Causus Wagler, 1830

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 : 315-320

Genus Causus Wagler, 1830

Causus bilineatus Boulenger, 1905 TWO-STRIPED NIGHT ADDER

Causus rhombeatus var. bilineatus Boulenger 1905:114 . Syntypes: BMNH 1905.5.29.36-40 (collector W.J. Ansorge). Type locality: “Duque de Braganca, Quissange, Caconda et Huilla” ( Bocage 1895a:146) [= Calandula, Quissange, Caconda and Huíla] Angola, and “Between Benguela and Bihé” ( Boulenger 1905:114), Angola.

Causus rhombeatus : Bocage (1895a:147); Boulenger (1896:468); Schmidt (1933:15), Hellmich (1957b:74), Loveridge (1957:299).

Causus bilineatus bilineatus : Laurent (1964a:125).

Causus bilineatus : Spawls et al. (2004:469), Broadley (1968b:407), Spawls and Branch (1995:140), Broadley and Cotterill (2004:46), Rasmussen (2005:1), Manaças (1981:33), Dobiey and Vogel (2007:125), Wallach et al. (2014:148).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from the plateau of South-Central Africa, in Angola, southern Democratic Republic of Congo and north-west Zambia, reaching Tanzania. Ocurrences in Angola (Map 275): The species occurs from west to eastern Angola.

Kwanza Norte: “Piri-Dembos” [-8.53333,

14.43333] ( Hellmich 1957b:74; Manaças

1981:33; Rasmussen 2005:14); “Canhoca”

[-9.25000, 14.68333] ( Boulenger 1905:114;

Manaças 1981:33). Malanje: “Duque de Bragança ” [-9.10000, 15.95000] ( Bocage

1866a:51, 1895a:145; Monard 1937b:141,

Manaças 1981:33; Rasmussen 2005:14).

Moxico: “ Rives du lac Calundo (Lago Calundo )” [-11.80000, 20.86667] ( Laurent

1964a:125; Manaças 1981:33; Rasmussen

2005:14). Bié: “Chitau” [-11.43333, 17.15000]

( Schmidt 1933:15; Rasmussen 2005:14).

Huíla: “Caconda” [-13.73333, 15.06667]

( Bocage 1895a:145; Monard 1937b:141;

Manaças 1981:33; Rasmussen 2005:14); MAP 275 Distribution of Causus bilineatus in Angola. “Capelongo” [-14.88333, 15.08333] (Manaças 1981:33); “Huíla” [-15.08333, 13.55000] ( Bocage 1895a:145; Boulenger 1896:468; Monard 1937b:141; Manaças 1981:33; Rasmussen 2005:14). Taxonomic and distributional notes: Bocage (1895a) in his remarks on Causus rhombeatus ( Lichtenstein, 1823) recognized a striped form based on several individuals from Angola deposit- ed in the Museu Bocage collection from “Duque de Bragança, Quissange, Caconda et Huilla,” although he did not recognize them as taxonomically distinct. Boulenger (1905) subsequently noted the same striped form described by Bocage based on specimens from “between Benguella and Bihé” and proposed the name C. rhombeatus var. bilineatus . McDiarmid et al. (1999) includ- ed Bocage’s (1895a) specimens among the type series, but Boulenger (1905) clearly refered only to the striped specimens collected by Ansorge and in the BMNH collection. For some time, the nomen bilineatus was neglected (e.g., Schmidt 1933; Monard, 1937b; Bogert, 1940; Hellmich 1957a,b) and the corresponding Angolan records were associated with C. rhombeatus . Laurent (1964a) revived the name bilineatus , and included his recently described Causus lineatus ( Laurent, 1956) , which he considered a dwarf subspecies of Causus bilineatus . However, his specimens differ little from those of the type series from “ Benguela to Bihe” (Broadley and Cotterill 2004). Loveridge (1957) included both bilineatus and lineatus in the synoymy of C. rhombeatus , but Broadley (1968b) placed C. b. lineatus in the synonymy of C. bilineatus without comment, where it has since remained ( Rasmussen 2005). Laurent (1964a) erroneously assigned some Angolan records of C. rhombeatus to C. bilineatus (e.g., “Pungo-Andongo” [ Boulenger 1905]; “Mombolo” [ Bogert 1940]; “Bela Vista”, “Entre Rios” [ Hellmich 1957b]) as did Manaças (1981). Most of the specimens cited by Laurent (1964a) were examined and revised by Rasmussen (2005).

