Agama Daudin, 1802

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 : 287-292

Genus Agama Daudin, 1802

Agama aculeata Merrem, 1820 WESTERN GROUND AGAMA

Agama aculeata Merrem 1820:53 . Lectotype: Specimen illustrated in Fig. 6 , plate 8 of Seba (1735) (collector unknown), probably lost, designated by Wagner et al. (2012). Type locality: “Promontorio bonae spei” [= the Cape of Good Hope] South Africa.

Agama aculeata : Bocage (1866a:43, 1867b:221), Branch (1998:212), Leaché et al. (2009:274), Wagner et al. (2012:177), Ceríaco et al. (2016a:58).

Agama armata : Bocage (1870:68, 1879c:88, 1895a:19, 1896a:110, 1896b:127), Boulenger (1905:110).

Agama hispida aculeata : Loveridge (1936a:52), Mertens (1938a:432), Hellmich (1957a:38, 1957b:51), Manaças (1963:229), Laurent (1964a:40).

Agama hispida var. aculeata : Monard (1937b:59).

Agama aculeata aculeata : Bates et al. (2014:303).

Global conservation status (IUCN): Least Concern.

Global distribution: Endemic to southern Africa where it has an extensive range from southern Angola, most of Namibia except the Namib Desert proper, western Botswana and the western half of South Africa.

Ocurrences in Angola (Map 247): The species widespreasd in Angola except for the more northerly provinces. Malanje: “ Duque de Bragança ” [-9.10000, 15.95000] ( Bocage

1866a:43, 1895a:19, 1896b:127). Lunda Sul:

“Alto Chicapa” [-10.88333, 19.23333] ( Laurent 1964a:40). Moxico: “ Lago Cameia”

[-11.71667, 20.80000] ( Manaças 1963:229),

“ environs du lac Claundo” [-11.80000,

20.86667] ( Laurent 1964a:40), “Calombe,

Luso” [-11.83333, 19.93333] ( Manaças

1963:229); “Cassamba” [-13.10000, 20.35000]

( Manaças 1963:229). Bié: “Silva Porto ”

[-12.38333, 16.95000] ( Manaças 1963:229);

“ Cubango basin (15)” [-12.78555, 16.75694]

( Conradie et al. 2016:8-9, 23). Huambo:

“Bimbi” [-11.81667, 15.83333] (Monard

MAP 247. Distribution of Agama aculeata in Angola. 1937b:59); “Galanga” [-12.06667, 15.15000]

( Bocage 1895a:19); “Bela-Vista (Sanguengue)” [-12.36667, 16.20000] ( Hellmich 1957b:51); “Nova Lisboa ” [-12.76667, 15.73333] ( Manaças 1963:229); “Santo-Amaro” [-12.70000, 15.85000] ( Monard 1937b:59); “Cuma” [-12.86667, 15.06667] ( Loveridge 1936a:52). Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:19) “Quissange” [-12.43333, 14.05000] ( Bocage 1895a:19); “Benguella” [-12.58333, 13.41667] ( Loveridge 1936a:52); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:59); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:38); “Cubal” [-13.03333, 14.25000] (Mertens 1938:432); “Hanha” [-13.30000, 14.20000] (Bocage 1896:110). Huíla: “ Rio Cuce” [-13.51667, 15.20000] ( Bocage 1895a:19); “Cassôco” [-13.65000, 15.01667] ( Bocage 1895a:19); “Caconda” [-13.73333, 15.06667] ( Bocage 1879c:88); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937b:59); “Kuvangu” [-14.46667, 16.30000] ( Monard 1937b:59); “Boca de Humpata, Sá da Bandeira” [-14.93333, 13.51667] ( Laurent 1964a:40); “Chibia, Rio Huíla ” [-15.20000, 13.68333] ( Hellmich 1957b:51); “Kambisa” [-15.31667, 16.21667] ( Monard 1937b:59); “Kuluï” [-15.41667, 15.73333] ( Monard 1937b:59).

Namibe: “Biballa” [-14.76667, 13.36667] ( Bocage 1895a:19; Ceríaco et al. 2016a:58); “Virei- Cahinde” [-15.73333, 12.95000] ( Hellmich 1957b:51; Ceríaco et al. 2016a:58); “Molundo” [-15.63333, 15.20000] ( Monard 1937b:59); “Chimporo” [-16.03333, 17.15000] ( Monard 1937b:59; Ceríaco et al. 2016a:58). Cunene: “Mupanda” [-17.13333, 15.76667] ( Monard 1937b:59); “Riv. Mbalé ” [-15.16667, 16.75000] ( Monard 1937b:59); “Kakindo” [-15.45000, 17.05000] ( Monard 1937b:59); “Humbe” [-16.68333, 14.90000] ( Bocage 1895a:19); “Mupa” [-16.18333, 15.75000] ( Monard 1937b:59). Undetermined Locality: “Between Bihé and Quillenges” (Ferreira 1905:110); “Between Benguela and Bihé” (Ferreira 1905:110); “Quando” ( Bocage 1895a:19).

