Viperidae

Branch, William R., 2018, When roads appear jaguars decline: Increased access to an Amazonian wilderness area reduces potential for jaguar conservation, Amphibian & Reptile Conservation (e 159) 12 (2), pp. 41-82 : 69-72

Family: Viperidae

Compared with other snake families, African Viperidae have had relatively little taxonomic change, with various new species described (e.g., Bitis harenna, Gower et al. 2016 ; Causus rasmusseni Broadley, 2014 ) or species boundaries readjusted (e.g., revival of Bitis rhinoceros, Lenk et al. 1999 ) but few generic re-arrangements. With the exception of confusion over night adder identifica- tions (see below), few taxonomic changes have affected Angolan vipers. As with the genus Naja , vipers of the genus Bitis had a number of subgenera proposed, and these are adopted in the following species accounts.

Variable Bush Viper

Atheris squamigera (Hallowell 1854)

Echis squamigera Hallowell, 1854 . Descriptions of new reptiles from Guinea. Proc. Acad. Nat. Sci. Philadelphia 1854: 193.

Bocage (1895) knew of no Angolan material except Peters’ (1881) record from Cuango. Later material was recorded from Gulongo Alto ( Ferreira 1904), Piri (Hellmich 1957), and Dundo and Luachimo River ( Laurent 1954, 1964). Atheris squamigera is widespread in the Congo Basin, and a highly variable species. Laurent (1964) only tentatively attributed four specimens from the Dundo region to A. squamigera , as he believed that there were two sympatric species in the lower Congo, and that A. anisolepis may be valid. Recognition of Mocquard’s anisolepis had been problematic, placed first in the synonymy of A. squamigera by Boulenger (1896), revived as a subspecies by Bogert (1940), treated again as a full species by Broadley (1998b), until finally again synonymized by Lawson and Ustach (2000).

Puff Adder

Bitis (Bitis) arietans (Merrem 1820)

Vipera (Echidna) arietans Merrem. 1820 . Versuch eines Systems der Amphibien I (Tentamen Systematis Amphibiorum). J. C. Kriegeri, Marburg: 152.

Although Bocage (1895) noted that the species was common in Angola, there are few records from the southwest region. Günther’s (1865) record from Moçâmedes (= Namibe) may have reflected the collector’s home base.

Horned Adder

Bitis (Calechidna) caudalis (Smith 1839)

Vipera (Cerastes) caudalis Smith, 1839 . Illustrations of the Zoology of South Africa, Reptilia. Smith, Elder, and Co., London: 7.

Although common through the western arid regions of southern Africa ( Branch 1998) there are few Angolan records of the species. Bocage (1867) first signalled its presence (as Cerastes caudalis ) with a specimen from Namibe collected by Anchieta. Subsequent records were added by Bocage (1895, Capangombe, Rio Curoca), and Laurent (1964, “ 35 km south of Namibe ”).

Gaboon Adder

Bitis (Macrocerastes) gabonica (Duméril, Bibron and Duméril 1854)

Echidna gabonica Duméril, Bibron and Duméril 1854 . Erpétologie générale ou histoire naturelle complète des reptiles. Tome septième. Deuxième partie, comprenant l’histoire des serpents venimeux. Paris, Librairie Encyclopédique de Roret: 1428.

At the time of his monograph Bocage (1895) knew this iconic species only from Lândana, Cabinda, and considered that the species was unlikely to occur south of the Kwanza River. However, a year later he reported two specimens from Hanha collected by Anchieta (Bocage 1896). Subsequently others have been recorded: Ferreira (1903), Laurent (1950, 1954, 1964), Tys van den Audenaerde (1967), Oliveira et al. (2016), and Conradie et al. (2017). It is widespread in Angola in suitable forest edge habitat.

Angolan Adder

Bitis (?) heraldica ( Bocage 1889)

Vipera heraldica Bocage, 1889 . Mélanges erpétologiques. II. Sur une vipère apparemment nouvelle d’Angola. Journ. Sci., Lisboa (2) 2: 127.