Causus lichtensteinii ( Jan, 1859) FOREST NIGHT ADDER

A [spidelaps] (Sepedon) Lichtensteinii Jan 1859:511 . Holotype: NMBA, lost (fide Kramer 1978; Golay et al. 1993), destroyed in 1943 during the World War II fide Wallach et al. (2014). Type locality: “Côte-d’Or” [= Ghana].

Causus lichtensteinii : Laurent (1950a:11, 1964a:127), Manaças (1981:34), Spawls and Branch (1995:143), Spawls et al. (2004:471), Rasmussen (2005:11), Chirio and LeBreton (2007:606), Dobiey and Vogel (2007:126), Akani et al. (2012:192), Wallach et al. (2014:149).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from the central African forests from Guinea and Liberia eastward through Ghana to Nigeria, Cameroon, the Central African Republic, south to Democratic Republic of Congo, northeastern

Angola, and northwestern Zambia, and east to

Uganda and western Kenya.

Ocurrences in Angola (Map 276): The species occurs in the northtern Angola, in the adjacent areas near the border with Democratic

Republic of Congo and in the Cabinda encalve,

although there are no published records from

Cabinda. Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1964a:127, Manaças

1981:34); “rivière Lukashi, afluente de la rive gauche de la Tshihumbwe (Rio Lukashi)”

[-7.53333, 21.08333] ( Laurent 1950a:11;

Manaças 1981:34).

Taxonomic and distributional notes:

Wallach et al. (2014) recorded this species as present in Benguela Province, although this almost certainly is based on a misidentifica- MAP 276. Distribution of Causus lichtensteinii in tion. Angola.

Causus maculatus ( Hallowell, 1842) SPOTTED NIGHT ADDER

Distichurus Maculatus Hallowell 1842:337 , pl. 19. Holotype: ANSP 6897 (collector S.M.E. Goheen). Type locality: “ Liberia, Westen Africa”.

Causus rhombeatus : Bocage (1895a:145), Parker (1936:126), Laurent (1950a:11, 1954a:61), Hellmich (1957b:74).

Causus maculatus : Laurent (1964a:124), Thys van den Audenaerde (1966:36), Manaças (1981:32), Spawls and Branch (1995:145), Rasmussen (2005:14), Chippaux (2006:241), Trapé and Mané (2006:206), Dobiey and Vogel (2007:127), Chirio and LeBreton (2007:608), Wallach et al. (2014:148).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from West-Central Africa from Mauritania to Ethiopia, and south to Angola.

Ocurrences in Angola (Map 277): The species occurs in the northern Angola. Lunda

Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1950a:11, 1954a:61, 1964a:124, Thys van den Audenaerde 1966:36, Manaças 1891:32;

Rasmussen 2005:14); “Barrage de la Luachimo” [-7.38333, 20.85000] ( Thys van den

Audenaerde 1966:36); “Muita (Luembe E)”

[-7.80000, 21.45000] ( Laurent 1950a:11,

1954a:61); “Sombo” [-8.68333, 20.95000]

( Laurent 1954a:61). Kwanza Norte: “PiriDembos” [-8.53333, 14.43333] ( Hellmich

1957b:74; Rasmussen 2005:14). Kwanza Sul:

“Quirimbo” [-10.68333, 14.26667] ( Parker

1936:126).

Taxonomic and distributional notes:

Bogert (1940) pointed out a possible taxonom- MAP 277. Distribution of Causus maculatus in Angola. ic difference between the savanna C. rhombeatus (typical form) and the forest dwellers, which was later confirmed by Laurent (1956, 1964a). Laurent (1960) identified C. maculatus as a forest form, distinguishing it from C. rhombeatus , which is sympatric in Angola (Manaças, 1981). Laurent (1964a) assigned the Angolan records previously refered to C. rhombeatus (e.g., “Dundo,” “Muita,” “Sombo” [ Laurent 1950a, 1954a], “Quirimbo” [ Parker 1936]) to C. maculatus , and refered the specimens cited in Bocage (1895a) from Congo and a part of northern Angolan to C. maculatus, Without further comment. According to Rasmussen (2005) two specimens from the Zoological Museum of the University of Hamburg previously identified as Causus rhombeatus ( Lichtenstein, 1823) from “Piri-Dembos” ( Hellmich, 1957b) correspond, in fact, to Causus maculatus .

