title: A molecular phylogeny of the African plated lizards, genus Gerrhosaurus Wiegmann, 1828 (Squamata: Gerrhosauridae), with the description of two new genera.
abstract: We constructed a molecular phylogeny of the African plated lizard family Gerrhosauridae using two mitochondrial markers (ND2, 732 bp; 16S, 576 bp) and one nuclear marker (PRLR, 538 bp). This analysis showed that the subfamily Gerrhosaurinae consists of five major clades which we interpret as representing five genera. The genera Tetradactylus and Cordylosaurus were each recovered as monophyletic, but Gerrhosaurus as currently conceived is paraphyletic, consisting of three distinct genus-level assemblages. The two clades consisting of Gerrhosaurus major Duméril, 1851 and Gerrhosaurus validus Smith, 1849 are both described here as new genera, namely Broadleysaurus Bates & Tolley gen. nov. and Matobosaurus Bates & Tolley gen. nov., respectively. Two subspecies of 'Gerrhosaurus major' that were historically separated on the basis of differences in colour pattern are not reciprocally monophyletic, so Gerrhosaurus bottegoi Del Prato, 1895 is relegated to the synonomy of Broadleysaurus major (Duméril, 1851) comb. nov., which is rendered monotypic. Gerrhosaurus validus maltzahni De Grys, 1938 is genetically and morphologically well differentiated from G. v. validus and the two taxa also occur in allopatry. We therefore re-instate the former as Matobosaurus maltzahni (De Grys, 1938) comb. nov., rendering Matobosaurus validus (Smith, 1849) comb. nov. a monotypic species. Our analysis also showed that Gerrhosaurus sensu stricto comprises two major subclades, one consisting of Gerrhosaurus typicus (Smith, 1837) + Gerrhosaurus skoogi Andersson, 1916, and the other containing the remaining species. In this latter subclade we show that west-Central African Gerrhosaurus nigrolineatus Hallowell, 1857 is most closely related to Gerrhosaurus auritus Boettger, 1887 rather than to G. nigrolineatus from East and Southern Africa. The west-Central African clade of G. nigrolineatus differs from the East and Southern African clade by a p-distance of 13.0% (ND2) and 6.9% (16S), and can be differentiated morphologically. We accordingly apply the name Gerrhosaurus intermedius Lönnberg, 1907 comb. nov. to populations from Kenya, Uganda, Rwanda, Tanzania, Malawi, Mozambique, Zimbabwe and South Africa previously identified under the name G. nigrolineatus. Our analysis also confirms that Gerrhosaurus bulsi Laurent, 1954 is a distinct species and sister taxon to a clade containing G. nigrolineatus, G. auritus and G. intermedius. The latter four taxa form a closely-related 'G. nigrolineatus species complex' with a widespread distribution in Africa. Most closely related to this complex of species is Gerrhosaurus flavigularis Wiegmann, 1828 which has an extensive range in East and Southern Africa, and displays genetic substructure which requires further investigation. The status of Gerrhosaurus multilineatus Bocage, 1866, and Angolan populations referred to G. nigrolineatus, remains problematic.
authors.affiliation: Department of Herpetology, National Museum, P.O. Box 266, Bloemfontein 9300, South Africa; Email: herp@nasmus.co.za.
authors.forename: Michael F
authors.initials: MF
authors.lastname: Bates
authors.affiliation: Applied Biodiversity Research Division, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa; and Department of Botany and Zoology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa; Email: K.Tolley@sanbi.org.za.
authors.forename: Krystal A
authors.initials: KA
authors.lastname: Tolley
authors.affiliation: Applied Biodiversity Research Division, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa; and Department of Botany and Zoology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa; Email: S.Edwards@sanbi.org.za.
authors.forename: Shelley
authors.initials: S
authors.lastname: Edwards
authors.affiliation: Applied Biodiversity Research Division, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa; Email: Z.Davids@sanbi.org.za.
authors.forename: Zoë
authors.initials: Z
authors.lastname: Davids
authors.affiliation: Applied Biodiversity Research Division, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa; and Department of Conservation Ecology and Entomology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa; Email: jessica.m.dasilva@gmail.com.
authors.forename: Jessica M
authors.initials: JM
authors.lastname: Da Silva
authors.affiliation: Department of Zoology, Nelson Mandela Metropolitan University, P.O. Box 77000, Port Elizabeth 6031, South Africa and Department of Herpetology, Port Elizabeth Museum, P.O. Box 13147, Humewood 6013, South Africa.; Email: wrbranch@bayworld.co.za.
authors.forename: William R
authors.initials: WR
authors.lastname: Branch
mesh_terms: D000349:Africa; D063147:Animal Distribution; D000825:Animal Structures / Q000033:anatomy & histology; D000818:Animals; D017753:Ecosystem; D005260:Female; D008116:Lizards / Q000033:anatomy & histology / Q000145:classification* / Q000235:genetics; D008297:Male; D008969:Molecular Sequence Data; D010802:Phylogeny