Hemidactylus Oken, 1817

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 : 182-188

Genus Hemidactylus Oken, 1817

Hemidactylus angulatus complex Hallowell, 1852 AFRICAN TROPICAL GECKO

Hemidactylus angulatus Hallowell 1852a:63 , fig. Holotype: ANSP 7431 (donated by H. Ford). Type locality:

“West coast of Africa” [= Gabon] by implication fide Loveridge (1947). Hemidactylus brookii angulatus : Laurent (1964a:29). Hemidactylus brooki : Spawls et al. (2004:86). Hemidactylus angulatus : Carranza and Arnold (2006:539), Bauer et al. (2006a:8), Trape et al. (2012:222),

Trape et al. (2012:222).

Hemidactylus bayonii Bocage, 1893 BAYÃO’S GECKO (Endemic)

Hemidactylus Bayonii Bocage 1893:116 . Holotype: MBL specimen number not known (collector F.A.P. Bayão), destroyed by fire 18 March 1978. Type locality: “Dondo, sur les bords du Quanza” [= Dondo, near Quanza river edges], Kwanza Norte, Angola.

Hemidactylus bayonii : Bocage (1895a:13, 1897a:193), Frade (1963:252), Laurent (1964a:30).

Hemidactylus brookii angulatus : Loveridge (1947:139, 1957:184).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known only from Angola.

Ocurrences in Angola (Map 130): The species is only known from western regions of the Kwanza watershed. Kwanza Norte: “Dondo” [-9.68333, 14.43333] ( Bocage 1893:116, 1895a:13, 1897a:193). Kwanza Sul: “ 31 km au nord-est de Novo Redondo” [-11.00000, 14.05000] ( Laurent 1964a:30).

Taxonomic and distributional notes: Loveridge (1947, 1957) considered the type specimen of Hemidactylus bayonii referable to H. angulatus . For some time this species was only known from the type locality, although Laurent (1964a) identified two individuals as Hemidactylus bayonii from “ Kwanza Sul, 31 km from Novo Redondo”, not far from the type locality. Laurent (1964a) also provided a discussion about Loveridge’s (1947) mistaken synonymy of H. bayonii

Hemidactylus benguellensis Bocage, 1893 BENGUELA GECKO (Endemic)

Hemidactylus benguellensis Bocage 1893:115 . Syntypes: MBL specimen numbers not known (collector J.A. d’Anchieta)”, destroyed by fire 18 March 1978. Type locality: “Cahata” [= Caota ], Benguela Province, Angola.

Hemidactylus benguellensis : Bocage (1895a:12, 1897a:193), Monard (1937b:52), Frade (1963:252).

Hemidactylus mabouia : Loveridge (1947:170, 1957:185).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from Angola.

Ocurrences in Angola (Map 131): The species is only known from southwestern regions of the country. Benguela: “Cahata”

[-12.35000, 14.81667] ( Bocage 1893:115,

1895a:12, 1897a:115; Loveridge 1947:170,

1957:185); “Ebanga” [-12.73333, 14.73333]

( Monard 1937b:52).

Taxonomic and distributional notes:

This species is generally regarded as a synonym of Hemidactylus mabouia ( Moreau De

Jonnés, 1818) ( Loveridge 1947, 1957; Uetz and Hošek 2017). However, H. mabouia is itself a complex of species (Bauer et al.,

unpublished) and we tentatively maintain the validity of H. benguellensis until both it and

H. mabouia can be thoroughly revised. MAP 131. Distribution of Hemidactylus benguellensis in Angola.

Hemidactylus longicephalus Bocage, 1873 LONG-HEADED TROPICAL GECKO Hemidactylus longicephalus Bocage 1873b:210 . Syntypes: MBL specimen numbers not known (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “l’intérieur de Mossamedes (Capangombe)” and “Catumbella, près Benguella” [= Capangombe (Namibe Province) and Catumbella (Benguela Province)], Angola.

Hemidactylus platycephalus : Bocage (1866a:42, 1866b:60, 1870:68; 1873b:209).

Hemidactylus bocagii : Boulenger (1885:125), Bocage (1895a:11, 1897a:193), Ferreira (1904:117). Hemidactylus longicephalus : Bocage (1887a:178), Schmidt (1933:4), Parker (1936:128), Hellmich (1957b:49), Manaças (1963:227), Loveridge (1947:187), Laurent (1964a:30), Ceríaco et al. (2014b:671), Ceríaco et al. (2016a:54).

Hemidactylus bocagei : Ferreira (1906:170).

Hemidactylus mabouia : Loveridge (1947:167).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from West Africa from Cameroon to Angola south into northern Namibia.

Ocurrences in Angola (Map 132): The species is known from throughout the western regions of the country. Cabinda: “Landana”

[-5.21667, 12.15000] ( Loveridge 1947:189);

“ Cabinda ” [-5.55000, 12.18333] ( Bocage

1866a:42, 1866b:60, 1895a:11, 1897a:193).

