Pachydactylus Wiegmann, 1834

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 : 192-199

Genus Pachydactylus Wiegmann, 1834

Pachydactylus angolensis Loveridge, 1944 ANGOLAN THICK-TOED GECKO (Endemic)

Pachydactylus scutatus angolensis ( Loveridge 1944a:3). Holotype: AMNH 47874 (collectors A. Vernay, H. Lang and R. Boulton). Type locality: “Hanha, Benguela Province,” Angola.

Pachydactylus scutatus angolensis : Barbour and Loveridge (1946:165), Loveridge (1947:357), Laurent (1964a:37).

Pachydactylus angolensis : Bauer et al. (2002a:26), Bauer and Lamb (2005:116), Bauer (2010:265); Heinicke et al. (2017:12), Ceríaco et al. (2016a:25), Branch et al. (2017:163).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is only known from Angola. Presumptive specimens from Namibia have been determined to be P. parascutatus Bauer, Lamb and Branch, 2002 .

Ocurrences in Angola (Map 139): The species occurs broadly in southwestern Angola,

below the eascarpment. Benguela: “ Lobito bay ” [-12.35000, 13.55000] (Loveridge

1944:3; Bauer et al. 2002a:23); “Hanha,

Benguela ” [-12.24500, 13.70750] (Loveridge

1944:3; Barbour and Loveridge 1946:165;

Bauer et al. 2002a:23; Bauer 2010:265; Branch et al. 2017:163); “ 24 km S Benguela ”

[-12.69747, 13.25222] ( Bauer 2010:265;

Branch et al. 2017:163); “ 30 km N of Dombe

Grande” [-12.73097, 13.23024] ( Bauer

2010:265; Branch et al. 2017:163). Namibe:

“Lucira” [-13.86667, 12.53333] ( Bauer

2010:265; Ceríaco et al. 2016a:55; Branch et al. 2017:163); “San Nicolau” [-14.26667,

12.36667] ( Bauer 2010:265; Ceríaco et al.

2016a:55); “Lungo” [-14.31667, 13.20000] MAP 139. Distribution of Pachydactylus angolensis in Angola .

( Bauer 2010:265; Ceríaco et al. 2016a:55;

Branch et al. 2017:163); “Bentiaba” [-14.26667, 12.38333] ( Branch et al. 2017:163); “Saco de Giraul” [-15.06833, 12.14222] ( Bauer 2010:265; Ceríaco et al. 2016a:55; Branch et al. 2017:163); “environs de Moçâmedes (au bord de la route de Sá da Bandeira et dans le plateau littoral de la Plage das Conchas” [-15.16667, 12.15000] ( Laurent 1964a:37; Ceríaco et al. 2016a:55); “Namibe-Lubango road, 2 km E (by road) of Mangueiras, south side of the road” [-15.04361, 13.16000] (Ceríaco et al. 2016a:25; Branch et al. 2017:163).

Taxonomic and distributional notes: Bauer et al. (2002a) re-examined the types of P. s. angolensis and revised the P. scutatus group, revalidating P. angolensis as a full species, although closely allied to P. scutatus . Bauer and Lamb (2005) did not include the species in their phylogenetic analyses of southern African Pachydactylus , but they suggested that it was a member of the “northwestern clade.” This has recently been verified by Heinicke et al. (2011), who subsequently (Heinicke et al. 2017) identified a sister group relationship between P. angolensis and P. caraculicus FitzSimons, 1959 . Ongoing molecular work has identified putative species-level diversity within the species.

Pachydactylus caraculicus FitzSimons, 1959 ANGOLA BANDED THICK-TOED GECKO

Pachydactylus caraculicus FitzSimons 1959:407 . Holotype: TM 22880 (collector C. Koch). Type locality: “Caracul, S. Angola ” [= Caraculo, Namibe Province, Angola].

Pachydactylus angolensis : Laurent (1964a:37), Haacke (1970:280), Branch (1998:253), Bauer and Lamb (2005:116), Bauer et al. (2002a:12), Bauer (2010:266), Heinicke et al. (2017:12), Mashinini and Mahlangu (2013:174); Ceríaco et al. (2016a:55).

Global conservation status (IUCN): Not Evaluated.

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

Ocurrences in Angola (Map 140): The species is known from the type locality “Caraculo” and nearby areas in Namibe Province .

