Catalog of the Family Cordylidae

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Catalog of the Family

Cordylidae

Harold De Lisle 2016

Cover photo : Platysaurus orientalis, by Nilsroe CC BY-SA 4.0 Back cover : Ouroborus cataphractus in defence posture, by Hardaker

Email: [email protected]

CONTENTS

INTRODUCTION …...... 1

CATALOG …...... 6

ACKNOWLEGEMENTS …...... 20

REFERENCES . …...... 21

TAXON INDEX …...... 32

The Cordylidae is a family of small to medium-sized lizards that occur in southern and eastern Africa. They are commonly known as girdled lizards, spinytail lizards or girdle-tail lizards.

Cordylid lizards are diurnal and mainly insectivorous. They are terrestrial, mostly inhabiting crevices in rocky terrain, although at least one species digs burrows and another lives under exfoliating bark on trees, and one genus lives mainly on grassy savannahs.

They have short tongues covered with long papillae, flattened heads and bodies. The body scales possess osteoderms and have large, rectangular, scales, arranged in regular rows around the body and tail. Many species have rings of spines on the tail, that aid in wedging the animal into sheltering crevices, and also in dissuading predators. The body is often flattened, and in most species is box-like on cross-section, with lateral fold.

The morphology and organization of osteoderms is highly variable in the Cordylidae. High-resolution X- ray Computed Tomography (CT) is ideal for recovering information about individual osteoderm structure, as well as overall arrangement of these features. The images above show just a small part of the variation in armor across the Cordylidae. Courtesy of the AMNH

Most species have four limbs, but those in the genus Chamaesaura are almost entirely limbless, with only tiny spikes in place of the hind limbs. The family includes both oviparous and ovoviviparous species.

Currently (2016) ten genera are recognized: Chamaesaura , with five species Cordylus, with twenty-one species Hemicordylus , with two species Karusasaurus, with two species Namazonurus, with five species Ninurta, with one species (montotypic) Ouroborus, with one species (montotypic) Platysaurus , with sixteen species Pseudocordylus , with six species Smaug , with eight species

The phylogeny of the Cordylidae is shown below (from Pyron et al., 2013) with position of each genus.

This Catalog is an attempt to summarize the genus and species taxonomic names of extant and recently extinct cordylid lizards. Several species concepts have been applied to these lizards, and changes in the species level taxonomy have been proposed and many more are to be expected for that reason alone. The systematic arrangement of the catalog is intuitive once a few principles are stated. Subspecific entries for each species are indented. Dates in brackets indicate that these have been subsequently determined from sources external to the original document. Misspellings are generally excluded except where they have been widely used or misinterpreted. Junior subjective synonyms are listed under each valid name. In order to enhance the usefulness of the present list, the type species of each genus has been added keeping in mind the famous quote of Harold Cogger, "Taxonomy is a matter of opinion" (Golay et al., 1993) based, of course, on the evaluation of the best evidence. The taxonomy used herein is my evaluation and we accept full responsibility for its use. We have deliberately omitted most lapsus calami, nomina nuda, nomina dubia, nomina oblita, nomina substituta , etc., from the synonymy because these names are not available for use. The synonymy herein is limited to new descriptions. Entries for species consist of species or subspecies name, genus, author, year of description, publication page, type locality, location of primary (name-bearing) type specimen(s), if known, and known distribution. Type localities are those in the original description, except those in obvious error or those amended subsequently by authoritative sources. Acronyms of type collections employed in the catalogue follow those used by American Museum of Natural History.

Abbreviations of museums holding the types are as follows

ANSP - Academy of Natural Sciences, Department of Herpetology, 19th and the Benjamin Franklin Parkway, Philadelphia, Pennsylvania 19103, USA.

BMNH – The Natural History Museum, Department of Zoology, Cromwell Road, London SW7 5BD, United Kingdom.

CAS - California Academy of Sciences, Department of Herpetology, Golden Gate Park, San Francisco, California 94118, USA

MBL - Museu Bocage, Universidade de Lisboa, Lisbon, Portugal [destroyed by fire in March 1978]

MCZ - Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, USA.

MD - Museu Regional do Dundo, Dundo, Luanda-Norte, Angola

MNHNP - Museum National dHistoire Naturelle, Laboratoire des Amphibiens et Reptiles, 25 Rue Cuvier, 75005 Paris, France

MSNG - Museo Civico di Storia Naturale di Genova Giacomo Doria, Via Brigata Liguria 9, 16121 Genova, Italy.

NHMW - Naturhistorisches Museum, Zoologische Abtheilung, Postfach 417, Burgring 7, 1014 Wien, Austria

NHRM - Naturhistoriska Rijkmuseet, Section for Vertebrate Zoology, Roslagsvägen 120, Box 50007, 104 05 Stockholm 50, Sweden.

NMB - National Museum Bloemfontein, Bloemfontein Central, Bloemfontein, 9301, South Africa

NMP – KwaZulu-Natal Museum, Pietermaritzburg 3201, KwaZulu-Natal, South Africa

NMZB - National History Museum of Zimbabwe, Bulawayo, Zimbabwe.

PEM - Port Elizabeth Museum, Port Elizabeth, Republic of South Africa.

QM - Queensland Museum, Gregory Terrace, Fortitude Valley, Brisbane, Queensland 4006, Australia.

RMNH - Naturalis (Nationaal Natuurhistorisch Museum; formerly Rijksmuseum van Natuurlijke Historie), Postbus 9517, 2300RA Leiden, The Netherlands

SAM - South African Museum, P.O. Box 61, Cape Town 8000, Western Cape Province, South Africa.

