Gerrhosaurus Wiegmann, 1828

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 : 230-235

Genus Gerrhosaurus Wiegmann, 1828

Gerrhosaurus auritus Boettger, 1887 KALAHARI PLATED LIZARD

Gerrhosaurus auritus Boettger 1887:148 , pl. V, figs. 3a–e. Holotype: SMF 13947 (formerly SMF 6120a) (collector H. Schinz) fide Boettger (1893) and Mertens (1922, 1967). Type locality: “Ondonga in Ovamboland” Namibia.

Gerrhosaurus auritus : Monard (1937b:78).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from Angola, Botswana, the Democratic Republic of Congo and Namibia.

Ocurrences in Angola (Map 187): The species occurs in northeastern Angola. Lunda Sul: “Lunda” [-10.96667, 20.06667] ( Monard 1937b:78)

Taxonomic and distributional notes: Monard considered one specimen from “Lunda” to be possibly referable to Gerrhosaurus multilineatus Bocage, 1866 , although Loveridge (1942) reject- ed this interpretation and treated this taxon as a subspecies of Gerrhosaurus nigrolineatus Hallowell, 1857 . Subsequent authors, including FitzSimons (1943), Mertens (1971) and de Witte (1953) have considered G. auritus a full species. However, de Witte’s specimens were, in fact, referable to G. bulsi Laurent, 1954 ( Bates et al. 2013), a species described by Laurent (1954) first as a subspecies of auritus . Broadley (1971) treated auritus as a subspecies of G. multilineatus and this has subsequently been followed by Bates et al. (2014). There has been much confusion regarding the applicability of names in Gerrhosaurus due to the lack of fresh material of Angolan

Gerrhosaurus bulsi Laurent, 1954 LAURENT’S PLATED LIZARD

Gerrhosaurus auritus bulsi Laurent 1954a:64 . Holotype: MD Ang. 2127 (collector A. Barros Machado). Type locality: “Dundo,” Lunda Norte Province, Angola.

Gerrhosaurus nigrolineatus nigrolineatus : Laurent (1950a:12).

Gerrhosaurus bulsi : Laurent (1964a:50), Branch and Conradie (2015:200).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from Angola, the southern Democratic Republic of Congo, and Zambia.

Ocurrences in Angola (Map 188): The species occurs in northeastern and central regions of Angola. Lunda Norte: “Dundo”

[-7.36667, 20.83333] ( Laurent 1950a:12,

1954a:64, 1964a:50); “Luachimo village”

[-7.38333, 20.83333] ( Bates et al. 2013:469

[ Table 1]); “ Lake Carumbo base camp”

[-7.74422, 19.95522] ( Bates et al. 2013:469

[ Table 1]); “Carumbo” [-7.74422, 19.95467]

(Branch and Conradie 2015:200). Lunda Sul:

“ Poste de Cacolo , Alto Cuílo ” [-10.15000,

19.28333] ( Laurent 1964a:51). Moxico: “environs du lac Calundo” [-11.80000, 20.86667]

( Laurent 1964a:50).

Taxonomic and distributional notes: de

Witte’s (1943) specimens of G. auritus

Boettger, 1887, are, in fact, referable to this

MAP 188. Distribution of Gerrhosaurus bulsi in Angola. species ( Bates et al. 2013). For some time

G. bulsi was treated as a synonym of the poorly known Gerrhosaurus multilineatus Bocage, 1866 (Broadley 1971), although it is currently regarded as valid (Broadley and Cotterill 2004; Broadley

2007). According to Bates et al. (2013) G. bulsi is a distinct species that is sister to the clade ( Gerrhosaurus intermedius [ G. auritus , G. nigrolineatus ]).

Gerrhosaurus multilineatus Bocage, 1866 KWANZA KEELED PLATED LIZARD (Endemic)

Gerrhosaurus multilineatus Bocage 1866a:44 . Holotype: MBL specimen number unknown (collector F.A.P. Bayão), destroyed by fire 18 March 1978. Type locality: “ Duque de Bragança ” [= Calandula], Malanje Province, Angola.

Gerrhosaurus multilineatus : Bocage (1866b:61, 1867b:221), Peters (1881:147).