Namibe: “Lungo, Vila Arriaga district ,

S. Angola ” [-14.76667, 13.36667] ( FitzSimons

1959:407; Ceríaco et al. 2016a:55); “ 36 mi northwest of Mocamedes [Namibe]”

[-14.82533, 12.54417] ( Bauer 2010:266;

Ceríaco et al. 2016b:55); “Caracul, S. Angola ”

[-15.01667, 12.66667] ( FitzSimons 1959:407;

Haacke 1970:280; Mashinini and Mahlangu

2013:174; Ceríaco et al. 2016a:55); “Cima,

Giraul River district ” [-15.06667, 12.15000]

( FitzSimons 1959:407; Ceríaco et al.

2016a:55).

Taxonomic and distributional notes: In the original description, FitzSimons (1959)

considered Pachydactylus caraculicus to be MAP 140. Distribution of Pachydactylus caraculicus in Angola.

geographically and phylogenetically intermediate between Pachydactylus scutatus angolensis Loveridge, 1944 (= Pachydactylus angolensis ) and Pachydactylus scutatus scutatus Hewitt, 1927 . However, this interpretation was challenged by Laurent (1964a) who noted that P. s. angolensis occurs in sympatry with P. caraculicus in southern Angola, and therefore must be specifically distinct (Bauer et al. 2002a). Bauer and Lamb (2005) and Heinicke et al. (2011) provided phylogenetic analyses that demonstrated that P. caraculicus is part of a diverse, primarily rupicolous “northwestern clade” of Pachydactylus , and this has been verified by Heinicke et al. (2017), who identified a sister group relationship of P. caraculicus with P. angolensis Loveridge, 1944 .

Pachydactylus oreophilus complex McLachlan and Spence, 1967 KAOKOLAND ROCK GECKO

Pachydactylus oreophilus McLachlan and Spence 1967:4 , figs.1–2. Holotype: PEM 1503 /67 [now PEM R1921 ] (collectors G.R. McLachland and J.M. Spence). Type locality: “ 20 miles West of Sesfontein ,” Kunene Region, Namibia.

Pachydactylus oreophilus : Branch (1998:258), Bauer and Lamb (2005:116), Bauer (2010:266), Heinicke et al. (2011:12), Herrmann and Branch (2012:98).

Pachydactylus cf. oreophilus : Ceríaco et al. (2016a:26, 55).

Global conservation status (IUCN): Not Evaluated.

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

Ocurrences in Angola (Map 141): The species occurs in Namibe Province and in the arid southwest of Benguela Province. Benguela: “ 35 km S Dombe Grande towards Lucira”

[-13.20979, 12.98017] ( Bauer 2010:266).

Namibe: “Assuñcao” [-14.86667, 13.10000]

( Bauer 2010:266; Ceríaco et al. 2016a:55);

“Caraculo” [-15.01667, 12.66667] ( Bauer

2010:266; Ceríaco et al. 2016b:55); “ 20 km W

Virei” [-15.66667, 12.76667] ( Bauer 2010:266;

Ceríaco et al. 2016a:55); “ 6 km S Rio Coroca towards Iona” ( Bauer 2010:266; Ceríaco et al.

2016a:55); “Saiona River, 25 km NW Cainde ”

[-15.40000, 13.20000] ( Bauer 2010:266;

Ceríaco et al. 2016a:55); “Mutiambo River on road to Lucira” [-15.78333, 12.46667] ( Bauer

2010:266; Ceríaco et al. 2016a:55); “Tambor”

[ -16.13556, 12.42972] ( Bauer 2010:266; MAP 141. Distribution of Pachydactylus oreophilus in Angola.

Ceríaco et al. 2016a:55); “ 7 km from Iona towards Oncocau, Iona Reserve” [-16.85831, 12.61275] ( Bauer 2010:266; Ceríaco et al. 2016a:55); “Furnas” ( Bauer 2010:266; Ceríaco et al. 2016a:55); “Omauha Lodge” [-16.19861, 12.40087] (Ceríaco et al. 2016a:26).

Taxonomic and distributional notes: Bauer and Lamb (2005) and Heinicke et al. (2011, 2017) demonstrated that P. oreophilus belongs to the diverse, largely rupicolous “northwestern clade” of Pachydactylus . Specimens currently allocated to this taxon, in fact, represent a nonmonophyletic group of superficially similar forms (Bauer et al., unpublished), with P. oreophilus sensu stricto limited to Namibia. There are further species level differences among the Angolan populations (Bauer et al., unpublished).

Pachydactylus punctatus complex Peters, 1854 SPECKLED THICK-TOED GECKO

Pachydactylus punctatus Peters 1854:615 . Lectotype: ZMB 4799, designated by Loveridge (1947:354) by implication (see Bauer and Günther 1991:294). Type locality: “Sena und Tette” [= Sena and Tete], Mozambique, restricted to “ Tete ” by implication fide Bauer et al. (1995:55).