Taxonomic and distributional notes: The “ holotype ” of A. aculeata was previously identified as ZMB 750 (Denzer et al. 1997). However, Wagner et al. (2012) demonstrated that this specimen, actually an Agama atra Daudin, 1802 , was one of four syntypes. They reviewed the complex history of these specimens and selected as a lectotype the only one of the syntypes actually referable to A. aculeata as currently recognized. The species Agama hispida (Kaup, 1827) and Agama armata Peters, 1854 have previously been included in the Angolan fauna by early authors. Agama hispida is, however, endemic to South Africa and adjacent southwestern Namibia ( Branch 1998; Bates et al. 2014) and Angolan citations to it are likely referable to A. aculeata or, in some cases, to the superficially similar Agama anchietae Bocage, 1896 , with which it occurs sympatricaly in Southern Angola and Namibia. Agama armata likewise is extralimital, with a distribution in southeastern Africa.

Agama anchietae Bocage, 1896 ANCHIETA’S AGAMA

Agama Anchietae ( Bocage 1896b:129) . Syntypes: MBL specimen numbers unknown (collectors J.A.

d’Anchieta [Benguella, Catumbella, Dombe] and H. Capello and R. Ivens [Mossamedes]), destroyed by fire 18 March 1978. Type locality: “Benguella, Catumbella, Dombe et Mossamedes,” Benguella and

Namibe provinces, Angola.

Agama aculeata : Bocage (1867b:221).

Agama armata : Bocage (1887b:203, 1895a:19).

Agama Anchietae : Bocage (1897a:194).

Agama anchietae : Boulenger and Power (1921:268), Monard (1937b:59), Frade (1963:253), Laurent

(1964a:42), Branch (1998:214), Leaché et al. (2009:274), Bates et al. (2014:304), Ceríaco et al. (2016a:21,

58).

Agama anchietae anchietae : Schmidt (1933:9), Parker (1936:131), Laurent (1947:5).

Global conservation status (IUCN): Not Evaluated.

Global distribution: This species has an extensive geographical range that extends from the northwestern part of South Africa northwards through Namibia and Angola to the southern Democratic Republic of the Congo.

Ocurrences in Angola (Map 248): The species occurs chiefly in southern lowland regions of Angola, however, there are two isolated records in Bié Province, central Angola, which require confirmation. Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1933:9); “Chitau” [-11.43333, 17.15000] ( Schmidt 1933:9). Benguela: “Lobito” [-12.35000, 13.55000] ( Monard 1937b:59); “Catumbella” [-12.43333, 13.55000] ( Bocage 1867b:221, 1895a:21, 1896b:129, 1897a:194; Boulenger and Power 1921:269; Laurent 1947:5; Monard 1937b:59); “Benguella” [-12.58333, 13.41667] ( Bocage 1867b:221, 1895a:21, 1896b:127, 1897a:194; Laurent 1947:5; Monard 1937b:59); “Huxe” [-12.71667, 13.20000] (Boulenger and Power 1921:269); “Dombe” [-12.95000, 13.10000] ( Bocage 1867b:221, 1895a:21, 1896b:127, 1897a:194; Laurent 1947:5; Monard 1937b:59). Huíla: “Caconda” [-13.73333, 15.06667] (Boulenger and Power 1921:269; Monard 1937b:59). Namibe: “Maconjo (= Fazenda Mucungo)” [-14.782192, 12.486557] (Boulenger and Power 1921:269; Monard

1937b:59; Ceríaco et al. 2016a:58);

“Mossamedes” [-15.20000, 12.15000]

( Bocage 1887b:203, 1895a:21, 1896b:127,

1897a:194; Monard 1937b:59; Ceríaco et al.

2016a:58); “ Iona National Park ” [-16.65753,

12.43807] (Ceríaco et al. 2016a:21); “Pico

Azevedo” [-15.53400, 12.49197] ( Ceríaco et al. 2016a:21); “ Namibe Natural Park”

[-15.77428, 12.33311] (Ceríaco et al.

2016a:21); “ 100 km southeast of Moçâmedes”

[-15.66515, 12.73503] ( Laurent 1964a:42;

Ceríaco et al. 2016a:58). Undetermined

Locality: “Cuangu” (Boulenger and Power

1921:269) (Malanje Province impossible to georeference: See History Section for more detailed information).