Gerrhosaurus bulsi : Ceríaco et al. (2016b:63).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 189): Published records exist for this species from central and western Angola. Malanje: “Duque de

Bragança ” [-9.10000, 15.95000] ( Bocage

1866a:44, 1866b:61); “ Malanje ” [-9.55000,

16.35000] ( Peters 1881:147); “Cangandala

National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:63). Benguela: “Catumbella”

[-12.43333, 13.55000] ( Bocage 1867b:221);

“Dombe” [-12.95000, 13.10000] ( Bocage

1867b:221).

Taxonomic and distributional notes:

Bocage (1866b) remarked that Gerrhosaurus multilineatus might be a well-characterized variety of G. nigrolineatus Hallowell, 1857 ,

however he considered the two species distinct based on color pattern. Loveridge (1942) and

FitzSimons (1943) subsequently relegated MAP 189. Distribution of Gerrhosaurus multilineatus in Angola.

G. multilineatus to the synonymy of G. nigrolineatus , the former considering it a color variant ( Loveridge 1942). Haagner et al. (2000) stated that Broadley (unpublished) considered G. multilineatus to be based on a hybrid specimen and that the name would, therefore, be unavailable. However, according to Article 17.2 of the Code ( ICZN 1999), even if the specimen was a hybrid, the name would still be available ( Bates et al. 2013). Currently this species is recognized as valid despite the obvious taxonomic confusion, the loss of the original type series to fire at the Museu Bocage and the lack of museum material, especially from Angola ( Bates et al. 2013). The status of Gerrhosaurus multilineatus and the assignment of Angolan populations referred to Gerrhosaurus nigrolineatus (e.g., Boulenger 1887, 1905; Ferreira 1900a, 1903, 1904; Angel 1923; Monard 1937b; Mertens 1938a; Hellmich 1957a, 1957b; Manaças 1963; Parker 1936; Schmidt 1933; Laurent 1954a) remains problematic and resolution must await the collection of material from the type locality for molecular analysis, and a detailed morphological evaluation of the complex ( Bates et al. 2013). Ceríaco et al. (2016b) referred to the specimen collected in Cangandala National Park (Malanje Province) as G. bulsi , but ongoing molecular works reveal that it shall be considered true multilineatus .

Gerrhosaurus cf. nigrolineatus Hallowell, 1857 BLACK-LINED PLATED LIZARD

Gerrhossaurus nigro-lineatus ( Hallowell 1857:49). Syntypes: ANSP 3729 , 8825 (collector H.A. Ford). Type locality: “Gaboon” [= Gabon] .

Gerrhosaurus nigrolineatus : Bocage (1866a:43; 1870:68, 1887c:210, 1895a:35, 1896a:111), Peters (1877a:613), Boulenger (1887:122, 1905:111), Ferreira (1900a:49, 1903:15, 1904:117), Angel (1923:159), Monard (1937b:78), Laurent (1954a:64), Grillitsch et al. (1996:26), Largen and Parker (2004:16), Ceríaco et al. (2014b:671), Ceríaco et al. (2016a:59).

Gerrhosaurus flavigularis nigrolineatus : Schmidt (1919:523,1933:11), Mertens (1926:152, 1938a:435), Loveridge (1936a:65), Parker (1936:133).

Gerrhosaurus nigrolineatus ahlefeldti Hellmich and Schmelcher 1956:203 , fig. 1. Holotype: ZMH R02563 (formerly ZMH 806 ) (collector G.A. von Maydell) fide Hallermann (1998). Type locality: “Piri- Dembos, Roca Nova Douro” [= Piri-Dembos, Roça Novo Douro, Fazenda Novo Minho], Kwanza Norte Province, Angola.

Gerrhosaurus nigrolineatus nigrolineatus : Hellmich (1957a:54, 1957b:58), Manaças (1963:236).

Gerrhosaurus nigrolineatus ahlefeldti : Hallermann (1998:208), Frazen and Glaw (2007:222), Böhme (2014:152).

Gerrhosaurus cf. nigrolineatus : Conradie et al. (2016:24) .

