Varanus Merrem, 1820

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 : 275-278

Genus Varanus Merrem, 1820

Varanus albigularis albigularis ( Daudin, 1802) WHITE-THROATED MONITOR

[Although we consider the two Angolan subspecies of V. albigularis to be pontentially valid we believe that it is not possible to meaningfully assign many locality records to one or the other. Thus we provide the occurences in Angola and taxonomic and distributional notes for the two combined.]

Tupinambis albigularis Daudin 1802 [An X]:72, pl. XXXII. Holotype: MNHN 6513 (collector unknown) fide

Brygoo (1987). Type locality: “de l’Afrique ou de l’Inde.” Varanus ocellatus : Bocage (1867b:220; 1870:68). Varanus albigularis : Bocage (1895a:27), Monard (1931:92), Themido (1941:7), Bayless (2002:1645). Varanus exanthematicus : Ferreira (1903:16), Themido (1941), Manaças (“1955” 1957:190), Frade

(1963:252). Varanus exanthematicus albigularis : Mertens (1926:152), Monard (1937b:63). Varanus albigularis albigularis : Ceríaco et al. (2016a:58).

Global conservation status (IUCN): Least Concern.

Global distribution: Southern Africa from Angola and Tanzania southwards.

Varanus albigularis angolensis Schmidt, 1933 ANGOLAN WHITE-THROATED MONITOR

Varanus albigularis angolensis ( Schmidt 1933:10, pl. II). Holotype: CM 5967 (collectors R. and L. Boulton). Type locality: “Gauca, Bihe,” Bié Province, Angola.

Varanus albigularis angolensis : Loveridge (1936a:59), McCoy and Richmond (1966:157), Bayless (2002:1645), Ceríaco et al. (2014b:671), Ceríaco et al. (2016a:39, 58).

Varanus exanthematicus angolensis : Mertens (1937a:9), Mertens (1938a:437), Laurent (1964a:48, 1964b:3), McCoy and Richmond (1966:157).

Varanus (Empagusa) exanthematicus angolensis : Mertens (1942a:104), Mertens (1942b:194), Mertens (1942c:359).

Varanus exanthematicus : Manaças (“1955” 1957:192).

Global conservation status (IUCN): Least Concern.

Global distribution: This subspecies is known from northern Angola and adjacent Zambia and the Katanga region of the Democratic Republic of Congo.

Ocurrences in Angola (Map 238):

Lunda Norte: “Dundo” [-7.36667, 20.83333]

( Laurent 1964a:48; Bayless 2002:1661).

Kwanza Norte: “Golungo” [-9.13333,

14.76667] ( Ferreira 1903:16; Monard

1937b:63; Mertens 1942c:359, 360; Manaças

“1955” 1957:192). Malanje: “Capanda”

[-9.72841, 15.34585] (Ceríaco et al.

2014a:671). Moxico: “Cazombo” [-11.88333,

22.91667] ( Laurent 1964a:48; Laurent

1946b:48). Bié: “Gauca” [-11.18333,

17.45000] ( Schmidt 1933:10; Loveridge

1936:59; Mertens 1942c:359; McCoy and

Richmond 1966:157; Bayless 2002:1661).

Huambo: “High-Huambo” [-12.76667,

15.73333] ( Mertens 1926:152, 1937a:9,

1938a:437, 1926:152, 1942a:104, 1942b:194,

210, 216, 1942c:359, 360; Bayless 2002:1661). MAP 238. Distribution of Varanus albigularis sbspp. in Angola.

Benguela: “Cuma” [-12.86667, 15.06667]

( Bayless 2002:1661); “Lobito bay” [-12.35000, 13.55000] ( Bocage 1895a:27; Mertens 1942c:359; Laurent 1964b:3-4; Bayless 2002:1662); “Catumbella” [-12.43333, 13.55000] ( Bocage 1895a:27; Mertens 1926:152, 1942a:104, 1942b:194, 216, 1942c:359, 360; Bayless 2002:1661); “Catumbella-Coporello” [-13.93333, 12.9666] ( Mertens 1926:152, 1942c:359; Bayless 2002:1661); “Corporello” [-13.93333, 12.9666] ( Mertens 1926:152; Bayless 2002:1661); “Quissange” [-12.43333, 14.05000] ( Frade 1963:253; Mertens 1942c:359; Bayless 2002:1662); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:27; Mertens 1942c:359; Bayless 2002:1662); “Benguella” [-12.58333, 13.41667] ( Bocage 1867c:220, 1895:27; Themido 1941:7, 8; Mertens 1942c:359; Manaças “1955” 1957:192; Bayless 2002:1661); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:63; Mertens 1942c:360; Bayless 2002:1661); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:437, 1942c:360; Bayless 2002:1661). “Hanha” [-13.30000, 14.20000] ( Bayless 2002:1661). Huíla: “Molundo” [-15.63333, 15.20000] ( Monard 1937b:63; Mertens 1942c:360; Bayless 2002:1662). Namibe: “ Rio Chimba” [-14.30000, 12.40000] ( Bocage 1895a:27; Mertens 1942c:359; Bayless 2002:1661); “Biballa” [-14.76667, 13.36667] ( Bocage 1895a:27; Mertens 1926:152, 1942c:359; Bayless 2002:1661); “ 7.5 km, NW (by road) of Pico Azevedo” [-15.47600, 12.69000] (Ceríaco et al. 2016a:39); “Caraculo” [-15.01667, 12.66667] ( Bayless 2002:1661); “Chiyaka district” [vic. -15.08333, 12.73333] ( Bayless 2002:1661); “Luciro-Inamangando road” [Undetermined Locality] ( Bayless 2002:1662); “ 7.5 km NW (by road) of Pico Azevedo, adult, found in a rock crevice” [-15.47600, 12.46150] (Ceríaco et al. 2016a:39). Cunene: “Mupa” [-16.18333, 15.75000] ( Monard 1937b:63; Mertens 1942c:360; Bayless 2002:1662); “Chimporo” [-16.03333, 17.15000] ( Monard 1931:92, 1937b:63; Mertens 1942c:359, 360; Bayless 2002:1661). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] ( Monard 1931:92, 1937b:63; Mertens 1942c:360; Bayless 2002:1661). “Kubango” ( Monard 1931:92; Mertens 1942c:359; Bayless 2002:1662). Undetermined localities: “Xangongo, 30 km SEE ” ( Bayless 2002:1662); “Cazuindo” ( Mertens 1942c:359; Bayless 2002:1661).