Zaire: “ S. Salvador do Congo ” [-6.26667,

14.23333] ( Bocage 1887a:178, 1895a:11;

Loveridge 1947:189). Uíge: “Fazenda Otília-

Encoge, Quitexe” [-7.55000, 15.03333]

( Manaças 1963:227). Luanda: “Loanda”

[-8.83333, 13.26667] ( Bocage 1866a:42,

1866b:60). Bengo: “Ambriz” [-7.844312,

13.106493] ( Boulenger 1885:125; Bocage

1895a:11; Loveridge 1947:189); “Minas de

Quizambil, Dande” [-8.31667, 13.73333]

( Manaças 1963:227); “Cunga” [-9.23333,

13.76667] ( Ferreira 1904:117; Loveridge MAP 132. Distribution of Hemidactylus longicephalus in Angola.

1947:189). Kwanza Norte: “Piri-Dembos”

[-8.56667, 14.50000] ( Hellmich 1957b:49); “Dondo” [-9.68333, 14.43333] ( Bocage 1873b:209). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] ( Parker 1936:128; Loveridge 1947:189); “Congulu” [-10.86667, 14.28333] ( Parker 1936:128; Loveridge 1947:189); “Gumba” [-11.26667, 14.28333] ( Ferreira 1904:117; Loveridge 1947:189). Malanje: “Duque de Bragança ” [-9.10000, 15.95000] ( Bocage 1895a:11, 1897a:193; Loveridge 1947:189); “Pungo-Andongo” [-9.66667, 15.58333] ( Boulenger 1885:125; Bocage 1895a:11; Loveridge 1947:189); “Capanda” [-9.72841, 15.34585] (Ceríaco et al. 2014b:671). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1933:4; Loveridge 1947:189). Benguela: “Catumbella” [-12.43333, 13.55000] ( Bocage 1873b:209, 1895a:11, 1897a:193; Loveridge 1947:189). Huíla: “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] ( Laurent 1964a:30). Namibe: “Capangombe” [-15.10000, 13.15000] ( Bocage 1873b:209, 1895a:11, 1897a:193; Loveridge 1947:189; Ceríaco et al. 2016a:54); “ Rio Coroca” [-15.78333, 12.06667] ( Bocage, 1895a:11, 1897a:193; Loveridge 1947:189; Ceríaco et al. 2016a: 54). Undetermined Locality: “Carangigo” ( Boulenger 1885:125; Bocage 1895a:11; Loveridge 1947:189); “Cuanza River ” ( Loveridge 1947:189).

Taxonomic and distributional notes: In the original description Bocage identified the species as Hemidactylus platycephalus , however, at the end of the description he mentioned that he had provisionally registered the species in the Museum catalogue under the name Hemidactylus longicephalus . Perret (1975) believed that H. longicephalus Bocage, 1873 was a junior synonym of H. muriceus Peters, 1870 (see below), but he had been unable to see the type of the former to confirm this. Hemidactylus bocagii Boulenger 1885 was recognized as valid for Angolan specimens by Bocage (1895a, 1897a) and Ferreira (1904, 1906), although this name was synonymized by Loveridge (1947). The taxonomy of H. longicephalus is unstable and several names that are currently placed in its synonymy require further investigation. This species is broadly sympatric with the rather similar Hemidactylus mabouia ( Moreau De Jonnès, 1818) , throughout much of its range in Angola (Ceríaco et al. 2014a).

Hemidactylus mabouia ( Moreau De Jonnès, 1818) TROPICAL HOUSE GECKO

Gecko Mabouia Moreau de Jonnès, 1818:138 . Holotype: MHNP 6573. Type locality: “en Amérique, dans les contrées continentales qui avoisinent au midi l’archipel des Antilles, et qu’il est également répandu dans les îles même de l’archipel, depuis la Trinité jusqu’a la Jamaïque. “ Restricted to “ St. Vincent ” [Lesser Antilles] by Stejneger (1904:600).

Hemidactylus mabouia : Peters (1877a:612), Boulenger (1885:122, 1905:110), Bocage (1895a:10), Ferreira (1904:117, 1906:170), Parker (1936:128), Loveridge (1947:167, 1957:185), Hellmich (1957a:34), Laurent (1950a:12, 1954a:63, 1964a:29), Kluge (1969:28), Spawls et al. (2004:88), Bates et al. (2014:114), Branch and Conradie (2015:200), Ceríaco et al. (2016b:59).

Hemidactylus mabouia mabouia : Chirio and LeBreton (2007:118).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from almost all sub-Saharan Africa regions, although it is absent from more arid and temperate areas except under anthropogenic conditions. It is also currently found in North, Central and South America and in the Caribbean, where it is an anthropophilic invasive ( Kraus 2009).

