Chamaeleo 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 : 279-283

Genus Chamaeleo Laurenti, 1768

Chamaeleo anchietae Bocage, 1872 ANCHIETA’S CHAMELEON

Chamaeleon Anchietae Bocage 1872:72 , fig. p. 72. Syntypes: MBL (originally 5 specimens) catalogue numbers unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978; Bauer et al. (2006:270) identified ZMB 7756 as a surviving syntype, exchanged from Bocage to Wilhelm Peters. Type locality: “Huilla, dans l’intérieur de Mossamedes” ( Bocage 1872:72), [= Huíla], Huíla Province, Angola.

Chamaeleo anchietae : Boulenger (1887:452), Tilbury and Tolley (2009:63).

Chamaeleon Anchietae : Bocage (1895a:62, 1897a:198),

Chamaeleon anchietae : Loveridge (1933:333), Frade (1963:252).

Chamaeleo senegalensis anchietae : Loveridge (1957:198).

Chamaeleo anchietae : Broadley and Cotterill (2004:40), Bauer et al. (2006:270), Tilbury (2010:447), Ceríaco et al. (2016a:58).

Global conservation status (IUCN): Least Concern.

Global distribution: The species occurs from the interior plateaus to montane grasslands in southwestern Angola through the former Katanga Province of the Democratic

Republic of Congo and the Albertine Rift to southwestern Tanzania and Namibe.

Ocurrences in Angola (Map 240): The species is known from southwestern Angola.

Huíla: “Lobango” [-14.91667, 13.50000]

( Bocage 1895a:62, 1897a:198); “Huilla”

[-15.05000, 13.55000] ( Bocage 1872:72,

1895a:62, 1897a:198; Loveridge 1933:333,

1957:198; Bauer et al. 2006:270). Namibe:

“ Namibe ” ( Tilbury 2010:451; Ceríaco et al.

2016a:58).

Taxonomic and distributional notes:

Based on Bocage’s (1872) somewhat ambigu-

ous locality Boulenger (1887) wrongly consid-

MAP 240. Distribution of Chamaeleo anchietae in Angola.

ered the terra typica to be “Mossamedes,

South-west Africa.” Bocage (1895a) restated the locality simply as “Huilla”.

Chamaeleo dilepis quilensis Bocage, 1886 QUILO FLAP-NECK CHAMELEON

Chamaeleon Dilepis Leach 1819:493 . Holotype: BMNH 1946.8 .13.75 (formerly BMNH 25.5 a) (presented by T.E. Bowditch). Type locality: Not stated. “ French Congo [= Congo]” fide Klaver and Böhme (1987).

Chamaeleo Capellii ( Bocage 1866a:42, 1866c:59). Holotype: MBL specimen number unknown (collector A. Capello), destroyed by fire 18 March 1978. Type locality: “ Benguela,” Benguela Province, Angola.

Chamaeleo dilepis var. quilensis Bocage 1866c:59 . Holotype: MBL specimen number unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “ Rio Quilo ,” Angola.

Chamaeleon dilepis : Bocage (1866a:42, 1866c:59, 1867b:219, 1870:68, 1895a:59, 1896a:112), Peters (1887:612), Boulenger (1887:450, 1905:112), Monard (1937b:99), Themido (1941:8).

Chamaeleo dilepis : Bocage (1879c:88, 1887a:178, 1887b:202, 1887c:209).

Chamaeleon parvilobus : Boulenger (1887:449).

Chamaeleon quilensis : Bocage (1895a:60, 1897a:198).

Chamaeleon quilensis : Boulenger (1905:112), Monard (1937b:99).

Chamaeleo dilepis quilensis : Angel (1923:165), Loveridge (1933:331, 1957:199), Glaw (2015:201). Chamaeleo dilepis : Schmidt (1933:12), Mertens (1937a:7, 1938a:435), Parker (1936:141), Manaças (1963:231), Branch and Conradie (2015:200).

Chamaeleo dilepis dilepis : Loveridge (1936a:76, 1957:199), Laurent (1950a:12, 1954a:65, 1964a:44), Hellmich (1957a:52, 1957b:53), Chirio and LeBreton (2007:182), Glaw (2015:200).

Chamaeleo quilensis : Laurent (1954a:65, 1964a:44), Chirio and LeBreton (2007:198).

Chamaeleo dilepis “ quilensis ”: Tilbury (2010:495).

Chamaeleo dilepis : Tolley (2014), Ceríaco et al. (2016a:58), Ceríaco et al. (2016b:73), Conradie et al. (2016: 24) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species as a whole is widely distributed throughout southern and eastern Africa, although the limits of its range in some areas are under dispute. There is general agreement that the distribution includes the area from Congo east to Tanzania, Kenya, Somalia and Ethiopia, and south to South Africa. However, records from Gabon and north and from Uganda have been questioned (Chirio and Ineich 2006; Pauwels and Vande weghe 2008), and it may be difficult to distinguish this taxon from its near relative C. gracilis , with which it is sympatric in parts of Central Africa.