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from a large distribution range, from Gabon and the lower Congo eastwards through southern Democratic Republic of the Congo eastern to Kenya, then southwards as far as northern Namibia, northern Botswana and north-eastern South Africa. Ocurrences in Angola (Map 190): The species is known from northeastern Angola.

Cabinda: “Chinchoxo” [-5.10000, 12.10000]

( Peters 1877a:613); “Molembo” [-5.33333,

12.20000] ( Bocage 1895a:35); “ Cabinda ”

[-5.55000, 12.18333] ( Bocage 1866a:43,

1895a:35). Lunda Norte: “Dundo près du

Musée” [-7.36667, 20.83333] ( Laurent

1954a:64). Zaire: “St. Antonio [do Congo]”

[-6.13333, 12.36667] ( Schmidt 1919:523-524).

Bengo: “Ambriz” [-7.844312, 13.106493]

( Boulenger 1887:122; Bocage 1895a:35; Grillitsch et al. 1996:26); “Cabiri” [-8.91667,

13.66667] ( Ferreira 1904:117). Luanda :

“ Luanda ” [-8.83333, 13.26667] (Largen and

Parker 2004:16). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich and

Schmelcher 1956:203; Hellmich 1957b:58; MAP 190. Distribution of Gerrhosaurus cf. nigrolineatus in Angola.

Hallermann 1998:208; Frazen and Glaw

2007:222; Böhme 2014:152); “Mucoso” [-9.53333, 14.65000] ( Hellmich 1957a:54); “Dondo” [-9.68333, 14.43333] ( Bocage 1895a:35). Kwanza Sul: “Libolo/Luati” [-9.98333, 14.90000] ( Hellmich 1957b:58), “Quirimbo” [-10.68333, 14.26667] ( Parker 1936:133); “Congulu” [-10.86667, 14.28333] ( Parker 1936:133). Malanje: “Duque de Bragança ” [-9.10000, 15.95000] ( Bocage 1895a:35; Ferreira 1903:15; Boulenger 1905:111); “ Malanje ” [-9.55000, 16.35000] ( Bocage 1895a:35); “Capanda” [-9.72841, 15.34585] (Ceríaco et al. 2014b:671); “Pungo-Andongo” [-9.66667, 15.58333] ( Boulenger 1905:111). Moxico: “ Lago Dilolo” [-11.50000, 22.01667] ( Manaças 1963:236); “ Lago Cameia” [-11.71667, 20.80000] ( Manaças 1963:236). Bié: “Gauca”

[-11.18333, 17.45000] ( Schmidt 1933:11); “Chitau” [-11.43333, 17.15000] ( Schmidt 1933:11). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937b:78); “Galanga” [-12.06667, 15.15000] ( Bocage 1895a:35; “Santo Amaro” [-12.70000, 15.85000] ( Monard 1937b:78); “Cuma” [-12.86667, 15.06667] (Loverdige 1936a:65). Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1887c:210); “Catumbella” [-12.43333, 13.55000] ( Bocage 1887c:210); “Quissange” [-12.43333, 14.05000] ( Bocage 1887c:210, 1895a:35); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:35); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:54); “Cubal” [-13.03333, 14.25000] (Mertens 1938:435); “ Benguela ” [-12.58333, 13.41667] ( Boulenger 1887:122; Bocage 1895a:35); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:111); “Catengue” [-13.03333, 13.73333] ( Mertens 1926:152). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:35); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937b:78); “Kapelongo” [-14.88333, 15.08333] ( Monard 1937b:78); “Quillengues” [-14.06667, 15.08333] ( Bocage 1895a:35); “Huilla” [-15.05000, 13.55000] ( Bocage 1895a:35); “Kampulu” [-15.21667, 16.11667] ( Monard 1937b:78); “Mulondo” [-15.63333, 15.20000] ( Monard 1937b:78). Namibe: “Capangombe” [-15.10000, 13.15000] ( Bocage 1895a:35; Ceríaco et al. 2016a:59). Cunene: “riv. Mbalé ” [-15.16667, 16.75000] ( Monard 1937b:78); “Chimporo” [-16.03333, 17.15000] ( Monard 1931:109, 1937b:148). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] ( Monard 1937b:78); “Kwito region, tributary of Kubango” ( Angel 1923:159); “Cubango basin (6a)” [-14.67155, 17.73525] ( Conradie et al. 2016:8-9, 24); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9- 10, 24); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016:9-10, 24); “Cubango basin (46)” [-14.58981, 16.907389] ( Conradie et al. 2016:9, 12, 24). Undetermined locality: “Carangigo” ( Bocage 1895a:35); “Between, Bihé and Quilenges” ( Boulenger 1905:111); “without precise location” ( Bocage 1870:68; Ferreira 1900:49).