Causus resimus (Peters, 1862) GREEN NIGHT ADDER

Heterophis resimus Peters 1862b:277 , pl., figs. 4, 4a, 4b. Holotype: ZMB 4370 (collector A. von Barnin and R. Hartman). Type locality: “Sennâr, vom Gebel-Ghule” [= Jebel Ghule, Sennar Province ], Sudan.

Heterophis resimus = Causus rostratus : Bocage (1870:68).

Causus resimus : Bocage (1887c:211), Ferreira (1904:116), Boulenger (1896:469, 1915:220), Loveridge (1936a:44, 1957:299), Parker (1936:127), Monard (1937b:141), Bogert (1940:97), Manaças (1981:33), Spawls and Branch (1995:142), Spawls et al. (2004:474). Rasmussen (2005:14), Chirio and LeBreton (2007:610), Dobiey and Vogel (2007:128), Akani et al. (2012:189), Wallach et al. (2014:149).

Causus resimus var. angolensis Bocage (1895a:148) . Syntypes: MBL 2025 , 2080 (Biballa) , MBL 2078 , 2084 a–b (Quissange) , MBL 2079 (Novo Redondo) , MBL 2082 a–b (Rio Chimba), 2083a–b (Dondo) (fide McDiarmid et al. 1999) and presumably others, (collectors Banyures [Rio Dande, Rio Bengo], F.A.P. Bayāo [Dondo], A. da Fonseca [Cazengo], F. Newton [Novo Redondo], J.A. d’Anchieta [Quissange, Rio Chimba, Biballa, Maconjo]), destroyed by fire 18 March 1978. Type locality: “ Rio Dande et Rio Bengo ... Dondo ... Cazengo … Novo Redondo … Quissange , Rio Chimba , Biballa et Maconjo. ”

Causus rhombeatus : Hellmich (1957b:74).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species has a strange and seemingly disjunct distribution in Africa, with isolated populations in northern Zaire, eastern Cameroon and the Cameroon-Chad border, Angola, presumably Nigeria, coastal Kenya, Somalia, central and south-western Sudan, and southeastern Ethiopia to Mozambique (see Hughes 1987; Spawls and Branch 1995).

Ocurrences in Angola (Map 278): The species occurs in western Angola , in dry and moist savannas and woodlands of coastal regions. Bengo: “Rio Bengo” [-8.71667,

13.40000] ( Bocage 1895a:146; Monard

1937b:141; Manaças 1981:33); “Cabicula”

[-9.16667, 13.56667] ( Ferreira 1904:116;

Monard 1937b:141; Manaças 1981:33);

“Cunga” [-9.23333, 13.76667] (Manaças

1981:33). Luanda: “ Rio Dande ” [-8.46667,

13.38333] ( Bocage 1895a:146; Monard

1937b:141; Loveridge 1957:299; Manaças

1981:33); Kwanza Norte: “Golungo Alto”

[-9.13333, 14.76667] ( Boulenger 1896:469;

Manaças 1891:33); “Cazengo” [-9.33333,

14.76667] ( Bocage 1895a:146; Ferreira

1904:116; Monard 1937b:141; Manaças

1981:33); “Caculo” [-9.38333, 14.91667] (Fer- MAP 278. Distribution of Causus resimus in Angola. reira 1904:116); “Dondo” [-9.68333, 14.43333] ( Bocage 1895a:146; Monard 1937b:141; Manaças 1981:33). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] ( Hellmich 1957b:74; Manaças 1981:33); “Novo Redondo” [-11.20000, 13.85000] ( Bocage 1895a:146; Monard 1937b:141; Manaças 1981:33); “Quirimbo” [-10.68333, 14.26667] ( Parker 1936:127; Manaças 1981:33); “Congulu” [-10.86667, 14.28333] ( Parker 1936:127; Manaças 1981:33). Benguela: “Quissange” [-12.43333, 14.05000] ( Bocage 1887c:211, Bocage 1895a:146; Monard 1937b:141; Manaças 1981:33); “Catengue” [-13.03333, 13.73333] ( Rasmussen 2005:14); “Hanha” [-12.25000, 13.75000] ( Bogert 1940:97; Manaças 1981:33; Rasmussen 2005:14). Namibe: “Rio Chimba” [-14.30000, 12.40000] ( Bocage 1895a:146; Monard 1937b:141; Manaças 1981:33); “Biballa” [-14.76667, 13.36667 -84] ( Bocage 1895a:146; Monard 1937b:141: Manaças 1981:33); “Maconjo (Macujo)” [-15.01667, 13.20000] ( Bocage 1895a:146; Monard 1937b:141; Manaças 1981:33). Undetermined Locality: “Without precise location” ( Bocage 1870:68).