Pachydactylus ocellatus : Bocage (1867b:220, 1895a:16), Boulenger (1885:205, 1905:110), Frade (1963:253).

Pachydactylus serval : Monard (1931:90, 1937b:54).

Pachydactylus punctatus brunnthaleri : Schmidt (1933:5).

Pachydactylus punctatus punctatus : Loveridge (1947:352), Laurent (1954a:63, 1964a:36), Hellmich (1957a:37).

Pachydactylus amoenoides : Laurent (1964a:36).

Pachydactylus punctatus : Bauer and Branch (1995a:70), Branch (1998:259), Bauer et al. (2006b:646), Bauer (1999:57), Bates et al. (2014:130), Ceríaco et al. (2016a:26, 55).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species as presently construed is widespread across southern Africa from the northwest and northeast of South Africa north to Malawi, the former Katanga Province of the Democratic Republic of Congo, and southern Angola.

Ocurrences in Angola (Map 142): Published species records are from southwestern

Angola, but members of this clade also occur in parts of central and southeastern Angola.

Benguela: “Hanha” [-12.25000, 13.75000]

(Bauer and Branch 1995a:82), “Lobito”

[-12.33333, 13.50000] ( Laurent 1954a:63,

1964a:36; Bauer and Branch 1995a:82);

“Catumbella” [-12.43333, 13.55000] ( Bocage

1895a:16; Loveridge 1947:356); “Benguella”

[-12.58333, 13.41667] ( Bocage 1867b:220,

1895a:16; Boulenger 1885:205; Loveridge

1947:356). Huíla: “Indungu” [-14.81667,

16.26667] ( Monard 1937b:54); “Capelongo”

[-14.91667, 15.08333] (Bauer and Branch

1995a:82); “Humpata, environs de Sá da Bandeira ” [-15.03333, 13.40000] ( Laurent

1964a:36); “Kuluї” [-15.41667, 15.73333] MAP 142. Distribution of Pachydactylus punctatus in Angola .

( Monard 1937b:54). Namibe: “ km 60 sur la route de Moçâmedes à Sá da Bandeira” [-15.00000, 12.66667] ( Laurent 1964a:36; Ceríaco et al. 2016a:55); “ 11 mi NE of Mocamede” [-15.08783, 12.26833] [Bauer and Branch 1995:82; Ceríaco et al. 2016a:55); “environs de Moçâmedes” [-15.16667, 12.15000] ( Laurent 1964a:36; Ceríaco et al. 2016a:55); “ 35 km south of Moçâmedes” [-15.50000, 12.16667] ( Laurent 1964a:36; Ceríaco et al. 2016a:55); “Pico Azevedo” [-15.55000, 12.51667] ( Schmidt 1933:5; Loveridge 1947:356; Ceríaco et al. 2016a:55); “Iona National Park” [-16.65669, 12.43672] (Ceríaco et al. 2016a:26); “Espinheira” [-16.78639, 12.35799] (Ceríaco et al. 2016a:26). Cunene: “Riv. Mbalé ” [-15.16667, 16.75000] ( Monard 1931:90); “Forte Roçadas” [-16.71667, 15.01667] ( Laurent 1964a:36). Undetermined Locality: “Cuanza” ( Boulenger 1905:110; Loveridge 1947:356); “Arid subregion” ( Frade 1963:253).

Taxonomic and distributional notes: Members of the Pachydactylus punctatus complex have been confused with a number of other southern African Pachydactylus , including the species P. geitje (Sparrman, 1778) and P. serval Werner, 1910 , South African and Namibian endemics, respectively ( Loveridge 1947; Bauer and Branch 1995a). Bocage’s (1867 b, 1895a), Boulenger’s (1885, 1905) and Frade’s (1963) Angolan specimens of P. ocellatus (= P. geitje ) and Monard’s (1931, 1937b) Pachydactylus serval specimens from “Mbalé,” “Indungu,” and “Kuluї” have all been redetermined as P. punctatus ( Loveridge 1947; Bauer et al. 2006b). Bauer and Lamb (2005) and Heinicke et al. (2011, 2017) identified P. punctatus as part of the “northwestern clade” of Pachydactylus , which comprises numerous morphologically diverse species that are widely distributed in southern Angola and northern Namibia. Laurent (1964a) considered Pachydactylus amoenoides Hewitt, 1935 to be distinct from P. punctatus based on their sympatry in southern Angola, and elevated it to a full species. Although occasionally considered a valid species or subspecies of P. punctatus, Bauer and Branch (1995a) tentatively included amoenoides in the synonymy of P. punctatus . Heinz (2011) identified deep, species-level divergences within P. punctatus sensu lato, and there are a minimum of four taxa in the complex that occur in Angola. A phylogeographic study of P. punctatus is being undertaken (Bauer et al. unpublished).