Bocage (1889) described this endemic Angolan snake from “sur les bords de la rivière Calae, l’un des afflu- ents du Cunene, entre le 13 et le 14 parallèle à l’est de Caconda, Angola ” (= Caluè River, a tributary of the Cunene River, east of Caconda, Huíla District, Angola). Soon after its description the species was incorrectly synonymized with the Bitis peringueyi , a dwarf adder from Namib Desert dune habitat ( Boulenger 1896), leading to over 50 years of confusion. Early records of ‘ B. peringueyi ’ in southern Angola were all based on Boulenger’s synonymization of B. heraldica : i.e., a female from “Between Benguela and Bié ” ( Boulenger 1905), one from Caluquembe ( Monard 1937), and another from “Mombolo, Angola ” from the Vernay, Lang, Boulton, expedition of 1925 ( Bogert 1940). The specific status of Bitis heraldica was revalidated by Mertens (1958) following the collection of a series of Bitis heraldica from Piri (Hellmich 1957) and the species confirmed as endemic to the Angolan highlands. Due to the absence of any fresh genetic material the subgenera status remains uncertain.

Rhinoceros Viper

Bitis (Macrocerastes) nasicornis (Shaw 1802)

Coluber nasicornis Shaw, 1802 . General Zoology or Systematic Natural History, vol. 3, Pt 1. Thomas Davison, London: 94.

Unknown to Bocage (1895) from Angola or Congo, it was first recorded from Angola from scarp forest at Quirimbo ( Parker 1936). That it could be regionally common is shown by Hellmich’s (1957) astonishing record of 53 specimens from Piri. Recent material was recorded from Uige (Ernst et al. 2016).

Two-lined Night Adder

Causus bilineatus ( Boulenger 1905)

Causus bilineatus Boulenger, 1905 . A list of the batrachians and reptiles collected by Dr. W. J. Ansorge in Angola with descriptions of new species. Ann. Mag. Nat. Hist. (7) 16: 114.

Night adders in Angola have been a source of great confusion, with Bocage (1895) and most subsequent authors recognizing only two species in Angola, C. rhombeatus and C. resimus . Bocage (1895), however, did note well-marked adders with lateral stripes from Calandula, Quissange, Caconda, and Huila, but made no taxonomic descision. Boulenger (1905) on receiving additional material from Anchieta noted these same features and proposed the name bilineatus . This name was overlooked by subsequent authors until Laurent (1955) described Causus lineatus for material from DRC. Later, when looking at material from Dundo that he realized was referable to Boulenger’s bilineatus and also conspecific with his DRC material, Laurent (1964) used trinomials and treated his DRC material as the subpecies C. bilineatus lineatus and Angolan material from Calundo, Moxico, as nominotypic C. b. bilineatus . Broadley (1968), based on unpublished analysis in his Ph.D. thesis ( Broadley 1966a), synonymized C. b. lineatus with C. bilineatus , and this was supported by Rasmussen (2005), who reviewed C. bilineatus , re-assessed and mapped Angolan night adders, and corrected many early misidentifications by Hellmich (1957) and Laurent (1964). He noted that C. bilineatus occurred in sympatry with C. rhombeatus in the Benguela-Bié area, Caconda, and Chitau, and with C. rhombeatus and C. maculatus at Piri.

Angola Green Night Adder

Causus resimus (Peters 1862)

Heterophis resimus Peters, 1862 . Über die von dem so früh in Afrika verstorbenen Freiherrn von Barnim und Dr. Hartmann auf ihrer Reise durch Aegypten, Nubien und dem Sennâr gesammelten Amphibien. Monatsber. Akad. Wiss., Berlin 1862: 277.

Bocage (1895) tentatively proposed the name C. resimus, var. angolensis for night adders from several localities in Angola, including Rio Dande, Rio Bengo, Cazengo, Sumbe, Quissange, Rio Chimha, Bibala, and Maconjo. Additional records of C. resimus have also been reported from Cazengo, Caculo, Cabicula ( Ferreira 1904), Quirimbo, Fazenda Congulo ( Parker 1936), and Hanha ( Bogert 1940). Few of these have been accepted by Rasmussen (2005). The Green Night Adder is currently distributed in four isolated populations around the Congo Basin, with the most isolated being that recorded from Angolan scarp forest refugia. The taxonomic status of the Angolan population, and the applicability of Bocage’s (1895) C. resimus, var. angolensis or C. nasalis Stejneger, 1893 , remain unresolved. A preliminary molecular phylogeny of Causus (Tolley et al. in prep.) indicates cryptic diversity within C. resimus , and supports the distinctiveness of Angolan material. Rasmussen (2005) mapped 13 localities for C. resimus in Angola, including one from Cabinda.