Ocurrences in Angola (Map 241):

Chamaeleo dilepis quilensis is widely distributed across the entire country.

Cabinda: “Chinchoxo” [-5.10000,

12.10000] ( Peters 1877a:612); “ Cabinda ”

[-5.55000, 12.18333] ( Bocage 1866a:42,

1866c:59, 1895a:59); Rio Quilo ” [-5.18333,

12.18333] ( Bocage 1866a:42, 1866b:59,

1895a:60, 1897a:198; Loveridge 1933:331,

1957:199; Chirio and LeBreton 2007:198).

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

14.23333] ( Bocage 1887a:178, 1895a:59;

1897a:198). Lunda Norte: “Dundo”

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

1954a:65); “Muita (Luembe E) “ [-7.80000,

21.45000] ( Laurent 1950a:121954a:65);

“Carumbo” [-7.74172, 19.95467] (Branch and

Conradie 2015:200). Lunda Sul: “Cacolo MAP 241. Distribution of Chamaeleo dilepis quilensis in Angola.

(Minungo)” [-10.15000, 19.28333] ( Manaças

1963:231). Moxico: “ Lago Dilolo” [-11.50000, 22.01667] ( Manaças 1963:231); “Teixeira de Sousa, distr. Villa Luso” [-10.70000, 22.23333] (Mertens 1937:8); “ Lago Cameia” [-11.71667, 20.80000] ( Manaças 1963:231); “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] ( Manaças 1963:231); “Calombe, Luso” [-11.83333, 19.93333] ( Manaças 1963:231); “environs du lac Calundo” [-11.80000, 20.86667] ( Laurent 1964a:44); “Calunda” [-12.11667, 23.46667] ( Laurent 1964a:44); “Macondo” [-12.55000, 23.76667] ( Laurent 1964a:44); “Cazombo” [-11.88333, 22.91667] ( Laurent 1954a:65). Malanje: “Duque de Bragança ” [-9.10000, 15.95000] ( Bocage 1895a:59); “Cangandala National Park” [-9.81858, 16.65403] (Ceríaco et al. 2016b:73). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] ( Bocage 1867b:219, 1895a:59). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1933:12); “Chitau” [-11.43333, 17.15000] ( Schmidt 1933:12); “General Machado (= Camacupa)” [-12.03333, 17.46667] ( Mertens 1937a:7); “Farm Goedecke on Conjo, 25 km north of General Machado (= Camacupa)” [-11.798563, 17.5114] ( Mertens 1937a:6;

Loveridge 1947:208). Huambo: “Bela-Vista (= Sanguengue)” [-12.36667, 16.20000] ( Hellmich 1957b:53); “ Mt. Moco ” [-12.41667, 15.18333] ( Parker 1936:141); “Cuma” [-12.86667, 15.06667] ( Loveridge 1936a:76). Benguela: “Quibula” [-12.28333, 14.68333] ( Bocage 1895a:59); “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:59); “Catumbella” [-12.43333, 13.55000] ( Bocage 1867b:219); “Quissange” [-12.43333, 14.05000] ( Bocage 1887b:209, 1895:59); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:59); “Benguella” [-12.58333, 13.41667] ( Bocage 1866a:42, 1866c:59, 1867b:219, 1895a:59-60; Loveridge 1957:200); “Dombe” [-12.95000, 13.10000] ( Bocage 1867b:219, 1895a:60); “Entre Rios/Chitidi” [-13.01667, 14.63333] ( Mertens 1938a:435, Hellmich 1957a:52); “Cubal” [-13.03333, 14.25000] ( Mertens 1937a:7, 1938a:435; Hellmich 1957b:53); “Marco de Canavezes, perto da barragem do rio Cubal” [-13.30000, 14.20000] ( Bocage 1896a:112); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:112); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:99). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1879c:88; Loveridge 1936a:76); “Lubango” [-14.91667, 13.50000] ( Bocage 1895a:59); “Huilla” [-15.05000, 13.55000] ( Bocage 1895a:59); “Mutundo” [-15.23333, 13.36667] ( Frade 1963:231); “Gambos” [-15.76667, 14.10000] ( Bocage 1895a:59); “Kampulu, environs de Kasinga” [-15.21667, 16.11667] ( Monard 1937b:99); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937b:99); “Kuluї” [-15.41667, 15.73333] ( Monard 1937b:99). Namibe: “Mossamedes” [-15.20000, 12.15000] ( Bocage 1867b:219, 1887b:202, 1895a:59; Boulenger 1887:451; Ceríaco et al. 2016a:58); “Chimba” [-15.20000, 13.68333] ( Hellmich 1957b:53; Ceríaco et al. 2016a:58). Cunene: “Mupanda” [-17.13333, 15.76667] ( Monard 1937b:99). Cuando Cubango: “Lwankundu, sub-tributary of Kwando” [Undetermined Locality] ( Angel 1923:165); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9-10, 24). Undetermined locality: “Between Bihé and Quilenges” ( Boulenger 1905:112); “Mayumba/Maiumba” ( Bocage 1887a:178, 1895a:60, 1897a:198); “Cuanza r.” ( Boulenger 1905:112). Without precise locality: “with no precise locality” ( Laurent 1964a:44).