Pachydactylus rangei ( Andersson, 1908) NAMIB WEB-FOOTED GECKO

Palmatogecko rangei Andersson 1908:299 , pl. 1, figs. a–c. Holotype: MWNH 460 (collector P. Range). “Lüderitzbuch in the German South-West Africa,” Karas Region, Namibia.

Palmatogecko rangei : Mertens (1937a:6), Haacke (1976b:22-23), Russel and Bauer (1990:198), Branch (1998:263), Bauer (1999:57).

Pachydactylus rangei : Bauer and Lamb (2005:116), Herrman and Branch (2013:98), Bates et al. (2014:140), Ceríaco et al. (2016a:55).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is restricted to the Namib Desert and immediately adjacent areas, from the Holgat River in the

Northern Cape, South Africa to extreme southwestern Angola.

Ocurrences in Angola (Map 143): The species occurs in the Nambi Desert, in southwestern Angola. Namibe: “Mossamedes”

[-15.20000, 12.15000] ( Haacke 1976b:22-23;

Ceríaco et al. 2016a:55); “Coroca River ”

[-15.78333, 12.06667] ( Haacke 1976b:22-23;

Ceríaco et al. 2016a:55); “ Porto Alexandre”

[-15.80000, 11.83333] ( Haacke 1976b:22-23;

Ceríaco et al. 2016a:55); “ Cunene mouth”

[-17.28333, 11.80000] ( Haacke 1976b:22-23);

“Lacrau” ( Haacke 1976b:22-23; Ceríaco et al.

2016a:55); “Namib desert” ( Mertens 1937a:6;

Hermann and Branch 2013:98; Ceríaco et al. MAP 143. Distribution of Pachydactylus rangei in Angola.

2016a:55).

Taxonomic and distributional notes: This species was initially described as the type species of the genus Palmatogecko Andersson, 1908 ( Bates et al. 2014). Joger (1985) used immunological data to infer its close affinities with Pachydactylus and Bauer and Lamb (2005) synonymized the genus Palmatogecko with Pachydactylus , within which it is most closely allied to P. vanzyli (Steyn and Haacke, 1966) and P. austeni Hewitt, 1923 (Heinicke et al. 2017). Pachydactylus rangei occupies compacted windward dune faces in the Namib sand seas and other areas of loose sand, including dry river beds, penetrating inland as far as the Namib fog belt ( Haacke 1976b; Russell and Bauer 1990; Branch 1998; Bauer 1999; Bates et al. 2014).

Pachydactylus scherzi Mertens, 1954 SCHERZ’S THICK-TOED GECKO

Pachydactylus scherzi Mertens 1954:175 , fig. 1. Holotype: SMF 45696 (collectors R. Mertens and E. Scherz). Type locality: “Welwitschia-Fläche der Namib am Südwestrand des Brandberg-Massivs, Damaraland, Südwest-Afrika” [= Welwitschia area of the Namib on the southwestern slope of the Brandberg massif, Damaraland], Namibia.

Pachydactylus scherzi : Bauer and Branch (1995a:69), Bauer and Lamb (2005:116), Heinicke et al. (2011:12),

Bauer and Branch (2012:84)

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from western Angola and Namibia, chiefly in association with the Great Western Escarpment.

Ocurrences in Angola (Map 144): The species occurs in southwestern Angola . Huíla: “ 10 mi. NE Sa da Bandeira ” [-14.81000, 13.61783] (Bauer and Branch 1995a:84) .

Taxonomic and distributional notes: Pachydactylus scherzi is a member of the P. punctatus complex of small mostly terrestrial geckos distributed throughout the semi-arid and arid regions of southern Africa (Bauer and Branch 1995a; Heinz 2011), which is, in turn a member of the “northwestern clade” of Pachydactylus (Bauer and Lamb 2005; Heinicke et al. 2011, 2017). Although long overlooked (e.g., Branch 1988), P. scherzi was treated by Bauer and Branch (1995a) as a full species, with several regional morphs. The single published Angolan record corresponds to Bauer and Branch’s (1995a) northern or “Sanitatas MAP 144. Distribution of Pachydactylus scherzi in

Angola. type form.” Heinz (2011) identified deep, species level divergences within P. punctatus sensu lato, and found that P. scherzi was embedded within this group. A phylogeographic study of the P. punctatus complex is being undertaken (Bauer et al., unpublished).