Rasmussen’s Night Adder

Causus cf. rasmusseni ( Broadley 2014)

Causus rasmusseni Broadley, 2014 . A new species of Causus Lichtenstein from the Congo /Zambezi watershed in northwestern Zambia ( Reptilia: Squamata : Viperidae ). Arnoldia Zimbabwe 10 (29): 342.

Broadley (2014) described C. rasmusseni based on four specimens from northern Zambia. It is weakly differentiated from sympatric C. rhombeatus by having slightly fewer ventrals in males (130–132, versus 134–150 in Zambian C. rhombeatus ) and reduced blotches or uniform dorsal coloration. Although the un-patterned dorsal pattern and low ventral counts of a specimen from Rio Longa ( Conradie et al. 2016) conform to Broadley’s concept of C. rasmusseni , we caution acceptance of its presence in Angola, and indeed its specific status. A preliminary molecular phylogeny of night adders (Tolley et al., in prep.) supports such caution.

(1895) considered material from Calandula, Quissange, Caconda, and Huila formed “une variété bien caractérisée du C. rhombeatus ” which was later described as C. bilineatus by Boulenger (1905). Rasmussen (2005) mapped the species in Angola, and corrected misidentified material (Hellmich 1957, Laurent 1964), following confusion with C. maculatus in northern populations.

West African Night Adder

Causus maculatus (Hallowell 1842)

Distichurus maculatus Hallowell, 1842 . Description of a new genus of Serpents from Western Africa. J. Acad. nat. Sci. Philadelphia 8: 337.

This species was confused with C. rhombeatus by Bocage, and the first material from Angola was noted by Laurent (1964) from Dundo. Rasmussen (2005) noted the importance of the lateral oblique scale row number in distinguishing between the two species, corrected earlier misindentifications, and showed that C. maculatus occurs in sympatry with both C. rhombeatus and C. bilineatus at Piri. It is restricted to the northern parts of Angola.

Lichtenstein’s Night Adder

Causus lichtensteini (Jan 1859)

Aspidelaps lichtensteinii Jan, 1859 . Additions et rectifications aux Plan et Prodrome de l’Iconographie descriptive des Ophidiens. Rev. Mag. Zool. 11: 511.

Laurent (1964) recorded the first and only specimens of C. lichtensteini in Angola from Dundo and the Lukashi River, 50 km east Dundo. In the Dundo region it occurs in sympatry with both C. rhombeatus and C. maculatus ( Rasmussen 2005) .

Rhombic Night Adder

Causus rhombeatus (Lichtenstein 1823)

Sepedon rhombeata Lichtenstein, 1823 . Verzeichniss der Doubletten des zoologischen Museums der Königl. Universität zu Berlin nebst Beschreibung vieler bisher unbekannter Arten von Säugethieren, Vögeln, Amphibien und Fischen. Königl. Preuss. Akad. Wiss. / T. Trautwein , Berlin : 106.

Bocage consistently used this name ( Bocage 1879, 1880, 1895) and considered it widespread in the interior of Angola. Additional material was noted from Golungo Alto ( Ferreira 1904), Mt Moco, Quirimbo ( Parker 1936), Calundo, Dundo, and Cossa ( Laurent 1964). Bocage

Species not confirmed for Angola that may occur

Hallowell’s House Snake

Boaedon virgatus (Hallowell 1854)

Coelopeltis virgata Hallowell, 1854 . Remarks on the geographical distribution of reptiles, with descriptions of several species supposed to be new, and corrections of former papers. Proc. Acad. Nat. Sci. Philadelphia 1854: 98–105.

Known from both Congo (Brazzaville) and Gabon ( Pauwels and Vande weghe 2008), and therefore likely to occur in forested habitats in Cabinda.

Plain Striped House Snake

Bothrophthalmus brunneus (Günther 1863)

Bothrophthalmus lineatus brunneus Günther, 1863 . Third account of new species of snakes in the collection of the British

Museum. Ann. Mag. Nat. Hist. (3) 12: 348.

Not known to Bocage (1895) from Angola. Although Bothrophthalmus lineatus was recorded from Dundo ( Laurent 1950, 1954, 1964; Tys van den Audenaerde 1967), the plain western subspecies B. l. brunnaeus , from Cameroon to Gabon, is now treated as a full species ( Pauwels and Vande weghe 2008). It is possible that B. brunnaeus may extend south to the Cabinda forests, and those in Angola just south of the border.