TMP - Transvaal Museum, P.O. Box 413, Pretoria 0001, Gauteng, South Africa.

UTEP - University of Texas at El Paso, Department of Biological Sciences and Centennial Museum, Laboratory for Environmental Biology, 500 W University Ave, El Paso, TX 79968, USA

ZMA - Universiteit van Amsterdam, Zoologisch Museum, Plantage Middenlaan 53, Amsterdam: C, Netherlands (now incorporated into NCB Naturalis, Leiden).

ZMB - Universität Humboldt, Zoologisches Museum, Invalidenstrasse 43, 10115 Berlin, Germany.

ZMH - Zoologisches Institut und Museum, Universität Hamburg, Martin-Luther-King Platz 3, D-2000 Hamburg 13, Germany.

Cordylidae Mertens, 1937 Mertens, R. 1937. Reptilien und Amphibien aus dem südlichen Inner-Afrika . Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft , Frankfurt, vol. 435, p. 1–23 Type genus : Cordylus Laurenti, 1768

Chamaesaura Schneider, 1801 Schneider, J.G. 1801. Historiae Amphibiorum naturalis et literariae. Fasciculus secundus continens Crocodilos, Scincos, Chamaesauras, Boas. Pseudoboas, Elapes, Angues. Amphisbaenas et Caecilias., Frommani, Jena. 364 pp. Type species: Lacerta anguina Linnaeus, 1758: 210

Chamaesaura aenea (Fitzinger, 1843)

Cricochalcis aenea Fitzinger 1843: 21 Type locality: South Africa Holotype: ZMB 1172 Distribution: South Africa: Northwest, Free State, Eastern Cape, KwaZulu-Natal provinces; Swaziland

Chamaesaura anguina Linnaeus, 1758

Chamaesaura anguina anguina (Linnaeus, 1758) Lacerta anguina Linnaeus 1758: 210 Type locality: Cape of Good Hope, South Africa Holotype: BMNH 1946.8.7.98 Distribution: South Africa: Eastern Cape, Western Cape, Gauteng, Mpumalanga, KwaZulu-Natal provinces

Chamaesaura anguina oligopholis Laurent, 1964 Chamaesaura anguina oligopholis Laurent, 1964: Type locality: Calonda, Lunda, Angola Holotype: MD ? Distribution: Angola

Chamaesaura macrolepis (Cope, 1862)

Mancus macrolepis Cope, 1862: 339 Type locality: Umvoti, KwaZulu-Natal, South Africa Syntypes: ANSP 9708-09 Distribution: South Africa: Northwest, Mpumalanga, KwaZulu-Natal provinces; Swaziland; southestern Zimbabwe

Chamaesaura miopropus Boulenger, 1895 Chamaesaura miopropus Boulenger, 1895 : 724 Type locality: Fwambo, northeastern Zambia Holotype: BMNH 1946.8.29.49 Distribution: northern Zambia; southern Tanzania, eastern DR Congo

Chamaesaura tenuior Günther, 1895 Chamaesaura tenuior Günther, 1895: 524 Type locality: Kampala, Uganda. Syntypes: : BMNH 1946.8.7.95-97 Distribution: Kenya: central highlands; northern Tanzania; eastern Rwanda; southern Uganda; northeastern DR Congo

Cordylus Laurenti, 1768 Laurenti, J.N. 1768. Specimen medicum , exhibens synopsin reptilium emendatam cum experimentis circa venena et antidota reptilium austracorum, quod authoritate et consensu. Vienna, Joan. Thomae, 217 pp. Type species : Cordylus verus Laurenti, 1768

Cordylus angolensis (Bocage, 1895)

Zonurus angolensis Bocage, 1895: 25 Type locality: Caconda, Huila District, Angola Holotype: MBL (probably lost in fire 1978) Distribution: Angola highlands ., northern Namibia, south- central DR Congo

Photo: Xesko

Cordylus aridus Mouton & Van Wyk, 1994 Cordylus aridus Mouton & Van Wyk, 1994: 15 Type locality: Farm Botterkraal, Prince Albert District, Western Cape Province, South Africa Holotype: SAM 50901 Distribution: South Africa: Western Cape Province

Cordylus beraduccii Broadley & Branch, 2002

Cordylus beraduccii Broadley & Branch, 2002: 22 Type locality: Ndukusi village, 10 km SE of Lolkisale, Kiteto District, Arusha Region, Tanzania . Holotype: NMZB 16923 Distribution: northern Tanzania and southern Kenya (plateau areas).

Photo: Petra Bejuk

Cordylus cloetei Mouton & Van Wyk, 1994

Cordylus cloetei Mouton & van Wyk, 1994: 12 Type locality: Farm De Hoek, Nuweveld Mountains, Fraserburg District, South Africa . Holotype: SAM 50893 Distribution: South Africa: Northern Cape Province Last seen 1993

Photo: Petra Bejuk

Cordylus cordylus (Linnaeus, 1758)

Lacerta cordylus Linnaeus, 1758: 202 Type locality: Africa Syntypes: NHRM 115-118 Synonyms: Cordylus verus Laurenti, 1768 Type locality: Africa Holotype: not located Cordylus dorsalis Cuvier, 1829 Type locality: Africa Holotype: MNHNP, not located Cordylus griseus Cuvier, 1829 Type locality: Africa Holotype: MNHNP, not located Zonurus vertebralis Gray, 1838 Type locality: Cape of Good Hope, South Africa Syntypes: BMNH V.2.a-e, V.3.f-g Zonurus cordylus flavus Rose, 1926 Type locality: Robben Island, Western Cape Province, South Africa Holotype: not located Zonurus cordylus tasmani Power, 1930 Type locality: Dunbrody, Uitenhage District, South Africa. Photo: A Shawka Syntypes: PEM R13539, 13513-14, 13522, 13518-21, 13582, 13583, 13515-17 Distribution: South Africa: Northern, Western. Eastern Cape southeastern Free State Provinces; Lesotho

