Elapidae

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 : 65-69

Family: Elapidae

The taxonomy of African elapids has changed considerably in recent years, particularly among cobras ( Naja ), although the generic relationships of allied species have also been affected and numerous new species have been described ( Broadley 1968, 1995; Broadley and Wüster 2004; Wüster and Broadley 2003, 2007; Wallach et al. 2009; Ceriaco et al. 2017; Wüster et al. 2018). Water cobras (Boulengerina) were demonstrated to be closely related to the Forest Cobra ( Naja melanoleuca ) and therefore synonymized with Naja ( Nagy et al. 2005; Wüster et al. 2007). Wallach et al. (2009) proposed four subgenera within Naja , with the subgenus Naja restricted to Asia, and with the three other subgenera used for African cobras, and Angolan species assigned to various subgenera: i.e., Naja (Boulengerina) melanoleuca , N. (Uraeus) anchietae , N. (Afronaja) mossambica , N. (Afronaja) nigricollis , and N. (Afronaja) n. nigricincta . Moreover, a recent revisions of forest cobras ( Ceriaco et al. 2017; Wüster et al. 2018) recognize a suite of five species, with typical N. (B.) melanoleuca entering northern Angola and the revived N. (B.) subfulva occuring in central Angola.

Coral Shield Cobra

Aspidelaps lubricus cowlesi ( Bogert 1940)

Aspidelaps lubricus cowlesi Bogert, 1940 . Herpetological results of the Vernay Angola Expedition. I. Snakes, including an arrangement of the African Colubridae . Bull. Amer. Mus. Nat. Hist. 77: 94.

Bogert (1940) described Aspidelaps lubricus cowlesi from a snake collected from Munhino (101 km east of Namibe, via railroad). Originally considered endemic to Angola, Mertens (1971) extended its range into the Kaokoveld. Broadley and Baldwin (2006) relegated it to the synonymy of the northern Namibia subspecies A. l. infuscatus, citing intermediates between A. l. cowlesi and A. l. infuscatus. Unusually they still presented an account for A. l. cowlesi even though synonymizing it. Moreover, no morphological data was presented to show that even A. l. infuscata was a valid taxon, or a molecular data presented to support the new taxonomic arrangement.

eastern subspecies D. j. kaimosae needs genetic assessment. Recent material was discussed by Vaz Pinto and Branch (2015).

Black Mamba

Dendroaspis polylepis (Günther 1864)

Dendraspis polylepis Günther, 1864 . Report on a collection of reptiles and fishes made by Dr. Kirk in the Zambesi and Nyassa Regions. Proc. Zool. Soc., London 1864: 310.

All early records (up to 1946) from Angola were discussed under D. angusticeps ( Peters 1881; Bocage 1888, 1895; Monard 1937; Bogert 1940), until D. polylepis was shown to be a valid species ( FitzSimons 1946). The species is nowhere common, but is relatively well-known in the central and southern regions. Bocage (1895) recorded it from the “hauts-plateaux de l’intérieur d’Angola,” although Bogert (1940) recorded it from Hanha. Fig. 29. Dendroaspis jamesoni, Gabela , Cuanza Sul.

Jameson’s Mamba

Dendroaspis jamesoni (Traill 1843)

Elaps jamesoni Traill, 1843 . Description of the Elaps jamesoni , a new species of serpent from Demerara. Edinburgh New. Phil. J. 34 (67): 53.