Taxonomic and distributional notes: Bocage (1866a) noted some distinct characters in a specimen collected by Anchieta in “Rio Quilo” relative to the typical form, Chamaeleo dilepis Leach, 1819 , however, variation across the C. dilepis species complex as a whole suggests that there are few if any consistent differences among these forms and the taxonomy of the C. dilepis complex remains unresolved ( Glaw 2015). The issue is exacerbated by the vague type locality of C. dilepis , “ Congo,”which, in the BMNH register is given as “Gaboon, French Congo ” (although Pauwels and Vande weghe 2008) stated that the species is absent from Gabon). Loveridge (1936a) refered all Angolan specimens from the Field Museum to C. d. quilensis , with the exception of one specimen from “Cuma,” which he regarded as typical dilepis ( FMNH 15365). Broadley (1966) and Tilbury (2010) considered quilensis a variant of the polymorphic C. dilepis , although Klaver and Böhme (1986) and Ullenbruch et al. (2007) identified differences in hemipenial morphology and regarded quilensis as subspecifically (Klaver and Böhme 1987) or even specifically distinct from nominotypical C. dilepis (Klaver and Böhme 1986) . Tilbury (2010) provided an extended discussion of the taxonomic and distributional issues surrounding the C. dilepis complex. Regardless if quilensis is taxonomically distinct from C. dilepis sensu stricto, it seems likely that all members of the complex form Angola are referable to the same form (although some authors have assigned some Angolan material to another doubtful form, C. d. roperi) and we here treat all Angolan specimens as C. d. quilensis pending a more detailed phylogeographic study of the group.

Chamaeleo gracilis etiennei Schmidt, 1919 ETIENNE’S GRACEFUL CHAMAELEON

Chamaeleo gracilis Hallowell 1842:324 , pl. 18. Holotype: ANSP 7325 (“purchased by the Rev. Charles Eden of Monrovia , of one of the African natives”). Type locality: “ Liberia, Western Africa.”

Chamaeleon etiennei Schmidt 1919:574 . Holotype: AMNH 11370 (collector H. Lang). Type locality: “hills near Banana at the mouth of the Congo,” Democratic Republic of Congo.

Chamaeleon gracilis : Bocage (1866a:41, 1867b:219, 1895a:61), Ferreira (1900a:50, 1903:16, 1904:117), Boulenger (1887:448, 1905:111), Monard (1937b:98), Themido (1941:8)

Chamaeleon senegalensis : Bocage (1870:68), Peters (1881:147).

Chamaeleon gracilis : Boulenger (1887:448, 1905:111), Bocage (1895a:61).

Chamaeleo senegalensis : Günther (1865a:480).

Chamaeleon etiennei : Parker (1936:140), Laurent (1964a:42).

Chamaeleo gracilis : Hellmich (1957a:53, 1957b:53), Laurent (1964a:44).

Chamaeleo gracilis gracilis : Loveridge (1957:198).

Chamaeleo gracilis etiennei : Loveridge (1957:197).

Chamaeleo senegalensis senegalensis : Loveridge (1957:197).

Chamaeleon gracilis etiennei : Tilbury (2010:507), Ceríaco et al. (2014b:670), Glaw (2015:202)

Global conservation status (IUCN): Least Concern.

Global distribution: Known from the sounthern Congo to northern Angola

Ocurrences in Angola (Map 242): Limit-

ed to areas from Kwanza Sul and Lunda Sul northwards, including Cabinda. Although we located no published historical records from

Lunda Norte, Tilbury (2010) plotted them there, and it is likely that they occur in this province as well as the immediately adjacent

Democratic Republic of Congo. Cabinda:

“Chinchoxo” [-5.10000, 12,10000] ( Peters

1877a:612; 1881:147; Bocage 1895a:61);

“ Cabinda ” [-5.55000, 12.18333] ( Bocage

1895a:61). Lunda Sul: “Alto Cuílo”

[-10.01667, 19.55000] ( Laurent 1964a:42);

“Alto Chicapa” [-10.88333, 19.23333] ( Laurent 1964a:42). Luanda: “Loanda” [-8.83333,

13.26667] ( Bocage 1867b:219). Kwanza

Norte: “Piri-Dembos” [-8.56667, 14.50000]

( Hellmich 1957b:53); “Ambaca” [-9.26667, MAP 242. Distribution of Chamaeleo gracilis etiennei in 15.18333] ( Bocage 1895a:61); “N’Dalla Angola.