Cordylus imkeae Mouton & Van Wyk, 1994

Cordylus imkeae Mouton & van Wyk, 1994: 14 Type locality: Lower slopes of the Rooiberg on the farm Wolfhok (30°24'27"S, 18°06'11" E), Garies District, Namaqualand, South Africa Holotype: SAM 50897 Distribution: South Africa:Western Cape Province: Rooiberg Mts, Namaqualand

Photo: Petra Beju

Cordylus jonesii (Boulenger, 1891)

Zonurus jonesii Boulenger, 1891: 417 Type locality: Murchison Range, northeast Transvaal, Northern Province, South Africa Holotype: BMNH 1946.8.29.40 Distribution: eastern Botswana; southern Zimbabwe; southern Mozambique; South Africa: Northern, Mpumalanga, KwaZulu- Natal Provinces; eastern Swaziland.

Photo: C.K. Willis

Cordylus machadoi Laurent, 1964 Cordylus vittifer machadoi Laurent, 1964: 49 Type locality: Leba, Humpata, near Sá da Bandeira, Húila, Angola. Holotype: MD ? Distribution: Angola: Huila Province; Namibia: extreme northern Kaokoveld

Cordylus macropholis (Boulenger, 1910)

Zonurus macropholis Boulenger, 1910: 494 Type locality: Little Namaqualand, South Africa. Holotype: SAM 434 Distribution: South Africa: Northern Cape Province: coastal areas of Little Namaqualand

Cordylus marunguensis Greebaum, Stanley, Kusamba, Moninga, Goldberg & Bursey, 2012 Cordylus marunguensis Greebaum, Stanley, Kusamba, Moninga, Goldberg & Bursey, 2012: 26 Type locality: Pepa (07°43’08.1’’S, 29°45’52.4’’E), Marungu Mountains, Katanga Province, DR Congo Holotype: UTEP 20374 Distribution: DR Congo: Katanga Province: Marungu Mountains

Cordylus mclachlani Mouton, 1986

Cordylus mclachlani Mouton, 1986: 319 Type locality: Koue Bokkeveld, near Ceres, South Africa. Holotype: SAM 47100 Distribution: South Africa: Western Cape Province: Koue Bokkeveld to Nieuwoudtville

Photo: Dorse & van Rooyen

Cordylus meculae Branch, Rödel & Marais, 2005 Cordylus meculae Branch, Roedel & Marais, 2005: 132 Type locality: Serra Mecula, Niassa Game Reserve (12°03'11"S, 37°38'49"E), Niassa Province, northern Mozambique Holotype: PEM R16166 Distribution: Mozambique: Serra Mecula

Cordylus minor Fitzsimons, 1943 Cordylus cordylus minor FitzSimons, 1943: 458 Type locality: just north of Matjesfontein,Western Cape Province, South Africa. Syntypes: TMP 19563-19568 Distribution: South Africa: Western Cape Province: southwestern Karoo

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

Branch, 2016: 12 Type locality: large rock outcrop near Caraculo (15°0'59.40"S, 12°38'31.30"E), on the road from Lubango and Namibe, Namibe province, Angola Holotype: CAS 254912 Distribution: Angola: south and central parts of Namibe Province, southwest of the Leba escarpment

Cordylus niger Cuvier, 1829

Cordylus niger Cuvier, 1829: 33 Type locality: Africa. Syntypes: MNHNP 0.2795 Synonyms: Calyptoprymnus verecundus De Vis, 1905 Type locality: unknown Holotype: QM ? Zonurus cordylus atrus Power, 1930 Type locality: Cape Peninsula, South Africa. Holotype: TMP ? Distribution: South Africa: Western Cape Province: Table Mountain, Cape Peninsula and Saldanha Photo: P Head

Cordylus nyikae Broadley & Mouton, 2000 Cordylus nyikae Broadley & Mouton, 2000: 169 Type locality: near Chelinda Bridge, Nyika National Park (10°35'S, 33°48'E), Malawi. Holotype: NMZB 24352. Distribution: northern Malawi: Nyika Plateau, Misuku Hills and at Chitipa

Cordylus oelofseni Mouton & Van Wyk, 1990

Cordylus oelofseni Mouton & van Wyk, 1990: 34 Type locality: Dasklip Pass near Porterville, Western Cape Province, South Africa. Holotype: SAM 47388 Distribution: South Africa: Western Cape Province: western borders of the Cape Fold Mountains

Photo: TH

Cordylus phonolithos Marques, Ceríaco, Stanley, Bandeira, Agarwal, Bauer, 2019 Cordylus phonolithos Marques, Ceríaco, Stanley, Bandeira, Agarwal, Bauer, 2019: 509 Type locality: within a crevice in a granite boulder in the vicinity of N’Dolondolo (13.80678ºS, 13.13507ºE), Namibe Province, Angola Holotype: CAS 263581 Distribution: Angola: Namibe Province, Serra de Neve

Cordylus rhodesianus (Hewitt, 1933) Zonurus cordylus rhodesianus Hewitt, 1933: 48 Type locality: Monte Cassino Mission, Macheke, Zimbabwe Syntypes: PEM 5294-96 Distribution: Zimbabwe: Eastern highlands; Mozambique: Chimanimani Mountains