The identification of historical records of Angolan mambas is complicated by confusion between green and black mambas in the early literature. Prior to 1946 the Black Mamba ( D. polylepis ) was considered a juvenile of the southern Green Mamba ( D. angusticeps ), and recognition of the Black mamba as a separate species was only confirmed by FitzSimons (1946). Bocage (1888) discussed the identification of mambas, reducing the number of species then known from seven to three, whilst also describing a new, overlooked species from Angola, D. neglectus (hence the unusual name). He gave detailed scale counts of the material he examined, and also presented a diagnostic key to the species recognized. From this it is evident that he applied the existing names wrongly: His D. jamesoni is now identified as D. viridis ; his D. angusticeps is really D. polylepis ; and his new species D. neglectus was already known as D. jamesoni . This arrangement continued to be reflected in his monograph (1895), where two species of mamba were correctly considered to occur in Angola, both unfortunately incorrectly named. Dendroaspis jamesoni has been recorded from Pungo-Andonga (the type of Dendroaspis welwitschii Günther, 1865 , which Bocage considered a synynoym of his own D. neglectus even though Günther’s name had priority). As D. neglectus its was recorded from north of the Kwanza River ( Bocage 1888) and from N’Dalatando ( Ferreira 1903); and recorded correctly as Dendroaspis jamesoni jamesoni from Piri, Bela Vista (Hellmich 1957) and Dundo ( Laurent 1954), although the status of the Günther’s Garter Snake

Elapsoidea guentherii (Bocage 1866)

Elapsoidea guntherii Bocage, 1866 . Lista dos reptis das possessões portuguezas d’Africa occidental que existem no Museu Lisboa. Jorn. Sci. Math. Phys. Nat., Lisboa 1: 50.

Bocage (1866) described both the genus and species Elapsoidea guentherii in one of his first herpetological papers. The description was based on material sent by Anchieta from Cabinda, and led to the long association beween these two icons of Angolan herpetology. Another adult from Bissau was included in the description, and to avoid confusion Parker (1949) later restricted the type locality to Cabinda. Additional material was discussed by Bocage (1895), Loveridge (1936), Bogert (1940), Hellmich (1957), Laurent (1964), and Broadley (1971b) who summarized its range from the northern parts of Angola, through Zambia to Zimbabwe.

Angolan Garter Snake

Elapsoidea semiannulata semiannulata (Bocage 1882)

Elapsoidea semi-annulata Bocage, 1882 . Reptiles rares ou nouveaux d’Angola. Journ. Sci., Lisboa 8(32): 303.

After describing the previous species, Bocage (1882) described Elapsoidea semiannulata from additional material from Caconda, but in his monograph ( Bocage 1895) treated his new species as E. guentheri var. semiannulata . Laurent (1964) described Elapsoidea decosteri huilensis from Humpata, which was relegated to the synonym of E. s. semiannulata by ( Broadley 1971b). However, Broadley (1971b) also treated a species described by Werner (1897) from Ghana as a northern race, Elapsoidea semiannulata moebiusi , that extended to Gabon but was unrecorded from Cabinda or northern Angola. All of Bocage’s northern localities (1866, 1895, 1897) were restricted to Bissau, but Broadley (1998) reassessed some of Laurent’s Congo-Kinshasa material and reassigned it to E. s. moebiusi. Although Broadley (1998) described the range of E. s. moebiusi as extending into northern Angola, he neither mapped nor noted any localities supporting this claim. His closest locality was a poorly defined “Bas Congo ” (Broadley 1998), and the race remains unknown from Angola, including Cabinda. Haacke and Finkeldey (1967) recorded the first record of E. s. semiannulata from southern Africa. Broadley (1971b) summarized and mapped the species in Angola, and recognized E. s. boulengeri Boettger, 1895 , from Mozambique as an eastern race, which he later raised to a full species (Broadley 1998). This is unrecorded from Angola, although known from adjacent regions in Namibia and Zambia, and parapatry between E. s. semiannulata and E. boulengeri occurs in the Caprivi area.

Anchieta’s Cobra

Naja (Uraeus) anchietae ( Bocage 1879)

Naja anchieta e Bocage, 1879. Reptiles et batraciens nouveaux d’Angola. J. Acad. Sci., Lisbon 7: 89.

As with the previous species, Bocage (1879) also described Naja anchietae from snakes sent from Caconda. Mertens (1937) relegated it to a subspecies of the Egyptian Cobra ( N. haje anchietae ), Broadley (1995) referred southern populations of N. haje to the Snouted Cobra ( N. annulifera ), retaining N. a. anchietae as a western race. Finally, Broadley and Wüster (2004) revalidated Bocage’s original name. Angolan specimens have thus been decribed under various names. Broadley and Würster (2004) refer Bocage’s (1895) N. haje material to N. anchietae . Known records were summarized and mapped in Broadley (1995) and Broadley and Wüster (2004), and its range was extended north to Capanda Dam by Ceriaco et al. (2014a).