Pachydactylus scutatus Hewitt, 1927 SCALY THICK-TOED GECKO Pachydactylus scutatus Hewitt 1927:395 , pl. 23, fig.

2. Holotype: SAM 17471 (collector R.F.

Lawrence). Type locality: “Kowaris, S.W.A.”

[= Kowares] Kunene Region, Namibia.

Pachydactylus scutatus : Bauer et al. (1993:37),

Branch (1998:260), Bauer (1999:56), Bauer

(2010:266), Ceríaco et al. (2016a:25).

Global conservation status (IUCN): Not

Evaluated.

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

Ocurrences in Angola (Map 145): The species occurs in southwestern Angola. Namibe: “ 6 km S Rio Coroca, Iona ” [-15.774892,

11.895472] ( Bauer 2010:266); “Espinheira”

[-16.79772, 12.35422] ( Bauer 2010:266;

Ceríaco et al. 2016a:25); “Iona, Iona Reserve ” MAP 145. Distribution of Pachydactylus scutatus in [-16.90000, 12.583333] ( Bauer 2010:266). Angola .

Taxonomic and distributional notes: Bauer et al. (2002a) re-examined the types of P. s. angolensis and revised the P. scutatus group, revalidating P. angolensis as a full species, although closely allied to P. scutatus . Ceríaco et al. (2016a) mistakenly stated that their Espinheira record was the first published for the country.

Pachydactylus vanzyli (Steyn and Haacke, 1966) NAMIB DESERT GECKO

Kaokogecko vanzyli Steyn and Haacke, 1966:6 . Holotype: CR 4000/7 (currently NHMW 1635 ) (collectors

W. Steyn, P.S. Swart, C.G. Coetzee and W.D. Haacke). Type locality: “About 18 miles SW of Orupembe,

Kaokoveld, South West Africa,” Kunene Region, Namibia. Kaokogecko vanzyli : Haacke (1976a:23). Palmatogecko vanzyli : Branch (1998:264). Pachydactylus vanzyli : Bauer and Lamb (2005:116), Ceríaco et al. (2016a:55).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from far northwestern Namibia and adjacent southern Angola.

Ocurrences in Angola (Map 146): The species occurs in southern areas of Namibe

Province. Namibe: “Espinheira” [-16.74477,

12.38186] ( Haacke 1976a:25-26; Ceríaco et al.

2016a:55); “Kakolo windmill” [-16.66667,

12.33333] ( Haacke 1976a:25-26; Ceríaco et al.

2016a:55).

Taxonomic and distributional notes:

This species was first described by Steyn and

Haacke (1966) in a new genus, Kaokogecko ,

the affinities of which have long been recognized to be with Pachydactylus , and P. rangei ,

in particular ( Haacke 1976a, 1976c; Joger

1985; Bauer 1990). Kluge and Nussbaum

(1995) synonymized Kaokogecko with

Palmatogecko and Bauer and Lamb (2005)

subsequently synonymized the latter genus with Pachydactylus . These authors, as well as MAP 146. Distribution of Pachydactylus vanzyli in Angola.

Heinicke et al. (2011, 2017), confirmed

K. vanzyli ’s sister group relationship to P. rangei .

Pachydactylus wahlbergii (Peters, 1869) WAHLBERG’S KALAHARI GECKO

Colopus Wahlbergii ( Peters 1869:57, pl. fig. 1). Holotype: NHR 385 (collector J. Wahlberg). Type locality: “Damaralande” [= Damaraland], Namibia.

Colopus wahlbergii wahlbergii : Haacke (1976b:30, 1984:173).

Colopus wahlbergii : Branch and Lamb (2005:118).

Pachydactylus wahlbergii : Heinicke et al. (2017:14).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from southern Africa from southeastern Angola, through Namibia and Botswana to western Zimbabwe and Limpopo Province, South Africa.

Ocurrences in Angola: The species occurs in southeastern Angola in Cuando Cubango Province, close to the Namibian border, although there appear to be no specific published records.

Taxonomic and distributional notes: This species was originally described in the monotypic genus Colopus Peters, 1869 . Bauer and Lamb (2005) retained Colopus as valid despite poor molecular phylogenetic support. Heinicke et al. (2017), however, have conclusively shown that Colopus wahlbergii is embedded within Pachydactylus . Haacke (1976b) cited Colopus wahlbergii from Angola, noting its occurrence in southeastern areas near the Namibian border, but without providing specific localities.