Cordylus rivae (Boulenger, 1896) Zonurus rivae Boulenger, 1896: 8 Type locality: Giacorsa, Gallaland, Ethiopia. Holotype: MSNG 28900 Distribution: Ethiopia: Sidamo region

Cordylus tropidosternum (Cope, 1869)

Zonurus tropidosternum Cope, 1869: 169 Type locality: Mozambique Holotype: MCZ R-5742 Synonyms: Zonurus frenatus Pfeffer, 1889 Type locality: Mhonda, Tanzania Holotype: ZMH, probably lost in WWII Zonurus parkeri Cott, 1934 Type locality: Amatongas, Sofala Province, Mozambique Holotype: BMNH 1946.8.29.42 Distribution: southern coastal Kenya; eastern Tanzania; to central Mozambique west to southeast DR Congo (Katanga); northern Zambia; Malawi; northeastern Zimbabwe

Photo: Petra Beju

Cordylus ukingensis (Loveridge, 1932) Zonurus ukingensis Loveridge, 1932: 378 Type locality: Tandala, Ukinga mountains, southwestern Tanzania Holotype: MCZ 30761 Distribution: Tanzania: southern highlands, Ukinga and Udzungwa Mountains

Cordylus vittifer (Reichenow, 1887)

Zonurus vittifer Reichenow, 1887: 25 Type locality: Transvaal, South Africa . Holotype: ZMB 10762 Photo: M Robinson Synonym: Zonurus tropidogaster Boulenger, 1910 Type locality: Barberton, Mpumalanga Province, South Africa Holotype: not located Distribution: southeastern Botswana; South Africa: Northwest, Northern, Gauteng, Mpumalanga, Free State,KwaZulu-Natal Provinces; Swaziland; southwestern Mozambique: eastern slopes of the Lebombo Range

Hemicordylus Smith, 1838 Smith, A. 1838. Contributions to South African zoology. Art. VI. Annals and Magazine of Natural History (1) 2: 30-33 Type species: Hemicordylus capensis Smith, 1838: 32

Hemicordylus capensis (Smith, 1838)

Cordylus capensis Smith, 1838: 32 Type locality: Hottentots Holland Mountains, Western Cape Province, South Africa Holotype: BMNH 1946.8.7.89 Synonym: Zonurus robertsi Van Dam, 1921 Type locality: Klaver, Van Rhynsdorp District, Western Cape Province, South Africa Holotype: TMP 3747 Distribution: South Africa: Western Cape Province

Photo: A Shawka

Hemicordylus nebulosus (Mouton & Van Wyk, 1995) Pseudocordylus nebulosus Mouton & van Wyk, 1995: 391 Type locality: northern slope of Landdroskop, Hottentots-Holland Mountains, South Africa Holotype: SAM 50905 Distribution: South Africa: Western Cape Province: Hottentots Holland Mountains

Karusasaurus Stanley, Bauer, Jackman, Branch & Mouton, 2011 Stanley, E.L., A.M. Bauer, T.R. Jackman, W.R. Branch & P.L.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 Type species: Cordylus polyzonus Smith, 1838: 34

Karusasaurus jordani (Parker, 1936) Zonurus jordani Parker, 1936: 133 Type locality: Hoffnung, near Windhoek, Namibia Holotype: BMNH 1946.8.7.92 Distribution: central Namibia, north of Aus

Karusasaurus polyzonus (Smith, 1838)

Cordylus polyzonus Smith, 1838: 31 Type locality: South Africa Syntypes: BMNH 1946.8.8.16-19 Distribution: South Africa: Northern Cape, Western Cape, Eastern Cape, Free State Provinces; southern Namibia

Photo: Gawie

Namazonurus Stanley, Bauer, Jackman, Branch & Mouton, 2011 Stanley, E.L., A.M. Bauer, T.R. Jackman, W.R. Branch & P.L.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 Type species: Zonurus pustulatus Peters, 1862: 19

Namazonurus campbelli (Fitzsimons, 1938)

Zonurus campbelli Fitzsimons, 1938: 189 Type locality: Farm Barby, 25 km west of Helmeringhausen , Great Namaqualand, Namibia Holotype: TMP 17635 Distribution: Namibia: Bethanie District

Photo: J Marais

Namazonurus lawrenci (Fitzsimons, 1939) Zonurus lawrenci Fitzsimons, 1939: 208 Type locality: Lekkersing, Richtersveld, Little Namaqualand, South Africa Holotype: SAM 18553 Distribution: South Africa: North Cape Prov., Richtersveld; southwestern Namibia

Namazonurus namaquensis (Methuen & Hewitt, 1914) Zonurus namaquensis Methuen & Hewitt, 1914: 137 Type locality: Farm Narudas Süd, Great Karasberg District, Great Namaqualand, Namibia Holotype: TMP 3163 Distribution: southern Namibia: Great Karasberg district

Namazonurus peersi (Hewitt, 1932) Zonurus peersi Hewitt, 1932: 116 Type locality: Garies, Little Namaqualand, South Africa Syntypes: SAM 43852 Distribution: South Africa: North Cape Prov: Little Namaqualand

Namazonurus pustulatus (Peters, 1862)

Zonurus pustulatus Peters, 1862: 19 Type locality: Otjimbingue, Namibia Syntypes: ZMB 4221-22, 4361 Distribution: central Namibia

Photo: Guy

Ninurta Stanley, Bauer, Jackman, Branch & Mouton, 2011 Stanley, E.L., A.M. Bauer, T.R. Jackman, W.R. Branch & P.L.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 Type species: Zonurus coeruleopunctatus Hewitt & Methuen,1913: 110

Ninurta coeruleopunctatus (Methuen & Hewitt, 1913)