Taxonomic and distributional notes: The species was long included within the synonymy of the savanna Monitor , Varanus exanthematicus (Bosc 1792) , now considered limited to the north of the equatorial African forest belt ( Bayless 2002). The subspecies angolensis was rejected by Broadley and Howell (1991), although it has been considered valid by many more recent authors (e.g., Branch 1998; Bayless 2002; Bates et al. 2014). The allocation of specimens from some Angolan localities to the nominotypical form appears to reflect their referral by earlier authors, rather than a critical assessment. Phillips (2004) regarded V. a. angolensis to be present throughout Angola. Pending a more thorough evaluation of the species in Angola we have not attempted to distinguish records of the two subspecies.

Varanus niloticus ( Linnaeus, 1758) NILE MONITOR

Lacerta monitor Linnaeus 1758:201 , nomen rejectum (ICZN, Opinion 540, 1959). Syntypes: Specimens figured in Seba (1734, pl. 94, figs. 1–2; 1735, pl. 82, fig. 2, pl. 105, fig. 1), and described by Linnaeus (1754:41) (collectors unknown), presumed lost. Type locality: “in Indiis. “ [= India].

Lacerta nilotica Linnaeus 1766:369 . Holotype: Specimen mentioned by Hasselquist (1757) (collector F. Hasselquist), presumed lost. Type locality: “in AEgypto” [= Egypt].

Varanus niloticus : Bocage (1866a:42, 1867b:220, 1895a:26, 1897b:210), Ferreira (1900a:50), Boulenger (1905:110), Manaças (“1955” 1957:192).

Monitor niloticus : Günther (1865a:480).

Monitor saurus : Bocage (1879b:95, 1887a:178), Peters (1877a:613)

Varanus niloticus : Bocage (1897b:210), Ferreira (1900a:50, 1903:15), Boulenger (1905:110), Monard (1937b:63), Mertens (1938a:436), Ceríaco et al. (2014b:671), Ceríaco et al. (2016b:87), Conradie et al. (2016:26) , Dowell et al. (2016:594).

Varanus (Polydaedalus) niloticus : Mertens (1942a:94, 1942c:321).

Varanus niloticus niloticus : Laurent (1950a:12; 1954a:63; 1964a:47), Hellmich (1957a:60; 1957b:69), Manaças (1963:239).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Senegal, west to Somalia and Egypt, and south to South Africa.

Ocurrences in Angola (Map 239): The species distribution comprises the entire country including Cabinda Province, with exception of the desert regions of far southwestern Angola . Cabinda: “Chinchoxo” [-5.10000,

12.10000] ( Peters 1877a:613); “ Rio Quilo ”

[-5.18333, 12.18333] ( Bocage 1866a:42,

1895a:26; Mertens 1942c:323; Manaças

“1955” 1957:193; Bayless 2002:1676);

“ Cabinda ” [-5.55000, 12.18333] ( Bocage

1866a:42, 1895a:26; Mertens 1942c:323;

Manaças “1955” 1957:193; Bayless 2002:

1675). Luanda: “Loanda” [-8.83333,

13.26667] ( Bocage 1866a:42, 1895a:26; Mertens 1942c:323; Manaças “1955” 1957:193).

Bengo: “ Cunga River ” [-9.23333, 13.76667]

( Boulenger 1905:110; Mertens 1942c:321,

MAP 239. Distribution of Varanus niloticus in 323; Bayless 2002:1675). Kwanza Norte:

Angola.

