Psammophiinae

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 : 60-63

Subfamily: Psammophiinae

The psammophiine genera ( Dipsina , Hemirhagerrhis , Malpolon , Psammophis , Psammophylax , Rhamphiophis ) are distributed throughout Africa, the Middle East, south-central Asia, and southern Europe ( Branch 1998; Kelly et al. 2008), with Mimophis restricted to Madagascar. Their monophyly is supported by morphological and molecular data ( Cadle 1994; Brandstätter 1996; Zaher 1999; Vidal and Hedges 2002 a; Kelly et al. 2008). Two recent genera have been synonymized: Dromophis with Psammophis ( Kelly et al. 2008) and Rhagerhis with Malpolon ( Figueroa et al. 2016) . Figueroa et al. (2016) consider Asian Psammodynastes to also be part of the Psammophiidae , but this species has a bifurcate, heavily ornamented hemipenis (see Fig. 17 in Zaher 1999) that contrasts with the well established synapomorphy of the simple, tubular and unadorned hemipenes of psammophines. We therefore continue to exclude Psammodynastes from the Psammophidae .

Viperine Rock Snake

Hemirhagerrhis viperina (Bocage 1873)

Psammophylax viperinus Bocage 1873 . Bocage JVB. 1873. Melanges erpetologiques. II. Sur quelques reptiles et batraciens nouveaux, rares ou peu connus d‘Afrique occidentale. Jorn. Acad. Sci., Lisboa 4: 222.

Bocage (1873) described Psammophylax viperinus from Dombe Grande, and later added further Anchieta material from Humbe, Maconjo, and Capangombe ( Bocage 1895), but deferred to Boulenger in treating this material as Psammophylax nototaenia . Bogert (1940) added new records from the Vernay Lang Angola expedition from Hanha, Huambo, and Munhino, and revived it as a subspecies, H. nototaenia vipernua , for all Angolan material. Laurent (1964) recorded a specimen from Humpata. Broadley (2000) revived H. viperina again as a full species, and noted no H. nototaenia from Angola.

Angolan Sand Snake

Psammophis angolensis ( Bocage 1872)

Amphiophis angolensis Bocage, 1872 . Diagnoses de quelques espéces nouvelles de Reptiles d’Afrique occidentale. Jorn. Sci., Lisbon 4: 82.

This dwarf psammophine was described by Bocage (1872) from “Dondo, (intérieur d’Angola).” Wallach et al. (2014) discuss confusion over the correct location of Bocage’s type locality “Dondo,” which has been presented in various forms; e.g., Danda, Loanda Distr. ( Loveridge 1940; Broadley 1962), Donda, Loanda Distr. ( FitzSimons 1962; Auerbach 1987), and Dondo, Luanda Distr. ( Crawford-Cabral and Mesquitela 1989). They note that it was correctly located by Loveridge (1957) and Hellmich (1957a) as Dondo, north bank of Kwanza River, SW Cuanza Norte Distr., NW Angola (09°41’S, 14°26’E, elevation 50 m). Additional material was not- ed ( Bocage 1895) from Caconda, Quindumbo, Humbe, Sumbe, Pungo-Andongo and Ambrizette (=N’Zeto), and later from Dundo, Cameia, and Lagoa Calundo ( Laurent 1964).

Ansorge’s Sand Snake

Psammophis ansorgii ( Boulenger 1905)

Psammophis ansorgii 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: 113.

Boulenger (1905) described Psammophis ansorgii on a snake collected by Anchieta between “Benguella to Bihe, Angola.” Hellmich (1957) recorded the species from Bela Vista, but of his seven specimens, five were just heads. However, this new material allowed him to note that the preocular was, at most, only in narrow contact with the frontal, and he therefore removed the species from the synonymy of P. jallae , where it had been placed by Loveridge (1940). This was followed by Broadley (1977, 2002), who noted no new material. Branch et al (2018a) discussed new material and phylogenetic relationships, and noted that it is one of the few snakes endemic to Angola.

Jalla’s Sand Snake

Psammophis jallae (Peracca 1896)

Psammophis jallae Peracca, 1896 . Retili et Anfibi raccolti a Kazungula e sulla strada da Kazungula a Buluwaio dal Rev. Luigi Jalla, Missionario Valdese nell’ alto Zambese. Boll. Mus. Zool. Comp. Anat. Univ. Torino 11(255): 2.

Loveridge (1940) removed P. jallae from the synonymy of P. crucifer , and Broadley (1977, 2002) discussed additional material, although noting that it remained known from Angola only from the type of Psammophis rohani Angel, 1921 (Lumuna River). A second Angolan record was noted by Conradie et al. (2017).