Zonurus coeruleopunctatus Methuen & Hewitt, 1913: 110 Type locality: Buffel’s Nek, between Knysna and Avontuur, South Africa Holotype: TMP ? Distribution: South Africa: Western Cape & Eastern Cape provinces: Mossel Bay to Witelsbos

Photo: Shaun Swanepoel

Ouroborus Stanley, Bauer, Jackman, Branch & Mouton, 2011 Stanley, E.L., A.M. Bauer, T.R. Jackman, W.R. Branch & P.L.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 Type species: Cordylus cataphractus Boie, 1828: 140

Ouroborus cataphractus (Boie, 1828) Cordylus cataphractus Boie, 1828: 140 Type locality: South Africa Holotype: RMNH 3603 Synonym: Cordylus nebulosus Smith, 1838 Type locality: South Africa Syntypes: BMNH 1865.5.4.72-73 Distribution: South Africa: southern Richtersveld, Northern Cape Prov., to Matjiesfontein, Western Cape Prov.

Platysaurus Smith, 1844 Smith, A. 1844. Illustrations of the zoology of South Africa, Reptilia. Smith, Elder, and Co., London Type species: Platysaurus capensis Smith, 1844: pl 40

Platysaurus attenboroughi Whiting, Branch, Pepper & Keogh, 2015 Platysaurus attenboroughi Whiting, Branch, Pepper & Keogh,

2015: 185 Type locality: Fish River Canyon (27 ̊52'21.7°S, 17 ̊31'15.7°E), Karasburg District, southern Namibia Holotype: TMP 85806 Distribution: South Africa: Northern Cape Province: lower Orange River from Goodhouse to the Richtersveld; southern Namibia: Hunsberg, Huamsib and Ploegberg Mountains and the Fish River Canyon Photo: M Whiting

Platysaurus broadleyi Branch & Whiting, 1997

Platysaurus broadleyi Branch & Whiting, 1997: 126 Type locality: Augrabies Falls National Park, Gordonia District, Northern Cape Province, South Africa Holotype: TMP 79828 Distribution: South Africa: Northern Cape Province: from Augribies to Klein Pella

Photo: B Dupont

Platysaurus capensis Smith, 1844

Platysaurus capensis Smith, 1844: pl 40 Type locality: Kamiesberg region, Namaqualand, Northern Cape Province, South Africa Lectotype: BMNH 1946.8.29.26 Distribution: South Africa: Northern Cape Province: greater Kamiesberg region, from Carolusberg to near Springbok south of Garies; southern Namibia: southern Lüderitz, Bethanie, and Karasburg districts

Platysaurus guttatus Smith, 1849

Platysaurus guttatus Smith, 1849: 8 Type locality: near the Limpopo River, South Africa Holotype: BMNH 1946.8.10.33 Distribution: South Africa: Northwest, Limpopo provinces; eastern Botswana; southwestern Mozambique; southern Zimbabwe

Photo: G Matz

Platysaurus imperator Broadley, 1962 Platysaurus imperator Broadley, 1962: 816 Type locality: Matowa, 60 km NE of Mtoko, Zimbabwe Holotype: NMZB 4819 Distribution: northeast Zimbabwe; northwestern Mozambique

Photo: ElinorD

Platysaurus intermedius Matschie, 1891

Platysaurus intermedius intermedius Matschie, 1891 Platysaurus intermedius Matschie, 1891: 606 Type locality: Station Mphome, near Haenertsburg, Limpopo Province, South Africa Lectotype: ZMB 3466 Distribution: South Africa: Limpopo Province: Soutpansberg Mountains

Platysaurus intermedius inopinus Jacobsen, 1994 Platysaurus intermedius inopinus Jacobsen, 1994: 139 Type locality: Farm Glen Alpine 304LR, 23° 12' S, 28° 40'E, Potgietersruts (= Mokopane) district, Limpopo Province, South Africa Holotype: TMP 70018 Distribution: South Africa: Limpopo Province: foothills of the Blouberg

Platysaurus intermedius natalensis Fitzsimons, 1948 Platysaurus guttatus natalensis Fitzsimons, 1948: 74 Type locality: Louwsburg, northern KwaZulu-Natal Holotype: TMP 21716 Distribution: South Africa: KwaZulu-Natal Province; Swaziland

Platysaurus intermedius nigrescens Broadley, 1981 Platysaurus intermedius nigrescens Broadley, 1981: 161 Type locality: Shoshong Hills (22°58’55”S, 26°22’80”E), 20km west of Mahalapye, Botswana Holotype: NMZB 33687 Distribution: northeastern Botswana Photo: Sputniktilt

Platysaurus intermedius nyasae Loveridge, 1953 Platysaurus guttatus nyasae Loveridge, 1953: 237 Type locality: rocks by roadside on Blantyre-State road south of Mpatamaya Bridge, Malawi Holotype: MCZ R-50656 Distribution: Malawi; west-central Mozambique

Platysaurus intermedius parvus Broadley, 1976 Platysaurus intermedius parvus Broadley, 1976: ? Type locality: Blouberg, Soutpansberg District, Limpopo Province , South Africa. Holotype: TMP ? Distribution: South Africa: Limpopo Province: Blouberg Massif

Platysaurus intermedius rhodesianus Fitzsimons, 1941 Platysaurus guttatus rhodesianus Fitzsimons, 1941: 31 Type locality: Vumba Mountain, Zim babwe Holotype: TMP ? Distribution: southern Zimbabwe; northeastern Botswana; South Africa: northern Limpopo Province