Mopane Racer

Mopanveldophis zebrinus ( Broadley and Schätti 2000)

Broadley, D.G. and Schätti, B. 2000. A new species of Coluber from northern Namibia ( Reptilia: Serpentes). Madoqua 19(2): 171.

This unusual and rare snake was described ( Broadley and Schätti 2000) from a single specimen collected near Ruacana on the Cunene River, western Owamboland, Namibia (17°25’S, 14°10’E). It is known from only three other specimens from the Kaokoveld (Bauer et al. 2001), and from the Kunene River mouth ( Cunningham et al. 2018), and was provisionally considered to form part of a Trans-Kunene mopaneveld fauna, and that it may therefore occur in southern Angola. Even at the time of its description, however, its inclusion in the genus Coluber was provisional as the genus was undergoing reassessment and division, particularly the African representatives (e.g., Schätti and Charvet 2003). Schätti and Utiger (2001) erected a monotypic genus for the unusual Socotran racer, Hemerophis socotrae , but deferred a descision on C. zebrinus as its phylogenetic relationships were unknown. However, it was subsequently prematurely placed in Hemerophis ( Wallach et al. 2014) , creating a 3,000 km + zoogeographic enigma, before being finally placed in the monotypic Mopaneveldophis ( Figueroa et al. 2016). This generic name is as non-euphonious as it is a misnomer, as Mopane veld has a much wider distribution than that of the snake, and there is as yet no confirmation that this attractive small racer is restricted to mopaneveld.

Bark Snake

Hemirhaggheris nototaenia (Günther 1864)

Coronella nototaenia Günther, 1864 . Contribution to the knowledge of snakes of tropical Africa. Ann. Mag. Nat. Hist. (6) 1: 309.

Hemirhaggheris in Africa was reviewed by Broadley and Hughes (2000), who showed that all historical Angolan records of H. nototaenia (e.g., Bocage 1895, Monard 1937) were confused with H. viperina . The species may enter extreme eastern Angola as it is recorded from Caprivi ( Broadley and Hughes 2000), the adjacent Ikelenge pedicel in extreme northwest Zambia ( Broadley 1991a), and Ngonye Falls, southwest Zambia (Pietersen et al. 2017).

Lined Grass Snake

Psammophis lineatus (Duméril, Bibron and Duméril 1854)

Dryophylax lineatus Duméril, Bibron and Duméril 1854 . Erpétologie générale ou histoire naturelle complète des reptiles. Tome septième. Deuxième partie, comprenant l’histoire des serpents venimeux. Paris, Librairie Encyclopédique de Roret: 1124.

Its presence in Angola is based on a single record (as Dromophis lineatus, Bogert 1940 ) from “ Angola,” (AMNH 50611, Vernay, Lang, Boulton 1925). The specimen lacks detailed locality data, and members of the Vernay-Lang Angola expedition did not visit eastern or northern Angola, although they visited other areas in Africa after the expedition (Hill and Carter 1940). The species is known from savannah habitats in adjacent countries ( Hughes 2004), but Bogert’s record should be treated with caution until the discovery of additional material. Dromophis was synonymized with Psammophis ( Kelly et al. 2008) .

Boulenger’s Garter Snake

Elapsoidea boulengeri (Boettger 1895)

Elapsoidea boulengeri Boettger, 1895 . Zwei neue Reptilien vom Zambesi. Zool. Anz. 18: 62.

Broadley (1998) mapped parapatry between Elapsoidea boulengeri and E. s. semiannulata from the eastern Caprivi and it is possible that E. boulengeri enters extreme southeast Angola.

Peringuey’s Adder

Bitis peringueyi (Boulenger 1888)

Vipera peringueyi Boulenger, 1888 . On new or little known South African reptiles. Ann. Mag. Nat. Hist. (6) 2: 141.

Haacke (1975) when reviewing the small adders of the western arid of southern Africa recorded no B. peringueyi from Angola, and no confirmed records have subsequently been recorded. Despite this, the species continued to be incorrectly listed for the country (e.g., Branch 1998; Dobiey and Vogel 2007; Uetz and Hozek 2017). Possibly suitable habitat occurs in the small extension of the Namib Desert into extreme southwest Angola, but there remain no records.