Leopard Sand Snake

Psammophis leopardinus (Bocage 1887)

Psammophis sibilans var. leopardinus Bocage 1887 . Melanges erpetologiques. I. Reptiles et Batraciens du Congo. II. Reptiles de Dahomey. III. Reptiles de l’Ile du Prince. IV. Reptiles et Batraciens de Quissange (Benguella) envoyés par M. J. d’Anchieta. Jorn. Sci. Math. Phys. Nat., Lisboa 11: 206.

Bocage (1887b) was confronted with diverse color patterns and scalation in sand snakes, and like many subsequent researchers ( Boulenger 1895; Loveridge 1940; Broadley 1977, 2002) found it difficult to resolve species boundaries. As was customary at the time he used varieties to characterize intraspecific grouping. Some of these have subsequently been demonstrated, both morphologically ( Broadley 1977, 2002) and genetically ( Kelly et al. 2008) to be valid species. One such is this species, first described by Bocage (1887) as a variety of P. sibilans (now restricted to North Africa), eventually as a western subspecies of P. brevirostris ( Broadley, 1977) , before being raised to a full species ( Broadley 2002). The latter lists numerous localities for western Angola, including the type locality Catumbela.

Namib Sand Snake

Psammophis namibensis (Broadley 1975)

Psammophis leightoni namibensis Broadley, 1975 . A review of Psammophis leightoni and Psammophis notosticus in southem Africa (Serpentes: Colubridae ). Arnoldia 7 (13): 9.

Some early Bocage Psammophis material from Mossamedes (= Namibe, MBL 1809) and Rio Curoca (MBL 1810) was studied by Broadley before the Lisbon fire. He assigned them to a new taxon, described initially as a western arid subspecies (Broadley 1975) of P. leightoni , but later treated as a full species, P. namibensis ( Broadley 2002) . Additional material, collected by Haacke from Cunene Forde 15 km NE, Iona Res, Foz do Cunene and Pico de Acezevedo, was also assigned to this species ( Broadley 2002). It has a much greater distribution in the western arid region, through Namibia to South Africa.

Karoo Sand Snake

Psammophis notostictus (Peters 1867)

Psammophis moniliger var. notostictus Peters 1867 . Peters, Wilhem Carl Hartwig 1867. Über eine Sammlung von Flederthieren und Amphibien aus Otjimbingue in Südwestafrica, welche Hr. Missionär Hahn dem zoologischen Museum zugesandt hat. Monatsber. königl. Akad. Wiss., Berlin 1867 (April): 237.

Bocage (1887) described Psammophis sibilans , var. nova. stenocephalus from “l’intérieur de Mossamedes” (= Rio Curoca), but this was later synonymized with P. notostictus by Broadley (1977) and in Angola is known from only a few localities in Namibe Province, but extending south to the Karoo of South Africa. Broadley (2002) noted material from Rio Sau Nicolau (= Rio Bentiaba), and Ceriaco et al. (2016a) from Espinheira and Pico de Azevedo.

Mozambique Grass Snake

Psammophis mossambicus (Peters 1882)

Psammophis sibilans var. mossambica Peters 1882 . Naturwissenschaftliche Reise nach Mossambique auf Befehl seiner Majestät es Königs Friedrich Wilhelm IV. in den Jahren 1842 bis 1848 ausgefeführt von Wilhelm C. Peters. Zoologie III. Amphibien. Berlin (Reimer), 122.

The large sand snakes of the Psammophis sibilansphilippsi-mossambicus complex have remained a recurrent taxonomic problem among African snakes for over 100 years; see discussion and shifting nomenclature in Bocage (1887, 1895), Boulenger (1895), Loveridge (1940), Broadley (1977, 2002), Hughes (1999), and Brandstätter (1995, 1996). Most Angolan Psammophis have previously been included in varied taxa within this complex. Broadley (2002) lists numerous localities for this species, which is widespread in savannah and secondary habitats in Angola. Whether all Angolan large sand snakes are referable to P. mossambicus , or whether northwest populations have affinities with West African P. phillippsi , requires fresh material and genetic assessment.

Western Strip-bellied Sand Snake

Psammophis subtaeniatus (Peters 1882)

Psammophis sibilans var. subtaeniata Peters 1882 . Naturwissenschaftliche Reise nach Mossambique auf Befehl seiner Majestät es Königs Friedrich Wilhelm IV. in den Jahren 1842 bis 1848 ausgefeführt von Wilhelm C. Peters. Zoologie III. Amphibien. Berlin (Reimer), 121.

