Cordylus Laurenti, 1768

Marques, Mariana P., Ceríaco, Luis M. P., Blackburn, David C. & Bauer, Aaron M., 2018, Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola Atlas of Historical and Bibliographic Records (1840 - 2017), Proceedings of the California Academy of Sciences 65, pp. 1-501 : 227-229

Genus Cordylus Laurenti, 1768

Cordylus angolensis (Bocage, 1895) ANGOLAN GIRDLED LIZARD (Endemic)

Zonurus angolensis Bocage 1895a:25 . Holotype: MBL specimen number unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Caconda,” Huíla Province, Angola. Zonurus cordylus : Boulenger (1897:277). Zonurus angolensis : Monard (1937b:61), Frade (1963:252). Cordylus cordylus angolensis : Loveridge (1944b:38). Cordylus tropidosternum tropidosternum : Broadley (1971:22) Cordylus angolensis : Broadley and Branch (2002:10), Branch et al. (2005:131), Stanley et al. (2011:67),

Greenbaum et al. (2012:36), Nielsen and Colston (2014:167), Stanley et al. (2016:23).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from Angola.

Ocurrences in Angola (Map 183): The species is known from the type locality “Caconda” in Huíla Province but its distribution range reaches the southern regions of Kwanza Sul Province. Kwanza Sul: “Mombolo” [-11.91667, 14.85000] (Greenbaum et al. 2012:36). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:24; Monard 1937b:61; Loveridge 1944b:39).

Taxonomic and distributional notes: In the description of Cordylus angolensis Bocage (1895a) expressed some doubts about the true identity of C. angolensis , leading him to name it in a “provisional manner.” The species was not cited in any further Bocage works after the description. With the destruction of the type specimen in the Lisbon museum fire, the taxonomic identity of the species became even MAP 183. Distribution of Cordylus angolensis in

Angola. more problematic. Broadley (1971b) considered it a synonym of Cordylus tropidosternum (Cope, 1869) . Loveridge (1944b) and Branch et al. (2005) noted that Bocage’s type description of C. angolensis lacked detail, although it is now considered a valid species endemic to central Angola (Broadley and Branch 2002; Stanley et al. 2011; Nielsen and Colston 2014; Stanley et al. 2016). Greenbaum et al. (2012) found three Cordylu s specimens in the collections of the American Museum of Natural History assignable to C. angolensis from “Mombolo”, approximately 190 km north-northwest of the type locality. These specimens agree with the original holotype description in some measurments and coloration,

although they differ in the scale counts of longitudinal rows of ventral scales (Greenbaum et al. 2012). Angolan Cordylus are currently being revised and the systematic position of C. angolensis will hopefully be resolved. Cordylus angolensis was described within the account of Cordylus cordylus ( Bocage 1895a). The two specimens treated as C. cordylus were from the Sociedade de Geografia de Lisboa, Portugal, and had only the locality “ Angola.” Monard (1937b) considered these specimens to be attributable to Cordylus vittifer ( Reichenow, 1887) , although Loveridge (1944b) considered this opinion untenable and considered them assignable to C. angolensis . The identity of these specimens, which have recently been re-discovered remains uncertain, but is under study.

Cordylus machadoi Laurent, 1964 MACHADO’S GIRDLED LIZARD

Cordylus vittifer machadoi Laurent 1964a:49 , fig. 12. Holotype: MD 1840-1 (collector A. Barros Machado). Type locality: “Leba, Humpata, environs de Sá da Bandeira, Alt. 1800 m, Huíla” [= Serra da Leba , Lubango], Huíla Province, Angola.

Cordylus vittifer machadoi : Broadley (1971b:22)

Cordylus machadoi : Branch (1998:195), Stanley et al. (2011:67), Greenbaum et al. (2012:23), Bates et al. (2014:192), Nielsen and Colston (2014:171), Stanley et al. (2016:202).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from southwestern Angola and northwestern Namibia.

Ocurrences in Angola (Map 184): The species is known from the type locality in

Huíla Province, however the species distribution may extend to Namibe Province. Huíla:

“Leba, Humpata, environs de Sá da Bandeira”

[-15.06667, 13.60000] ( Laurent 1964a:49);

“Humpata” [-14.96167, 13.34972];

[-15.17750, 13.32139E ( Stanley et al. 2011:57;

Stanley et al. 2016:209); “Nascente do Tchi-

viuguira” [-15.16667, 13.30000] ( Stanley et al.

2016:209).

Taxonomic and distributional notes:

The holotype of this taxon has recently been rediscovered in the collection at Dundo. The single juvenile paratype, was exchanged from this collection and is now MCZ R-74120 (fide

Stanley et al. 2016). Branch (1998) elevated

C. machadoi to full species, an action recently

MAP 184. Distribution of Cordylus machadoi in Angola. supported by recently Stanley et al. (2016) who based their conclusions on two C. machadoi specimens collected at the type locality. These were recovered as sister to all East African species of Cordylus ( Stanley et al. 2011; Greenbaum et al. 2012; Nielsen and Colston 2014). A cryptic species allied to C. machadoi , was recently discovered at lower elevation ( Stanley et al. 2016). Significant genetic variation across such a constrained geographic area suggests that its northern congener, C. angolensis (Bocage, 1895) , which has a comparatively broad geographic distribution, may itself represent a species complex ( Stanley et al. 2016).

Cordylus namakuiyus Stanley, Ceríaco, Bandeira, Valério, Bates and Branch, 2016 KAOKOVELD GIRDLED LIZARD (Endemic)

Cordylus namakuiyus ( Stanley et al. 2016:212, figs. 1, 4, 5). Holotype: CAS 254912 (collectors L. Stanley, S. de Sá, S. Bandeira, H. Valério, A. Kuhn, J. Vindum and L. Ceríaco). Type locality: “large rock outcrop near Caraculo, road from Lubango and Namibe, Namibe Province,” Angola.

Cordylus namakuiyus : Ceríaco et al. (2016a:38, 59).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 185): The species is known from Huíla and Namibe provinces in southwestern Angola. Huíla:

“ large rock outcrop near Caraculo, on the road from Lubango and Namibe ” [-15.01650,

12.64203] ( Stanley et al. 2016:212; Ceríaco et al. 2016a:38, 59). Namibe: “outskirts of Pico

Azevedo” [-15.47589, 12.46269] ( Stanley et al.

2016:212; Ceríaco et al. 2016a:38, 59); “low rock outcrop bordering the road between

Namibe and Omahua lodge” [-15.99681,

12.40683] ( Stanley et al. 2016:212; Ceríaco et al. 2016a:38, 59).

Taxonomic and distributional notes:

Stanley et al. (2016) recently described this form as a new species endemic to the arid lowlands west of the southern Angolan escarpment and sister to the geographically proximate, MAP 185. Distribution of Cordylus namakuiyus in Angola.

high-elevation species Cordylus machadoi

Laurent, 1964. All phylogenetic analyses recovered the Angolan specimens as monophyletic, forming the earliest diverging lineage of the northern Cordylu s clade ( Stanley et al. 2016). These authors analyzed the morphology of AMNH Angolan specimens identified as Cordylus cordylus ( Linnaeus, 1758) collected during the Vernay expedition in 1925 and showed that they cluster closely with C. namakuiyus . Although the Vernay specimens lack specific locality information, the expedition field notes mention that significant numbers of unidentified lizards were collected at “Pico Azevedo” and “ 100 km east of Moçâmedes,” the same areas where the eight specimens of the C. namakuiyus were collected, leading the authors to assign the Vernay material to the new species ( Stanley et al. 2016).