Platysaurus intermedius subniger Broadley, 1962 Platysaurus guttatus subniger Broadley, 1962: 814 Type locality: Watsomba, a granite peak 16 miles NNW of Umtali, Zimbabwe Holotype: NMZB 1392 Distribution: northeastern Zimbabwe

Platysaurus intermedius wilhelmi Hewitt, 1909 Platysaurus wilhelmi Hewitt, 1909: 32 Type locality: Nelspruit, Barberton District, Mpumalanga province, South Africa Holotype: TMP ? Distribution: South Africa: Mpumalanga and KwaZululand-Natal provinces; Swaziland

Platysaurus lebomboensis Jacobsen, 1994

Platysaurus lebomboensis Jacobsen, 1994: 140 Type locality: Mananga hill (25°58'S, 31°52'E), Barberton district, Mpumalanga province, South Africa Holotype: TMP 70037 Distribution: South Africa: KwaZululand Province: Middle Lebombo Range; southern Mozambique; Swaziland

Photo: J Marais

Platysaurus maculatus Broadley, 1965 Platysaurus maculatus maculatus Broadley, 1965 Platysaurus maculatus maculatus Broadley, 1965: 3 Type locality: Mitucué Mountain, Niassa Province, Mozambique Holotype: NMZB 8054 Distribution: southern Tanzania; northern Mozambique

Platysaurus maculatus lineicauda Broadley, 1965 Platysaurus maculatus lineicauda Broadley, 1965: 3 Type locality: 14 mi. west of Morrumbala, Zambezia Province, Mozambique Holotype: NMZB ? Distribution: Mozambique: Zambezia Province

Platysaurus minor Fitzsimons, 1930

Platysaurus guttatus minor Fitzsimons, 1930: 30 Type locality: at Vygeboompoort, Waterberg District, north Limpopo Province, South Africa Syntypes: TMP 2109, 2367 Distribution: South Africa: Limpopo Province: Waterberg Massif to the Blouberg Massif

Photo: R van Huyssteen

Platysaurus mitchelli Loveridge, 1953 Platysaurus mitchelli Loveridge, 1953: 234 Type locality: Ruo River Forest, Mt Mlanje, Malawi Holotype: MCZ R-50657 Distribution: southern Malawi: Mulanje Mountains

Platysaurus monotropis Jacobsen, 1994 Platysaurus monotropis Jacobsen, 1994: 136 Type locality: Farm Blackhill 317 LR, Potgietersrust district, Limpopo Province, South Africa Holotype: TMP 70003 Distribution: South Africa: Limpopo Province, foothills of the Blouberg

Platysaurus ocellatus Broadley, 1962

Platysaurus ocellatus Broadley, 1962: 810 Type locality: western slopes of the Chimanimani Mts., above Dead Cow Camp, 8 miles east of Melsetter, Zimbabwe Holotype: NMZB 1199 Distribution: east-central Zimbabwe: Chimanimani Mountains; west-central Mozambique

Photo: Brian Gratwicke

Platysaurus orientalis Fitzsimons, 1941

Platysaurus orientalis orientalis Fitzsimons, 1941 Platysaurus minor orientalis Fitzsimons, 1941: 280 Type locality: Sekororo, on the lower slopes of the Drakensberg, about 40 miles south of Leydsdorp, Limpopo Province, South Africa Syntypes: TMP 4527-4530 Distribution: South Africa: Drakensberg Mountains, southern Limpopo Province, South Africa

Platysaurus orientalis fitzsimonsi Loveridge, 1944 Platysaurus guttatus fitzsimonsi Loveridge, 1944: 88 Type locality: Lydenburg, Mpumalagna Prov., South Africa Holotype: MCZ 8982 Distribution: South Africa: Mpumalagna Escarpment, Mpumalagna Prov. Photo: Joachim

Platysaurus pungweensis Broadley, 1959 Platysaurus pungweensis pungweensis Broadley, 1959 Platysaurus guttatus pungweensis Broadley, 1959: 314 Type locality: Pungwe River (near the Mozambique border), eastern Zimbabwe Holotype: NMZB 1975 Distribution: eastern Zimbabwe: Eastern Highlands

Platysaurus pungweensis blakei Broadley, 1964 Platysaurus intermedius blakei Broadley, 1964: 1 Type locality: 15 miles south-east of Vila de Manica, Mozambique Holotype: NMZB ? Distribution: western Mozambique: Chimoio Plateau, Manica Prov., Photo: Milii Platysaurus relictus Broadley, 1976

Platysaurus relictus Broadley, 1976: ? Type locality: Waterpoort, Soutpansberg District, Limpopo Prov., South Africa Holotype: TMP 39637 Distribution: South Africa: Limpopo Province: northern side of the Soutpansberg Photo: R van Huyssteen

Platysaurus torquatus Peters, 1879

Platysaurus torquatus Peters, 1879: 10 Type locality: Tete, Mozambique Syntypes: ZMB 1129-1131, 9302-9303, MCZ R-37203, NMHW 15048 Distribution: northeastern Zimbabwe, west-central Mozambique, sothern Malawi

Photo: Matthijs

Pseudocordylus Smith, 1838 Smith, A. 1838. Contributions to South African zoology. Art. VI. Annals and Magazine of Natural History (1) 2: 30-33 Type species: Cordylus montanus Smith, 1838: 32

Pseudocordylus langi Loveridge, 1944

Pseudocordylus langi Loveridge, 1944: 73 Type locality: Mont-aux-Sources, Drakensberg, Lesotho Holotype: MCZ R-46835 Distribution: South Africa: KwaZulu-Natal: Drakensberg mountains; Lesotho

Photo: Alexander R

Pseudocordylus melanotus (Smith, 1838)

