populations recorded for P. broadleyi (MJW unpubl. data). All Platysaurus have a fixed clutch of two eggs (Broadley, 1978). The reproductive cycles of P. capensis, P. broadleyi and P. attenboroughi sp. nov. were collectively studied when these were considered a single species (Van Wyk & Mouton, 1996). The minimum size at sexual maturity is 64 mm (sex not specified); eggs are likely laid in summer (November–December) (Van Wyk & Mouton, 1996). While we know

very little about the diet of P. attenboroughi sp. nov., the closely related P. broadleyi is an omnivore and lives in similar habitat. The marked sexual dichromatism suggests a classic sexual selection system in which males compete heavily for females, as is the case in P. broadleyi (Whiting et al., 2003; Whiting et al., 2006). Males do have UV-reflective throats, which suggests a role of this colour signal in either settling contests (as in P. broadleyi) or in mate choice, although this remains to be tested. In the two males we measured, their throats had violet and blue and less pure UV than we would typically see in adult male P. broadleyi (Figure 5) (Whiting et al., 2006). Measurement of spectral reflectance in additional individuals is necessary for a proper comparison.

Etymology. We name this new species in honour of Sir David F. Attenborough (Fig. 3), in recognition of his immense contribution to the public understanding and appreciation of animals, plants, ecosystems and nature in general. David Attenborough made flat lizards, specifically the closely related Platysaurus broadleyi, famous in the BBC documentary series Life in Cold Blood.

Whiting, Martin J., William R. Branch, Mitzy Pepper & J. S. Keogh. 2015. A New Species of Spectacularly Coloured Flat Lizard Platysaurus (Squamata: Cordylidae: Platysaurinae) from southern Africa. Zootaxa. 3986(2): 173–192. DOI: 11646/zootaxa.3986.2.2

http://zoobank.org/urn:lsid:zoobank.org:pub:986B0CFE-77D7-453F-8708-5F60F79624BE

http://publicationslist.org/data/martin.whiting/ref-146/Whiting_Branch_Pepper_Keogh_2015_Zootaxa_Platysaurus_attenboroughi_new_species.pdf

New African flat lizard named for David Attenborough  http://whitinglab.com/?p=4930