Taxonomic and distributional notes: No subspecies are currently recognized ( Wallach et al. 2014).

Causus rhombeatus ( Lichtenstein, 1823) RHOMBIC NIGHT ADDER

Sepedon rhombeata Lichtenstein 1823:106 . Syntypes: ZMB 2768-69 (collector G.L.E. Krebs) and possibly ZMB 2770 (donor [?] M.H.C. Lichtenstein, possibly G.L.E. Krebs). Type locality: “Prom. B. sp.” [= Promontorium Bonae Spei = Cape of Good Hope], South Africa. Restricted to “area contained within the magisterial districts of Uitenhage, Kirkwood and Port Elizabeth, Eastern Cape Province, South Africa ” by Bauer (2000:55, 57).

Causus rhombeatus : Günther (1865a:480), Bocage (1866a:51, 1879b:95, 1887a:189, 1887b:207, 1895a:145, 1896a:113, 1897b:211), Peters (1877a:618, 1881:150), Ferreira (1897b:245, 1900a:53, 1906:169), Boulenger (1896:468, 1905:114, 1915:220), Schmidt (1933:15), Parker (1936:126), Loveridge (1936:44), Monard (1937b:141-142), Mertens (1938a:442), Bogert (1940:96), Themido (1941:11), Laurent (1964a:123), Hellmich (1957a:75, 1957b:74), Thys van den Audenaerde (1966:36), Manaças (1973:197, 1981:32), Spawls and Branch (1995:147), Rasmussen (2005:14), Dobiey and Vogel (2007:129), Hampton (2010:235), Broadley (2014:350), Wallach et al. 2014:150), Branch and Conradie (2015:200).

Causus cf. rasmusseni : Conradie et al. (2016:23) .

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species has an extensive range in moist savannas of south, central and east Africa, from eastern Nigeria, southern Sudan and Ethiopia southwards to the Swellendam area in the Western Cape, South Africa, but is absent from much of the arid western half and much of the central part, of southern Africa.

Ocurrences in Angola (Map 279): The species occurs in the entire country with exception of the southern regions and probably

Cabinda (see below). Cabinda:? “Chinchoxo”

[-5.10000, 12.10000] ( Peters 1877a:618; Bocage 1895a:145);? “Landana” [-5.21667,

12.15000] ( Bocage 1895a:145). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] ( Bocage 1887a:189, 1895a:145). Luanda: “Loanda”

[-8.83333, 13.26667] ( Ferreira 1900a:53).

Bengo: “Cacolo até ao Rio Bengo ” (Ferreira

1900:53). Malanje: “ Malanje ” [-9.55000,

16.35000] ( Peters 1881:150; Bocage

1895a:145; Monard 1937b:141); “PungoAndongo” [-9.66667, 15.58333] ( Günther

1865a:480; Boulenger 1905:114; Manaças

1981:33; Rasmussen 2005:14). Lunda Norte:

“Dundo” [-7.36667, 20.83333] (Laurent MAP 279. Distribution of Causus rhombeatus in Angola. 1964a:123; Manaças 1981:32; Thys van den Audenaerde 1966:36; Broadley 2014:350); “Carumbo” [-7.74422, 19.95467] (Branch and Conradie 2015:200); “Cossa” [-7.90000, 21.36667] ( Laurent 1964a:123; Manaças 1981:32; Broadley 2014:350); “Rivière Chicapa, près du pont, 50 km à l’Ouest de Dundo” [-8.10000, 20.51667] ( Thys van den Audenaerde 1966:36); “R. Chimenji, affl. de la Chiumbe” [-7.66667, 20.83333] ( Thys van den Audenaerde 1966:36); “Cassange” [-9.58333, 17.86667] ( Bocage 1895a:145; Monard 1937b:141). Lunda Sul: “Tyihumbwé” [-10.96667, 20.06667] ( Monard 1937b:142). Moxico: “Lago Dilolo” [-11.50000, 22.01667] ( Manaças 1973:197); “environs du lac Calundo” [-11.71667, 20.80000] ( Laurent 1964a:123; Manaças 1981:32; Broadley 2014:350); “Calombe, Luso” [-11.83333, 19.93333] ( Manaças 1973:197); “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] ( Manaças 1973:197). Kwanza Norte: “Cazengo” [-9.33333, 14.76667] ( Ferreira 1900a:53); “Piri-Dembos” [-8.56667, 14.50000] ( Hellmich 1957b:74; Rasmussen 2005:14); “Golungo Alto” [-9.13333, 14.76667] ( Boulenger 1896:468; Ferreira 1906:169; Monard 1937b:141; Rasmussen 2005:14); “Canhoca” [-9.25000, 14.68333] ( Boulenger 1905:114; Rasmussen 2005:14); “Território de Ambaca” [-9.26667, 15.18333] ( Ferreira 1900a:53). Kwanza Sul: “Mombolo” [-12.16667, 14.83333] ( Bogert 1940:96; Manaças 1981:33; Rasmussen 2005:14). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1933:15; Rasmussen 2005:14). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1937b:142); “Bela-Vista” [-12.36667, 16.20000] ( Hellmich 1957b:74; Manaças 1981:33; Rasmussen 2005:14); “ Mt. Moco ” [-12.41667, 15.18333] ( Parker 1936:126); “Santo-Amaro” [-12.70000, 15.85000] ( Monard

1937b:141-142). Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:145; Monard 1937b:141); “ Benguela ” [-12.58333, 13.41667] ( Loveridge 1936a:44); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:145; Monard 1937b:141); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:141-142); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:75; Manaças 1981:33); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:442; Rasmussen 2005:14); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113, 1897b:211); “Cabal” ( Rasmussen 2005:14). Huíla: “Caconda” [-13.73333, 15.0666] ( Bocage 1895a:145; Monard 1937b:141; Rasmussen 2005:14); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937b:141-142); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] ( Monard 1937b:141-142); “Capelongo” [-14.91667, 15.08333] ( Bogert 1940:96; Rasmussen 2005:14); “Huilla” [-15.05000, 13.55000] ( Hampton 2010:235). Namibe: “ Rio Cuce” [-13.51667, 15.20000] ( Ferreira 1897b:245; Monard 1937b:141); “Mossamedes” [-15.20000, 12.15000] ( Bocage 1887a:189, 1895a:145). Cuando Cubango: “Kutatu” [-14.36667, 16.48333] ( Monard 1937b:141-142); “Kakindo” [-15.45000, 17.05000] ( Monard 1937b:141-142); “Kayundu” [-15.70000, 17.45000] ( Monard 1937b:141-142); “riverine habitat Rio Longa (57)” [-15.45969, 18.76833] ( Conradie et al. 2016:8, 12, 23). Undetermined Locality: “Between Benguella and Bihé” ( Boulenger 1905:114; Rasmussen 2005:14).

Taxonomic and distributional notes: Hughes and Barry (1969) and McDiarmid et al. (1999) incorrectly stated that the types were lost, following information from G. Peters. Bogert (1940) pointed out a possible taxonomically relevant difference between the savanna C. rhombeatus (typical form) and its forest dwelling congeners, which was later confirmed by Laurent (1956, 1964a). Laurent (1964a) assigned the Angolan records previously refered to C. rhombeatus (e.g., “Dundo,” “Muita,” “Sombo” [ Laurent 1950a, 1954a], “Quirimbo” [ Parker 1936]) to C. maculatus and also considered specimens cited in Bocage (1895a) from Congo and a part of northern Angolan to be maculatus, without further comment. The species is widely distributed in Angola, however the Cabinda records, “Chinchoxo” and “Landana,” probably represent misidentifications (Spawls and Branch 1995; Rasmussen 2005; Dobiey and Vogel 2007). Conradie et al. (2016) cited a single specimen from “Rio Longa,” Cuando Cubango as Causus cf. rasmusseni Broadley, 2014 , although its taxonomic status remains equivocal, since the topotypic material from C. rasmusseni is not available, and the “Rio Longa” specimen is only weakly differentiated from South African material.