Taxonomic and distributional notes: Bates et al. (2013) suggested that G. nigrolineatus was not monophyletic and resurrected G. intermedius Lönnberg, 1907 for eastern populations ( Kenya, Uganda, Rwanda, Tanzania, Malawi, Mozambique, Zimbabwe and South Africa). However, accurate determination of geographical boundaries for nigrolineatus and intermedius , especially in Central Africa ( Angola, Democratic Republic of Congo, Zambia, northern Botswana and northern Namibia) will require additional sampling, as well as additional morphological analysis of specimens from throughout their extensive ranges. Bates et al. (2013) questioned the assignment of Angolan Gerrhosaurus multilineatus Bocage, 1866 specimens to G. nigrolineatus (e.g., Boulenger 1887, 1905; Ferreira 1900, 1903, 1904; Angel 1923; Monard 1937b; Mertens 1938; Hellmich 1957a, 1957b; Manaças 1963; Parker 1936; Schmidt 1933; Laurent 1954), and their relationship

(see G. multilineatus account). The status of the G. nigrolineatus complex in Angola is currently under investigation (M. Bates, pers. comm.) and the records plotted here should be considered as only tentatively assigned to this species. Gerrhosaurus nigrolineatus ahlefeldti (Hellmich and Schmelcher, 1956) is now considered a synonym of G. nigrolineatus , the holotype and four paratypes are present in the Zoological Museum in Hamburg and two paratypes are at the Zoologische Staatssammlung, München ( Hallermann 1998; Franzen and Glaw 2007).

Gerrhosaurus skoogi Andersson, 1916 DESERT PLATED LIZARD

Gerrhosaurus skoogi Andersson 1916:10 , fig. 2. Holotype: GNM 1387 (donor Lt. D. Fortunato). Type locality: “ Port Alexander , Portuguese West Africa ” [= Tômbua], Namibe Province, Angola.

Angolosaurus skoogi : Loveridge (1942:495), FitzSimons (1953:215), Branch (1998:177), Ceríaco et al. (2016a:39, 59).

Global conservation status (IUCN): Least Concern.

Global distribution: A deserticolous species that may form loosely-structured colonies restricted to the Namib Desert, in southwestern

Angola and northwestern Namibia.

Ocurrences in Angola (Map 191): The species occurs in southwestern Angola and is restricted to the Namib Desert. Namibe: “Porto

Alexandre, between Mossamedes and the mouth of the Kunene river” [-15.80000,

11.83333] ( Andersson 1916; Loveridge

1942:495; FitzSimons 1953:215; Ceríaco et al.

2016a:59); “ Praia do Navio coastal dunes, ca

12 km SSW of Namibe ” [-16.27233, 11.83164]

(Ceríaco et al. 2016a:39).

Taxonomic and distributional notes:

The Genus Angolosaurus was established by

FitzSimons (1953) to accommodate the species

Gerrhosaurus skoogi Andersson, 1916 . Lang

(1991) considered Angolosaurus skoogi the probable sister taxon to mainland African ger- MAP 191. Distribution of Gerrhosaurus skoogi in Angola.

rhosaurids (Lang et al. 2003) and Nance (2007)

supported a position as the most basal member of Gerrhosauridae based on morphology. Howev- er, Lamb et al. (2003) showed, using mitochondrial DNA data, that the species is embedded within Gerrhosaurus and this was confirmed by a multilocus phylogeny by Lamb and Bauer (2013). Consequently Angolosaurus is a synonym of Gerrhosaurus (Adolphs and Bates 2010) .