Cordylus melanotus Smith, 1838: 32 Type locality: South Africa, hills between the main branches of the Orange River east of Philippolis, Orange Free State Holotype: lost Distribution: South Africa: northern KwaZulu-Natal, Gauteng, eastern Mpumalanga, northeastern Free State Provinces

Photo: DK

Pseudocordylus microlepidotus (Cuvier, 1829)

Pseudocordylus microlepidotus microlepidotus (Cuvier, 1829) Cordylus microlepidotus Cuvier, 1829: 33 Type locality: South Africa Holotype: iconotype, based on Seba pl 62 Synonyms: Zonurus wittii Schlegel, 1834 Type locality: South Africa Holotype: not located Photo: nvlooi Cordylus montanus Smith, 1838 Type locality: South Africa Holotype: BMNH, not located Distribution: South Africa: Cape Fold Mountains, from Cedarberg to Port Elizabeth, Western Cape and Eastern Cape Provinces.

Pseudocordylus microlepidotus fasciatus (Smith, 1838) Cordylus fasciatus Smith, 1838: 32 Type locality: South Africa, rocky hills near Grahamstown Holotype: lost Synonym: Cordylus algoensis Smith, 1838 Type locality: Algoa Bay, Cape Province, South Africa Holotype: BMNH 1946.8.8.49 Distribution: South Africa: Eastern Cape Province

Pseudocordylus microlepidotus namaquensis Hewitt, 1927 Pseudocordylus microlepidotus namaquensis Hewitt, 1927: 392 Type locality: Namaqualand, Northern Cape Prov., South Africa Holotype: SAM 872 Distribution: South Africa: Northern Cape Prov.: Nuweveld Mountains, from Sutherland to Beaufort West

Pseudocordylus spinosus Fitzsimons, 1947

Pseudocordylus spinosus Fitzsimons, 1947: 116 Type locality: Cathkin Peak area, Drakensberg, KwaZulu- Natal Province, South Africa Holotype: TMP 21267 Distribution: South Africa: lower slopes of Drakensberg, from Giant's Castle in KwaZululand-Natal to Golden Gate in Free State

Photo: T Ping

Pseudocordylus subviridis (Smith, 1838) Cordylus subviridis Smith, 1838: 33 Type locality: South Africa Holotype: lost Distribution: South Africa: from Amatola Mountains, Eastern Cape Province, to Mont-Aux-Sources (Drakensberg), Lesotho

Photo: Armada44

Pseudocordylus transvaalensis Fitzsimons, 1943

Pseudocordylus subviridis transvaalensis Fitzsimons, 1943: 469 Type locality: Woodbush, Pietersburg District, Limpopo Province, South Africa Syntypes: TMP 1695, 1699 Distribution: South Africa: southern Limpopo Province

Photo: Arno Meintjs

Smaug Stanley, Bauer, Jackman, Branch & Mouton, 2011 Stanley, E.L., A.M. Bauer, T.R. Jackman, W.R. Branch & P.L.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 Type species: Cordylus giganteus Smith, 1844: pl 35

Smaug barbertonensis (Van Dam, 1921)

Zonurus barbertonensis Van Dam, 1921: 240 Type locality: Barberton, Mpumalanga Province, South Africa Holotype: TMP 4273 Distribution: South Africa: southeastern Mpumalanga, northern KwaZululand-Natal; eastern Swaziland

Photo: DR McKenzie

Smaug breyeri (Van Dam, 1921)

Zonurus breyeri V Van Dam, 1921: 239 Type locality: Farm Geelhoutkop, Waterberg District, Limpopo Province, South Africa Holotype: ZMA 10975 Distribution: South Africa: Limpopo Province: Waterberg Massif area

Photo: L Ghoul

Smaug depressus (Fitzsimons, 1930)

Zonurus barbertonensis depressus Fitzsimons, 1930: 24 Type locality: Newgate, near Louis Trichardt, Limpopo Province, South Africa Holotype: TMP 4621 Distribution: South Africa: Limpopo Province: middle and eastern Soutpansberg, to northern Krüger National Park and south to Duiwelskloof

Smaug giganteus (Smith, 1844)

Cordylus giganteus Smith, 1844: 35 Type locality: Interior districts of South Africa on pinnacles of Quathlamba Mountains Syntypes: BMNH 1946.8.10.32, 1946.8.8.48 Synonym: Zonurus derbianus GRAY, 1845 Type locality: South Africa Syntypes: BMNH 1946.8.8.20-21 Distribution: South Africa: northeastern Free State, extreme KwaZululand-Natal, southeastern Mpumalanga

Smaug mossambicus (Fitzsimons, 1958)

Cordylus warreni mossambicus Fitzsimons, 1958 : ? Type locality: Gorongoza Mountain, near Vila Paiva, Mozambique Holotype: NMP 1237 Distribution: Mozambique: Gorongoza Mountains; Zimbabwe: Chimanimani Mountains

Photo: KS Bader

Smaug regius (Broadley, 1962)

Cordylus warreni regius Broadley, 1962: 808 Type locality: Dora, 8 miles S.S.W. of Umtali, Zimbabwe Holotype: NMZB 1210 Distribution: Zimbabwe: southeastern Mutare District: Bvumba Mountains

Photo: J Pfleiderer

Smaug swazi Bates, Stanley, 2020

Smaug swazi Bates, Stanley, 2020: 11 Type locality: 320 m SSE of car park, Maguga Dam (26°04 ′57 ″S, 31° 15 ′59 ″E) , Hhohho Region, Eswatini (=Swaziland) Holotype: NMB R9201 Distribution: Swaziland: Highveld and Middleveld of Eswatini in Hhohho, Manzini and Shiselweni Regions; South Africa: eastern Mpumalanga and northern KwaZulu–Natal, Provinces