Tando” [-9.30000, 14.91667] ( Ferreira 1904:111); “Cazengo” [-9.33333, 14.76667] ( Ferreira 1903:16); “Mucoso” [-9.53333, 14.65000] ( Hellmich 1957a:53); “Dondo” [-9.68333, 14.43333] ( Bocage 1895a:61); “Canhoca” [-9.25000, 14.68333] ( Ferreira 1904:111; Parker 1936:140). Lunda Sul: “Lunda” [-10.96667, 20.06667] ( Monard 1937b:98); “Mutianvo” [-11.45000, 19.33333] ( Themido 1941:8). Moxico: “environs du lac Calundo” [-11.80000, 20.86667] ( Laurent 1964a:42). Kwanza Sul: “Lembu, Serra de Selles” [-12.86667, 14.11667] ( Ferreira 1904:117); “Congulu” [-10.86667, 14.28333] ( Parker 1936:140); “Condo” [-11.28333, 14.33333] ( Günther 1865a:480; Boulenger 1887:448; Parker 1936:140). Benguela: “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:98). Undetermined locality: “Quango = Cuango” ( Peters 1877a:612, 1881:147; Bocage 1895a:61) (Malanje Province, impossible to georeferenced. See History Section for more detailed information); “Carangigo” ( Boulenger 1887:448).

Taxonomic and distributional notes: Records of Chamaeleo senegalensis Daudin, 1802 from Angola are referable to C. g. etiennei (see Peters Bocage 1895a). Both Tilbury (2010) and Glaw (2015) recognized Chamaeleo gracilis etiennei as a subspecifically distinct forest-dwelling form. The more southerly records listed above require confirmation and may refer to C. dilepis .

Chamaeleo namaquensis Smith, 1831 NAMAQUA CHAMELEON

Chaemelio Namaquensis Smith 1831:17 . Syntypes: BMNH 65.5.4.61–62, MNHP 8017 (collector A. Smith)

(see notes below). Type locality: “western coast of South Africa, near to the mouth of the Orange river ”

[Namaqualand]. Chamaeleon namaquensis : Bocage (1867c:227, 1870:68, 1895a:62), Boulenger (1887:462), Frade

(1963:253). Chamaeleo tuberculiferus : Günther (1865a:480). Chamaeleo namaquensis = C. tuberculiferus : Bocage (1872:72). Chamaeleo namaquensis : Branch (1998:228), Tilbury (2010:519), Glaw (2015:203), Ceríaco et al.

(2016a:58).

Global conservation status (IUCN): Least Concern ( Carpenter 2011).

Global distribution: The species ranges from western South Africa through Namaqualand and the Namib Desert to southern Angola.

Ocurrences in Angola (Map 243): The species occurs only in the arid portion of southwestern Angola mainly in Namibe Province.

Namibe: “Mossamedes” [-15.20000,

12.15000] ( Günther 1865a:480; Bocage

1867c:227, 1872:72, 1895a:62; Boulenger

1887:462; Ceríaco et al. 2016a:58).

Taxonomic and distributional notes:

Smith (1831) did not specify the number of the type specimens, nor their disposition. There is a long history of one stuffed specimen from

Andrew Smith being present in Paris (Duméril and Duméril 1851; Duméril 1852; Brygoo

1983, although not mentioned by Guibé 1954)

and this specimen has generally been regarded as a syntype and is, in fact, the only syntype mentioned by Klaver and Böhme (1987) and

Glaw (2015). However, FitzSimons (1937) MAP 243. Distribution of Chamaeleo namaquensis in Angola.

mentioned two specimens in the BMNH that he regarded as types. These specimens correspond to BMNH 65.5.4.61–62, which although not marked as types in the BMNH registers, were obtained from Andrew Smith and have the explicit locality, “mouth of Orange River.” We regard these specimens as part of the original type series. Klaver and Böhme (1997) also noted the possibility that additional types might yet be found in the collection of the National Museums of Scotland. This species represents a distinct phylogenetic lineage within Chamaeleo and some authors have suggested that this taxon could be recognized as generically distinct (Townsend and Larson 2002), however it is strongly supported as a member of the Chamaeleo clade ( Bates et al. 2014).