Banded Water Cobra

Naja (Boulengerina) annulata (Peters, 1876) Naja annulata Buchholz and Peters in Peters, 1876. Eine zweite Mittheilung über die von Hrn. Professor Dr. R. Buchholz in Westafrica gesammelten Amphibien. Monatsber. königl. Akad. Wiss. Berlin 1876 (February): 119.

Bocage (1985) mentioned Naja annulata from the Congo, but knew of no Angolan records. The first and only record for the country came from Lagoa Carumbo region (Branch and Conradie 2012). Wüster et al. (2018) noted that Aspidelaps bocagei Sauvage, 1884 (type locality: Gabon and Majumba) is not a synonym of N. melanoleuca , as recorded by various authors, but instead of Naja annulata .

Central African Forest Cobra

Naja (Boulengerina) melanoleuca (Hallowell 1857)

Naja haje var. melanoleuca Hallowell 1857 . Notes of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proc. Acad. Nat. Sci. Philadelphia 9: 61.

Bocage (1895) had a very different understanding of species boundaries within African cobras compared with modern nomenclature, but the loss of so much of his material has made assignment of his records to modern species difficult. Much of his cobra material was grouped as varieties under the spitting cobra N. nigricollis , e.g., Naja nigricolls var. melanoleuca . Fortunately, Broadley (1974) examined this material before it was destroyed in the Lisbon Museum fire (1978) and corrected the identity of the individual apecimens of which none was referrable to true Naja melanoleaca (Hallowell) . To stabilize taxonomy Broadley (1974) designated MBL 1972 from “Caconda” as the lectotype of Bocage’s (1895) Naja nigricolls var. melanoleuca , and treated it as a synonym of N. nigricollis . Whether Bocage knew true forest cobras from Angola remains debatable, but subsequent records confirming its presence are from Pungo Andongo ( Boulenger 1905), Piri, Calulo, Sangeunge (Hellmich 1957), Dundo ( Laurent 1954, 1964) and Soyo ( Wüster et al. 2018). Genetic material from the latter validated the presense of true N. melanoleuca in northwest Angola ( Wüster et al. 2018). Laurent (1964) considered that true N. melanoleuca in northeast Angola was probably confined to forest galleries with N. subfulva living in secondary forests and the intervening savannas.

Brown Forest Cobra

Naja (Boulengerina) subfulva ( Laurent 1955)

Naja melanoleuca subfulva Laurent, 1955 . Diagnoses preliminaires des quelques Serpents venimeux. Rev. Zool. Bot. Afr. 51: 127–139.

The taxonomic status of the different populations of the forest cobra has long been problematic. Don Broadley (in litt., July 2013), in conjunction with Wolfgang Wüster and colleagues, recognized at least six species-level taxa within the forest cobra complex, including an endemic species on São Tome. Description of these cryptic taxa was delayed by Broadley’s illness, and the São Tome population subsequently described as Naja (Boulengerina) peroescobari ( Ceriaco et al. 2017) . The delay also led to some nomenclatural confusion for other forest cobra populations, with various authors prematurely adopting some of the taxonomic findings. Broadley and Blaylock (2013) referred southern populations to Laurent’s (1955) savannah-inhabiting subspecies N. m. subfulva , and not- ed that unpublished molecular data indicated it deserved specific status. They also used subgenera within Naja as proposed by Wallach et al. (2009), but some authors subsequently elevated these subgenera to full genera, e.g., Wallach et al. (2014) and Ceriaco et al. (2016b), even though the intention of using subgenera was to avoid medical issues possibly arising from nomenclatural changes in the names of medically important snakes. Chirio and Ineich (2006) and Ceriaco et al. (2017) also revived N. subfulva , the latter based on genetic divergence of Mozambique material. Wüster et al. (2018) mapped populations of N. subfulva from large parts of Angola. However, the assignment was presumably based on historical aspects, as no analysis was presented for Angolan N. subfluva morphology or genetic divergence compared with other populations. The status and distribution of N. subfulva in Angola therefore requires further study.

