Psammophis mossambicus, (Peters, 1882)

Trape, Jean-François, Crochet, Pierre-André, Broadley, Donald G., Sourouille, Patricia, Mané, Youssouph, Burger, Marius, Böhme, Wolfgang, Saleh, Mostafa, Karan, Anna, Lanza, Benedetto & Mediannikov, Oleg, 2019, On the Psammophis sibilans group (Serpentes, Lamprophiidae, Psammophiinae) north of 12 ° S, with the description of a new species from West Africa, Bonn zoological Bulletin 68 (1), pp. 61-91 : 73-75

PSAMMOPHIS MOSSAMBICUS (Peters, 1882)

Olive Whip Snake, Psammophis olivâtre, Olivenfarbige Sandrennnatter

Psammophis sibilans (not Linnaeus) Mocquard, 1887: 78 (Brazzaville); Bocage, 1895: 114 (part, var. C); Tornier, 1896: 82; Boulenger, 1905: 113; 1906: 214 (part, Fernand Vaz); Sternfeld, 1911: 250; Boulenger, 1915a: 213; 1915b: 631; Loveridge, 1916: 85; 1918: 328; Sternfeld, 1917: 478; Schmidt, 1923: 111; Loveridge, 1923: 886; Loveridge, 1928: 39; 1933: 255; Witte, 1933a: 123; 1933b: 93; Corkill, 1935: 20 (part); Loveridge, 1936a: 38; 1936b: 262; Uthmoller, 1937: 119; Mertens, 1955: 59; Laurent, 1960: 55; Trape & Roux-Estève, 1995: 41.

Psammophis sibilans var. mossambica Peters, 1882 , Reise nach Mossambique ...3: 122. Type locality: Mozambique Island. Lectotype: ZMB 2468A.

Psammophis sibilans var. tettensis Peters, 1882 , Reise nach Mossambique ...3: 122. Type locality: Tete, Mozambique.

Psammophis sibilans var. intermedius Fischer, 1884: 14 (Arusha, Tanzania).

Psammophis irregularis (not Fischer) Sauvage, 1884: 201.

Psammophis sibilans var. occidentalis Werner, 1919: 504 . Type locality: Congo. Holotype NMW 19245:2.

Psammophis notostictus (not Peters) Witte, 1933a: 123; 1933b: 93.

Psammophis brevirostris (not Peters) Witte, 1933a: 123; 1933b: 93.

Psammophis sibilans phillipsii Loveridge, 1940: 41 (part); Witte 1962: 117 (part); Perret 1961: 136; Böhme 1975: 40; Stucki-Stirn, 1979: 434.

Psammophis subtaeniatus (not Peters) Witte, 1941: 212 (part, Bitshumbi, P.N. Virunga); Joger, 1990: 97, Fig. 6 (Bangui, CAR).

Psammophis sibilans sibilans (not Linnaeus) Bogert, 1940: 79 (part); Loveridge, 1940: 30 (part); 1942a: 110; 1942b: 8; 1956: 48 (part); Witte, 1941: 213; 1955: 220; 1962: 117; 1975: 87; Laurent, 1950: 8; 1954: 59; 1956: 249, Pl. xxiv, fig. 1; 1964: 113; Monard, 1951: 164; Villiers, 1966: 1765; Vesey-FitzGerald, 1958: 60; Robertson et al., 1962: 428; Bourgeois, 1964: 81; Roux-Estève, 1965: 72; Thys van den Audenaerde, 1965: 381 (Kinshasa); Broadley, 1971: 88 ( Zambia); Pitman, 1974: 156; Stucki-Stirn, 1979: 430; Pakenham, 1983: 26; Spawls, 1978: 8; Rasmussen, 1991: 177.

Psammophis subtaeniatus sudanensis (not Werner) Laurent, 1956: 248, Pl. xxiv, fig. 2 (part, Bitshumbi); Perret, 1961: 136; Witte, 1962: 117; 1975: 88.

Psammophis phillipsi Joger, 1982: 331 , 1990: 97; Hughes, 1983: 346, 353 (part); Butler & Reid, 1990: 32 (part); Broadley, 1991: 530; Broadley & Howell, 1991: 28; Rasmussen, 1991: 177; Brandstätter, 1995: 75 (part), 1996: 55 (part); Trape & Roux-Estève, 1995: 41; Chippaux, 1999: 164 (part); Luiselli et al., 2004: 415 (part); Chirio & Ineich, 2006: 52: Jackson et al., 2007:77.

Psammophis mossambicus Branch, 1998: 92 ; Haagner et al., 2000: 15; Hughes & Wade, 2002: 77; Spawls et al., 2002: 405; Broadley et al., 2003: 167, Pl. 110–1; Marais, 2004: 153; Bates et al., 2014: 377.

Psammophis phillipsi occidentalis Hughes & Wade, 2004: 129 , Fig. 1 .

Psammophis occidentalis Chirio & LeBreton, 2007: 532 ; Wallach et al., 2014: 589.