Ocurrences in Angola (Map 133): The species occurs mainly in northern Angola, but also along the coast, including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] ( Peters 1877a:612; Bocage 1895a:10;

Loveridge 1947:179); “ Cabinda ” [-5.55000,

12.18333] ( Bocage 1895a:10; Loveridge

1947:179). Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1950a:12, 1964a:29);

“Carumbo, Lucapa” [-8.42278, 20.73917]

(Branch and Conradie 2015:200). Luanda:

“Loanda” [-8.83333, 13.26667] ( Bocage

1895a:10; Loveridge 1947:179; Hellmich

1957a:34). Bengo: “Cabiri” [-8.91667,

13.66667] ( Ferreira 1903:117; Loveridge

1947:179); “Catete” [-9.11667, 13.70000]

( Ferreira 1903:117; Loveridge 1947:179).

Kwanza Norte: “Cambondo” [-9.15963,

14.65827] (Ferreira:1906:170; Loveridge

1947:179); “Mucoso bei Dondo” [-9.53333,

14.65000] ( Hellmich 1957a:34); “Dondo”

[ -9.68333, 14.43333] ( Bocage 1895a:10; Love- MAP 133. Distribution of Hemidactylus mabouia in Angola.

ridge 1947:179). Malanje: “Duque de Bragança ” [-9.10000, 15.95000] ( Bocage 1895a:10; Loveridge 1947:179); “Pungo-Andongo” [-9.66667, 15.58333] ( Boulenger 1905:110; Loveridge 1947:179); “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:59). Benguela: “Lobito” [-12.35000, 13.55000] ( Parker 1936:128; Loveridge 1947:179; Laurent 1954a:63); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:34). Undetermined Locality: “Carangigo” ( Boulenger 1885:122; Loveridge 1947:179); “Serra de Stombe” ( Ferreira 1903:117; Loveridge 1947:179).

Taxonomic and distributional notes: The type locality of H. mabouia was initially restrict- ed to “Antilles” by A. Duméril in Duméril and Duméril (1851) and then more precisely to “ St. Vincent ” by Stejneger (1904). However, as noted by Kluge (1969), the holotype, which he believed was actually collected in Catragena, Colombia, is not conspecific with the gecko now typically regarded as H. mabouia , and would appear to be a specimen of H. angulatus (Carranza and Arnold 2006) . Kluge recommended the maintenance of prevailing usage of the name. Hemidactylus mabouia as currently recognized includes several synonyms. However, it is known to be part of a complex of many species and the taxonomy of the complex is in flux. In Angola, H. mabouia is broadly sympatric with the rather similar Hemidactylus longicephalus Bocage, 1873 throughout much of its range (Ceríaco et al. 2014b). Kluge (1969) mapped several Angolan localities, but did not provide specific localities.

Hemidactylus cf. muriceus Peters, 1870 GUINEAN SPINY TROPICAL GECKO

Hemidactylus muriceus Peters 1870:641 . Holotype: ZMB 6921 (collector R. Schomburgk). Type locality: “Keta (Guinea)” [= Keta, Ghana].

Hemidactylus muriceus : Peters (1881:147), Bocage (1895a:13), Henle and Böhme (2003:33), Bauer et al. (2006a:10).

Hemidactylus cf. muriceus : Ceríaco et al. (2016b:59).

Global conservation status (IUCN): Not Evaluated.

Global distribution: This species occurs in West Africa forests, from Ghana to Cameroon, east to the Central African Republic and south to northern Angola.

Ocurrences in Angola (Map 134): The species is known from northcentral Angola.

Malanje: “ Cangandala National Park ”

[-9.84606, 16.72233] (Ceríaco et al.

2016b:59). Undetermined Locality: “Cuango

= Quango” ( Peters 1881:147; Bocage 1895a:

13) (Malanje Province impossible to georeference: See History Section for more detailed information).

Taxonomic and distributional notes:

The type was incorrectly cited as ZMB 69201

by Perret (1975). According to Perret (1975),

Henle and Böhme (2003) and Bauer et al.

(2006a) Hemidactylus muriceus Peters, 1870

has frequently been confused with other species. Perret (1975) attributed the confusion to Tornier’s (1902) use of the name for

H. intestinalis Werner, 1897 , which he consid- MAP 134. Distribution of Hemidactylus cf. muriceus in Angola.

ered as senior synonym of H. ansorgii

Boulenger, 1901. However, Henle and Böhme (2003) have argued that H. intestinalis is indeed a synonym of H. muriceus , and that H. ansorgii is the name applicable to another species. Perret (1975) also believed that H. longicephalus Bocage, 1873 was a junior synonym of H. muriceus , but he had been unable to see the type of the former to confirm this. Because of those frequent misidentifications, literature records for the species are in many cases unreliable. The identity of Angolan specimens is unclear but they are probably not referable to H. muriceus sensu stricto.