Taxonomic and distributional notes: MAP 248. Distribution of Agama anchietae in Angola.

Boulenger and Power (1921) refered to two specimens from “Caconda” and “Catumbella” in the British Museum collection sent by Bocage, stating that the “Caconda” material belonged to the type series. However, it is clear ( Bocage 1896b, 1897a) that this cannot be the case. Specimens of A. anchietae can be confused with the similarly-sized ground dwelling congener A. aculeata , so all older records should be reexamined.

Agama congica Peters, 1877 CONGO AGAMA

Agama colonorum var. congica ( Peters 1877a: 612). Syntypes: ZMB 9169 (3 specimens) fide Bauer et al .

(1995). Type locality: “Chinchoxo,” Cabinda Province, Angola. Agama colonorum : Bocage (1866a:42, 1895a:17). Agama planiceps : Bocage (1887a:178, 1895a:18, 1896a:110. Agama planiceps planiceps : Loveridge (1936a:56). Agama colonorum var. congica : Denzer et al.

(1997:313).

Global conservation status (IUCN): Not

Evaluated.

Global distribution: The distribution of this species has not been established, but it is probably limited to the region of northern

Angola and adjacent regions of central Africa.

Ocurrences in Angola (Map 249): The species is known from northwestern Angola.

Cabinda: “Chinchoxo” [-5.10000, 12.10000]

( Peters 1877a:612; Loveridge 1957:191; Wolfgang et al. 1997:313); “ Cabinda ” [-5.55000,

12.18333] ( Bocage 1866a:42, 1887a:178,

1895a:17). Zaire: “S. Salvador do Congo”

[-6.26667, 14.23333] ( Bocage 1895a:17, 18).

MAP 249. Distribution of Agama congica in Angola.

Taxonomic and distributional notes: Agama congica was described by Peters (1877) as a variety of Agama colonorum . On the basis of the recommendation of Philipp Wagner, we have recognized A. congica as specifically distinct. The distribution and affinities of this taxon are currently under study. Ceríaco et al. (2014b) considered that specimens of Capanda Dam, Malanje Province, were true congica , but recent reanalysis suggests that these specimens should be considered as representatives of an as yet undescribed endemic form (see Agama sp. account).

Agama mucosoensis Hellmich, 1957 MUCOSO AGAMA (Endemic)

Agama agama mucosoënsis ( Hellmich 1957a:44, pl. 8). Holotype: ZSM 117 /53 (collector W. Hellmich). Type locality: “Mucoso bei Dondo” [= Mucoso, Dondo] Kwanza Norte Province, Angola.

Agama agama mucosoënsis : Hellmich (1957b:50).

Agama mucosoensis : Wagner et al. (2012:184), Ceríaco et al. (2014b:670).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is only known from Angola.

Ocurrences in Angola (Map 250): The species is known only from the type locality and near areas in western Angola . Kwanza

Norte: “Mucoso” [-9.53333, 14.65000]

( Hellmich 1957a:44; Wagner et al. 2012. 18);

“Dondo” [-9.66667, 14.41667] ( Hellmich

1957b:50; Wagner et al. 2012. 18). Kwanza

Sul: “Libolo/Luati” [-9.98333, 14.90000]

( Hellmich 1957b:50; Wagner et al. 2012. 18).

Taxonomic and distributional notes:

Hellmich (1957a) described Agama agama mucosoënsis based on a type series of 72 specimens collected in “Mucoso bei Dondo”,

Kwanza Norte Province, recently revalidated by Wagner et al. (2012) as a full species. The only published data for this species are from

Kwanza Norte Province, although specimens have recently been deposited in the Museu MAP 250. Distribution of Agama mucosoensis in Angola. Nacional de História Natural e da Ciência Lisboa, Portugal from “Açucareira”(Bengo) and “Quifangondo” (Luanda), Angola (L. Ceríaco and M. Marques, pers. obs.). Ceríaco et al. (2014b) also suggested that the species might occur in Malanje Province. There is a possibility of confusion between this species and A. congica , both of which exhibit sexual dimorphism, and all historical records should be reexamined with this in mind .

Agama planiceps Peters, 1862 NAMIB ROCK AGAMA

Agama planiceps Peters 1862a:15 . Syntypes: ZMB 4200–4201 (collector C. H. Hahn). Type locality: “Neu-Barmen” [= Gross Barmen, Otjozondjupa Region], Namibia.

Agama planiceps : Schmidt (1933:9), Ceríaco et al. (2016a:21, 58).

Global conservation status (IUCN): Not Evaluated.

Global distribution: Agama planiceps occurs in the northwestern quadrant of Namibia and the southwestern quadrant of Angola.