Photo:T. Busschau

Smaug vandami (Fitzsimons, 1930)

Zonurus vandami Fitzsimons, 1930: 25 Type locality: Gravelotte, near Leydsdorp, Limpopo Province, South Africa Holotype: TMP 7407 Synonym: Zonurus vandami perkoensis Fitzsimons, 1930 Type locality: Farm Perkoe, Olifant River District, Mpumalanga Province Holotype: TMP 12804 Distribution: South Africa: Mpumalanga Province: eastern escarpment of Mpumalanga Photo: scavenger

Smaug warreni (Boulenger, 1908) Zonurus warreni Boulenger, 1908: 232 Type locality: Ubombo, Zululand Holotype: BMNH 1946.8.8.1 Distribution: South Africa: Lebombo Range, southeastern Mpumalanga, northern KwaZululand-Natal Provinces; eastern Swaziland

Photo: James Culverwell

ACKNOWLEGEMENTS

This work could not have been completed without the help of numerous individuals. We thus acknowledge the help and advice of Patrick D Campbell (BMNH), Esther Dondorp (RMNH), Lemmy Mashinini (TMP), Werner Conradie (PEM), Jan Grathwohl

The author expresses his to the photographers who have contributed to this work, in particular, James Culverwell, J. Pfleiderer, K. S. Bader, Petra Bejuk, Abu Shawka, C.K. Willis and many others.

If any of the readers have any of the missing data (designated by ?), I would appreciate if they would contact me ([email protected])

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TAXON INDEX

A aenea (see Chamaesaura ) algoensis (see Pseudocordylus microlepidotus ) angolensis (see Cordylus ) anguina (see Chamaesaura ) aridus (see Cordylus ) atrus (see Cordylus niger ) attenboroughi (see Platysaurus )

B barbertonensis (see Smaug ) beraduccii (see Cordylus ) blakei (see Platysaurus pungweensis ) breyeri (see Smaug ) broadleyi (see Platysaurus )

C campbelli (see Namazonurus ) capensis (see Hemicordylus ) capensis (see Platysaurus ) cataphractus (see Ouroborus ) cloetei (see Cordylus ) coeruleopunctatus (see Ninurta ) cordylus (see Cordylus ) D derbianus (see Smaug giganteus ) depressus (see Smaug ) dorsalis (see Cordylus cordylus )

F fasciatus (see Pseudocordylus microlepidotus ) fitzsimonsi (see Platysaurus ) flavus (see Cordylus cordylus ) frenatus (see Cordylus tropidosternum )

G giganteus (see Smaug ) griseus (see Cordylus cordylus ) guttatus (see Platysaurus )

I imkeae (see Cordylus ) imperator (see Platysaurus ) inopinus (see Platysaurus intermedius ) intermedius (see Platysaurus )

J jonesii (see Cordylus ) jordani (see Karusasaurus )

L langi (see Pseudocordylus ) lawrenci (see Namazonurus ) lebomboensis (see Platysaurus ) lineicauda (see Platysaurus maculatus )

M machadoi (see Cordylus ) macrolepis (see Chamaesaura ) macropholis (see Cordylus ) maculatus (see Platysaurus ) marunguensis (see Cordylus ) mclachlani (see Cordylus ) meculae (see Cordylus ) melanotus (see Pseudocordylus ) microlepidotus (see Pseudocordylus ) minor (see Cordylus ) minor (see Platysaurus ) miopropus (see Chamaesaura ) mitchelli (see Platysaurus ) monotropis (see Platysaurus ) montanus (see Pseudocordylus microlepidotus ) mossambicus (see Smaug )

N namakuiyus (see Cordylus ) namaquensis (see Namazonurus ) namaquensis (see Pseudocordylus microlepidotus ) natalensis (see Platysaurus intermedius ) nebulosus (see Hemicordylus ) nebulosus (see Ouroborus cataphractus ) nigrescens (see Platysaurus intermedius ) niger (see Cordylus ) nyasae (see Platysaurus intermedius ) nyikae (see Cordylus )

O ocellatus (see Platysaurus ) oelofseni (see Cordylus ) oligopholis (see Chamaesaura ) orientalis (see Platysaurus )

P parkeri (see Cordylus tropidosternum ) parvus (see Platysaurus intermedius ) peersi (see Namazonurus ) perkoensis (see Smaug vandami ) phonolithos (see Cordylus ) polyzonus (see Karusasaurus ) pungweensis (see Platysaurus ) pustulatus (see Namazonurus )

R regius (see Smaug ) relictus (see Platysaurus ) rhodesianus (see Cordylus ) rhodesianus (see Platysaurus intermedius ) rivae (see Cordylus ) robertsi ( see Hemicordylus capensis )

S spinosus (see Pseudocordylus ) subniger (see Platysaurus intermedius ) subviridis (see Pseudocordylus ) swazi (see Smaug )

T tasmani (see Cordylus cordylus ) tenuior (see Chamaesaura ) torquatus (see Platysaurus ) transvaalensis (see Pseudocordylus ) tropidosternum (see Cordylus ) tropidogaster (see Cordylus vittifer )

U ukingensis (see Cordylus )

V vandami (see Smaug ) verecundus (see Cordylus niger ) vertebralis see Cordylus cordylus ) verus (see Cordylus cordylus ) vittifer (see Cordylus )

W warreni (see Smaug ) wilhelmi (see Platysaurus intermedius ) wittii (see Pseudocordylus microlepidotus )