“Piri-Dembos” [-8.56667, 14.50000] ( Hellmich 1957b:60); “Golungo Alto” [-9.13333, 14.76667] ( Ferreira 1903:15; Mertens 1942c:321; Manaças “1955” 1957:193); “N’dalla Tando” [-9.30000, 14.91667] ( Ferreira 1903:15; Mertens 1942c:321, 323; Bayless 2002:); “ Rio Membege River, perto de N’dalla Tando” [-9.43333, 14.76667] ( Ferreira 1900a:50; Mertens 1942c:321; Manaças “1955” 1957:193; Bayless 2002:1675); “Mucoso” [-9.53333, 14.65000] ( Hellmich 1957a:61); “Dondo” [-9.68333, 14.43333] ( Bocage 1895a:26; Mertens 1942c:323; Manaças “1955” 1957:193; Bayless 2002:1675). Malanje: “Duque de Bragança (Duque do Cunene [Bayless])” [-9.10000, 15.95000] ( Bocage 1895a:26; Mertens 1942c:323; Manaças “1955” 1957:193; Bayless 2002:1675); “Capanda” [-9.72841, 15.34585] (Ceríaco et al. 2014b:671); “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:87); “ Rio Loando” [-11.55000, 22.01667] ( Bocage 1879a:95; Mertens 1942c:323; Bayless 2002:1675). Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1954a:63, 1964a:47; Manaças “1955” 1957:193); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950:12). Moxico: “ Lago Dilolo” [-11.50000, 18.10000] ( Manaças 1963:239). Kwanza Sul: “Condo” [-11.28333, 14.33333] ( Günther 1865a:480; Mertens 1942c:323; Bayless 2002:1675). Benguela: “Catumbella (Catumbinda river [ Bayless 2002])” [-12.43333, 13.55000] ( Bocage 1867b:220, 1895a:26; Mertens 1942c:323; Manaças “1955” 1957:193; Bayless 2002:1675); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:69); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:436; Mertens 1942a:94, 95; Mertens 1942c:324; Bayless 2002:1675); “Hanha” [-13.30000, 14.20000] ( Bocage 1897b:210; Mertens 1942c:323; Manaças “1955” 1957; Bayless 2002:1675). Huíla: “ Rio Cuce” [-13.51667, 15.20000] ( Bocage 1895a:26; Mertens 1942c:323; Bayless 2002:1675); “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:26; Mertens 1942c:323; Bayless 2002:1675); “Quillangues” [-14.06667, 15.08333] ( Bocage 1895a:26; Mertens 1942c:323; Bayless 2002:1675); “Tyitunda” [-14.36667, 16.48333] ( Monard 1937b:63; Mertens 1942c:322, 323; Bayless 2002:1676); “Vila da Ponte (= Kuvangu (Kubangu))” [-14.46667, 16.30000] ( Monard 1931:93, 1937b:63; Mertens 1942c:322, 32; Manaças “1955” 1957:193; Bayless 2002:1675); “Kuluї” [-15.41667, 15.73333] ( Monard 1937b:63; Mertens 1942c:323; Bayless 2002:1675). Cunene: “Riv. Mbalé ” [-15.16667, 16.75000] ( Monard 1931:93, 1937b:63; Mertens 1942c:322, 323; Manaças 1957 “1955”:193); “Humbe” [-16.68333, 14.90000] ( Bocage 1895a:26; Mertens 1942c:323; Bayless 2002:1675); “Foz de Cunene ” [-17.28333, 11.80000] ( Bayless 2002:1675); “Sighting near Cunéné ” ( Monard 1937b:63; Mertens 1942c:323; Bayless 2002:1675). Cuando Cubango: “Cunga” [-17.30000, 20.80000] ( Dowell et al. 2016:594); “Cuito basin (29)” [-17.87291, 19.83333] ( Conradie et al. 2016:8-9, 26); “Cuito basin (35) not collected” [-16.62322, 19.05352] ( Conradie et al. 2016:9-10, 26); “Cuando basin (43) not collected” [-17.53500, 23.18916] ( Conradie et al. 2016:9-10, 26); “Cuando basin (44a) not collected” [-17.57333, 23.26000] ( Conradie et al. 2016:9-10, 26); “Cubango basin (45) not collected” [-16.88350, 18.01180] ( Conradie et al. 2016:9, 12, 26). Undetermined Locality: Rio Quanza” ( Bocage 1895a:26; Mertens 1942c:323); “Mayumba” (Manaças “1955” 1957:193); “ Rio Quango” ( Bocage 1895a:26; Mertens 1942c:323; Manaças “1955” 1957:193; Bayless 2002:1675); “Kubango Gebiet” ( Mertens 1942c:322).

Taxonomic and distributional notes: The name Lacerta monitor Linnaeus, 1758 initially associated with this and other species of varanids, was rejcted by the ICZN under Opinion 540 and Lacerta nilotica , based on a specimen described by Hasselquist (1757) is the accepted name. Varanus niloticus comprises three major clades of monitors, with the sole Angolan sample includ- ed clustering with a very broadly distributed southern clade, also including virtually all samples south of 3°N ( Dowell et al. 2016). This includes those specimens in this geographic range that had previously been assigned to Varanus ornatus (Daudin, 1803) which had been elevated to full specific status by Böhme and Ziegler (1997) based on external features, particularly color.