Mozambique Cobra

Naja (Afronaja) mossambica (Peters 1854)

Naja mossambica Peters, 1854 . Diagnosen neuer Batrachier, welche zusammen mit der früher (24. Juli und 17. August) gegebenen Übersicht der Schlangen und Eidechsen mitgetheilt werden. Ber. Bekanntmach. Geeignet. Verhandl. Königl.-Preuss. Akad. Wiss., Berlin 1854: 625.

Broadley (1974) first signaled the presence of N. mossambica in Angola when he identified a specimen from Maconjo (MBL 1964, now lost) that Bocage (1895) had referred to N. nigricollis var. fasciata . Broadley (1968) treated mossambica as a full species after finding sympatry between N. nigricollis and N. mossambica in Zambia. The species is poorly known in Angola and the few records are restricted to the southern provinces ( Conradie et al. 2016).

Western Barred Spitting Cobra

Naja (Afronaja) nigricincta nigricincta ( Bogert 1940)

Naja nigricollis nigricincta Bogert, 1940 . Herpetological results of the Vernay Angola Expedition. I. Snakes, including an arrangement of the African Colubridae . Bull. Amer. Mus. Nat. Hist. 77: 89.

Broadley (1974), after examination of Bocage’s original material, assigned Bocage’s (1895) N. nigricollis var. fasciata to Bogert’s N. nigricollis nigricinta , and designated MBL 1968 (now lost) from Benguela as the lectotype. He was also unable to find Bocage’s var. fasciata from Dondo, and this record may be in error. Molecular studies later supported the elevation of N. nigricinta to a full species, with a southern subspecies N. n. woodi (Wuster et al. 2007). However, no Angolan material was includ- ed in this analysis and the status of Namibian material needs confirmation, as the type locality for N. nigricincta is “Munhino (101 km east of Mossamedes via railroad,” and nominotypical Angolan N. nigricinta have a different color pattern to those in Namibia.

Black Spitting Cobra

Naja (Afronaja) nigricollis (Reinhardt 1843)

Naja nigricollis Reinhardt, 1843 . Beskrivelse af nogle nye Slangearter. Danske Vidensk. Selsk. Afhandl. 10: 369.

Broadley (1968) had reinstated N. mossambica as a full species after confirming sympatry with N. nigricollis in eastern Zambia. He treated pallida, katiensis, nigricincta and woodi as subspecies of N. mossambica , whilst various other names, e.g., crawshayi, occidentalis, and atriceps were considered synonyms of N. nigricollis . Later, Broadley (1974) assigned Bocage’s (1895) N. nigricollis var. occidentalis to N. nigricollis nigricollis , and designated MBL 1963 (now lost) from Dondo as the lectotype. He also noted that Naja nigricollis var. melanoleuca Bocage, 1895 , described from Angola, was preoccupied by N. melanoleuca (Hallowell) . Finding additional sympatry between N. mossambica and N. nigricollis nigricincta in Angola, he treated nigricincta and the all-black woodi from western South Africa and southern Namibia as subspecies of N. nigricollis . Following a molecular analysis of African spitting cobras, Wuster et al. (2007) validated N. nigricinta as a valid species, with N. nigricincta woodi as a southern race. The Black Spitting Cobra, N. nigricollis , therefore reverted to binomials. It is widespread in Angola, but generally absent from closed-canopy forest.

Gold’s Tree Cobra

Pseudohaje goldii ( Boulenger 1895)

Naja goldii Boulenger, 1895 . On some new or little-known reptiles obtained by W. H. Crosse Esq. on the Niger. Ann. Mag. Nat. Hist. (6) 16: 34.

Parker (1935, as Naja goldii ) recorded the first examples of this arboreal cobra from south of the Congo River and for Angola. The specimens were collected from remnant scarp forest near Quirimbo. The only other historical Angolan material is from Peri (Hellmich 1957) and Dundo ( Laurent 1950, 1954).