Bocage (1895) recognized five forms of Psammophis sibilans (vars. A–E), of which var. A he contrasted with Peters’ var. subtaeniata . He noted that his specimens from Rio Bengo, Catumbela, Bibala, Maconjo, Humbe and Cunene, all collected by Anchieta, looked like var. subtaeniata . Broadley (2002) list numerous localities for P. subtaeniatus in Angola, including many from the Lisbon Museum subsequently lost in the fire. The species is restricted to the semi-arid scrubland and mopane woodland, above and below the escarpment in southwest Angola. Broadley (2002) includes Psammophis bocagii Boulenger, 1895 as a synonym.

Western Sand Snake

Psammophis trigrammus (Günther 1865)

Psammophis trigrammus Günther, 1865 . Fourth account of new Species of Snakes in the Collection of the British Museum. Ann. Mag. Nat. Hist. (3) 15: 95.

For many years this snake was known only from the type locality (“São Nicolau, Mossamedes”). Bocage (1887) seemed to consider it distinct, but only as a variety of Psammophis sibilans and not identical to the other varieties he described at the time (e.g., stenocephalus and leopardinus ). Later he seems to have overlooked the species as he did not discuss it in his monograph ( Bocage 1895). This may be why Monard (1937) also overlooked it, as its taxonomic status has never been challenged. Broadley (2002) plotted the limited extenstion into southwest Angola.

Zambezi Sand Snake

Psammophis zambiensis (Hughes and Wade 2000)

Psammophis zambiensis Hughes, 2000 . On the African leopard whip snake, Psammophis leopardinus Bocage, 1887 (Serpentes, Colubridae ), with the description of a new species from Zambia. Bull. nat. Hist. Mus. Lond (Zool.) 68(2): 75.

Only recently described ( Hughes and Wade 2002), the species was first reported from Angola during the NGOWP surveys in the headwater region of the Angolan Okavango catchment (Conradie et al. 2017; Conradie and Branch 2017).

Striped Beaked Skaapstekker

Psammophylax acutus ( Günther 1888)

Psammophis acutus Günther, 1888 . Contribution to the knowledge of snakes of tropical Africa. Ann. Mag. Nat. Hist. (6) 1: 327.

First recorded from Angola by Bocage (1873) from material sent by Capello and Ivens from Cassange (= Baixa de Cassange). Unfortunatey he misidentified it as Rhagerhis tritaeniata and later, after Günther (1888) had described it as a new species on a specimen from Pungo Andongo, apologized to the collectors ( Bocage 1895) and noted more specimens from Caconda and Huila. Additional material was noted from Benguela to Bié ( Boulenger 1905), Bela Vista (Hellmich 1957), Alto Chicapa, Alto Cuilo, and Dundo ( Laurent 1964). Broadley (1971c) revised the species, which was later transferred to Psammophylax ( Kelly et al. 2008) .

Huila Skaapstekker

Psammophylax ocellatus (Bocage 1873)

Psammophylax ocellatus Bocage 1873 . Melanges erpetologiques. II. Sur quelques reptiles et batraciens nouveaux, rares ou peu connus d‘Afrique occidentale. Jorn. Acad. Sci., Lisboa 4: 221.

Bocage (1873) described Psammophylax ocellatus on the basis of an adult snake from “l’intérieur de Mossamedes (Gambos).” Additional material was recorded (as P. rhombeatus ) from Humbe ( Bocage 1895; Boulenger 1896) and Tundavala ( Baptista et al. 2018a). Broadley (1977) reviewed the Angolan material and revived P. r. ocellatus as a northern subspecies. Branch et al. (2018a) extending the known range to the Chela escarpment region and also validated P. ocellatus as a full species.

Striped Skaapstekker

Psammophylax tritaeniatus (Günther 1868)

Rhagerrhis tritaeniatus Günther, 1868 . Sixth account of new species of snakes in the collection of the British Museum. Ann. Mag. Nat. Hist. (4) 1: 423.

First recorded from Dondo by Bocage (1873), and later from the Rio Kwanza, Quissange, Cahata, Quindumho, Caconda, Huila, Gambos, and Humbe by Bocage (1895), who noted it was one of the most common and widespread snakes on the highlands of Angola. Additional records from Cuvango, Mupanda, Vila da Ponte (= Cuvango) ( Monard 1937), Capelongo ( Bogert 1940), Alto Cubal (Hellmich 1957), Cazombo, Forte Roçadas ( Laurent 1964), and Calombe ( Manaças, 1973). Broadley (1977) reviewed the genus.

Figures

Fig. 17. Boaedon capensis-fuliginosus-lineatus complex, near Menongue, Cuando Cubango.