Ocurrences in Angola (Map 251): The species is known the true desert areas of Namibe province. Namibe: “Pico Azevedo” [-15.55000, 12.51667] ( Schmidt 1933:9; Ceríaco et al.

2016a:58); “Lion Cave’s at 3.4 km SW of

Espinheira camp” [-16.81250, 12.33969]

(Ceríaco et al. 2016a:21–22); “Omauha Lodge camp” [-16.19872, 12.40008] (Ceríaco et al.

2016a:22); “ Iona National Park , north of Tambor ” [-15.99611, 12.40667] (Ceríaco et al.

2016a:22); “Iona National Park, south side of

Curoca River crossing” [-16.30433, 12.41710]

(Ceríaco et al. 2016a:22); “Pediva Hot

Springs” [-16.12214, 12.56111] (Ceríaco et al.

2016a:22); “ Namibe Lubango road, road mark-

er 59, 1.8 km west by road from Caraculo,

north side of the road” [-15.01600, 12.64356]

(Ceríaco et al. 2016a:22); “Pico Azevedo”

[-15.53400, 12.49197] (Ceríaco et al.

2016a:22).

Taxonomic and distributional notes:

Bauer et

al. (1995) incorrectly gave the current MAP 251. Distribution of Agama planiceps in Angola.

name of the type locality of A. planiceps (Neu Barmen) as “Otjimbingue,” which is actually anoth- er Rhenish Mission site to the southwest of Neu Barmen. In Angola the species appears to be confined to the true desert areas in Namibe, being replaced by A. schacki in the northern and eastern parts of the province. However, Namibian A. planiceps occur broadly throughout arid and semi-arid areas where suitable rocky habitats occur. Monard’s (1937b) record of “ A. planiceps ” from Cuando Cubango Province is certainly in error and is not included in our map.

Agama schacki Mertens, 1938 SCHACK’S ROCK AGAMA (Endemic)

Agama planiceps schacki ( Mertens 1938a:433). Holotype: SMF 25299 (collector W. Schack). Type locality:

“Cubal, 900m H. Prov. Benguella, Angola ” [= Cubal], Benguela Province, Angola. Agama planiceps : Bocage (1887c:210, 1895a:18, 1896a:110), Boulenger (1885:358, 1905:110), Mertens

(1926:152), Monard (1937b:60). Agama colonorum : Boulenger (1885:357), Ferreira (1905:117, 1906:170). Agama planiceps schacki : Hellmich (1957a:47, 1957b:52), Mertens (1967:61), Manaças (1963:230), Laurent

(1964a:40), Branch (1998:218).

Global conservation status (IUCN): Not Evaluated

Global distribution: Agama schacki occurs in the southwestern quadrant of Angola.

Ocurrences in Angola (Map 252): The species occurs along the southern areas of the escarpment (Benguela and Huíla provinces), as well as in the lowlands of Benguela Province. Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:18); “Morro de Pundo” [-12.38333, 13.88333] ( Parker 1936:129); “Quissange” [-12.43333, 14.05000] ( Bocage 1887c:210, 1895a:18); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:18); “Benguella” [-12.58333, 13.41667] ( Boulenger 1885:357; Bocage 1895a:18); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:60); “Lembu, Serra de Selles” [-12.86667, 14.11667] (Ferreira 1905:117, 1906:170); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:47); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:433, 1967:61; Hellmich 1957b:52); “Marco de Canavezes” [-13.08333, 14.33333] ( Laurent 1964a:40); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:110); “Catengue” [-13.03333, 13.73333] (Mertens 1927:152). Huíla: “Senhora do Monte, Sá da Bandeira” [-14.93333, 13.43333] ( Laurent 1964a:40); “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] ( Laurent 1964a:40). Namibe: “Biballa” [-14.76667, 13.36667] ( Bocage

1895a:18; Ceríaco et al. 2016a:58).

Taxonomic and distributional notes:

Mertens (1938a) diagnosed A. planiceps schacki from the nominotypical form on the basis of a larger, more robust body, a greater number of midbody scale rows (80–90 vs. 63–

76), and a black (rather than red) tail tip.

Branch (1998) stated that A. p. schacki replaces the typical form in Angola. Recently collected molecular and morphological data suggests that both forms occur in Angola and that planiceps and schacki , although closely related, are two distinct species. A detailed review of the group is being prepared. Records from Benguela ( Boulenger 1885) and Lembu

( Ferreira 1906, 1906) originally referred to

MAP 252. Distribution of Agama schacki in Angola. Agama colonorum have been tentatively assigned to A. schacki on the basis of their distribution.

Figures

FIGURE 6. Annual mean precipitation in Angola (after Hijmans et al. 2005).