Psammophis sp.1 Chirio & LeBreton, 2007: 538.

Psammophis cf. phillipsi Pauwels & Vande weghe, 2008: 220 .

Description. (431 specimens examined) Nostril pierced between 2 (rarely 3) nasals; preocular 1, usually widely separated from frontal; postoculars 2 (rarely 1 or 3); temporals usually2+2/3 but fusions or partial fusions frequent; supralabials 8 (very rarely 6, 7 or 9), the fouth and fifth & (rarely third & fourth or fifth and sixth) entering orbit; infralabials usually 10 (rarely 9 or 11), the first 4 (rarely 5, e.g. sequenced specimen IRD 2226.N) in contact with anterior sublinguals; dorsal scales in 17-17-13 rows; ventrals 154–188; cloacal divided (rarely entire); subcaudals 84–122.

Dorsum brown or greenish brown often uniform ( Fig. 22 ), sometimes yellowish posteriorly, sometimes with scattered black scales (rarely more black scales than olive ones). Other specimens have black-edged dorsal scales, a vertebral chain and a yellow or whitish dorsolateral stripe on scale rows 4 and 5 ( Fig. 23 ), this pattern almost constant in specimens from coastal areas of Gabon, Republic of the Congo and Democratic Republic of the Congo. Top of head uniform or reticulated, which fades out in adults. Supralabials uniform or speckled with black. Venter yellow or whitish, uniform or with lateral rows of black spots or short streaks or irregular black speckling, sometimes delimiting a mid-ventral band of grey ofuscation. A specimen from coastal Gabon with a contrasted head and body pattern is illustrated in Pauwels & Vande weghe (2008, Figs 333–334), and a specimen from Kenya with uniform dorsum is illustrated in Spawls et al. (2002: 405). Roux-Estève (1965) provided a detailed description of the two types of patterns of the populations of southern Central African Republic. De Witte (1966) provided scale counts of specimens of Garamba National Park in northern Democratic Republic of the Congo.

Size. Largest specimens for the species, including south of 12°S: NMK 3233/1 – Kassa B, Sennar, Sudan, 1,320 + 460 = 1,780 mm; IRD 2226.N – Baïbokoum, Chad, 1,227 + 508 = 1,735 mm; IRD 2136.N – Baïbokoum, Chad, 1,235 + 515 = 1,750 mm; NMZB 16031 – Ndau School, Western Province, Zambia, 1,260 + 500 = 1,760 mm.

Remarks. ( Broadley, 1977, 1983) first applied the name P. phillipsi to populations in eastern Africa, but Brandstätter (1995) and Hughes (1999) restricted the name to the uniform olive form with an entire cloacal shield in West Africa, and our molecular data support this view. Branch (1998) first used the name P. mossambicus Peters for the southern African populations after Broadley selected a lectotype in Berlin. Hughes & Wade (2004) used the name P. occidentalis for populations from Cameroon to Uganda with a divided cloacal, but our data show little molecular divergence between southern African populations and those from north of the Congo forest block, despite a great variety of colour patterns in both regions. Another available name for this species was P. irregularis Fischer, 1856 , based on a specimen from Peki in former German Togo with a divided cloacal and extensive black dorsal patches on the anterior third of the body, decreasing posteriorly, but our molecular study show that such specimens from Togo are molecularly identical to typical P. phillipsi from West Africa. Data for specimens south of 12°S were published by Broadley (2002). Further molecular studies are needed to investigate if additional species of this complex occur around the Congo forest block (e.g. sequencing specimens with a grey mid-ventral band from Central African Republic and other areas), and to clarify the situation in Nigeria where both P. phillipsi and P. mossambicus occur, possibly in sympatry.

Habitat. Moist savannas and grasslands, especially riparian habitats, swamps, reed beds and cultivated areas from sea level to 1,500 metres.

Distribution. Southeastern Nigeria, Cameroon, southern Chad, Central African Republic, Sudan, South Sudan, Gabon, Republic of Congo, Democratic Republic of Congo, Rwanda, Burundi, Uganda, Kenya, Tanzania, Angola, Zambia, Mozambique, Malawi, northeastern Namibia, northern Botswana, Zimbabwe and the Northern, Mpumalanga and KwaZulu-Natal provinces of South Africa.

Figures

Fig. 6. Psammophis phillipsi “irregularis”. Sequenced specimen IRD 44.T. Vicinity of Kpalimé (Togo).

Fig. 1. Phylogenetic tree (16S, Cyt. B, ND4) of the African species of Psammophis inferred with maximum likelihood from a concatenated dataset. Numbers on branches are bootstrap support values. Black lines separate the major lineages, and red lines clades within lineages 1–3 which include most species of the Psammophis sibilans group.

Fig. 22. Psammophis mossambicus. Sequenced specimen IRD 2238.N. Baïbokoum (Chad).

Fig. 23. Psammophis mossambicus. Sequenced specimen LC.4366. Gamba (Gabon).