Psammophis Boie, 1825

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 : 345-354

Genus Psammophis Boie, 1825

Psammophis angolensis ( Bocage, 1872) DWARF SAND SNAKE

Amphiophis angolensis Bocage 1872:82 . Holotype: MBL 1822 (collector F.A.P. Bayão), destroyed by fire 18 March 1978. Type locality: “Dondo dans l’intérieur d’Angola” [= Dondo ], Kwanza Norte, Angola.

Ablabes Homeyeri Peters (1877a:620) . Holotype: ZMB 9209 (collector A. von Homeyer). Type locality: “Pungo Andongo,” Malanje Province, Angola.

Amphiophis angolensis : Bocage (1895a:113, 1897a:201).

Psammophis angolensis : Boulenger (1896:170), Boettger (1898:104); Boulenger (1915:213); Schmidt (1933:14), Loveridge (1933:252, 1940:68, 1957:280), Laurent (1950a:9, 1954a:59, 1964a:114), FitzSimons (1962:235), Frade (1963:253), Auerbach (1987:171), Brandstätter (1996:40), Broadley (1962a:834, 1977b:26, 1990:148, 2002a:98), Broadley and Cotterill (2004:50), Spwals et al. (2004:403), Kelly et al. (2008:1053), Bates et al. (2014:374), Wallach et al. (2014:575).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is widespread in sub-Saharan Africa, from Ethiopia and Tanzania, through the Democratic Republic of Congo, through Zambia, Malawi, Mozambique, westwards to Angola and southwards through northern Namibia, Botswana, and Zimbabwe to northeastern South Africa (Limpopo and Mpumalanga provinces).

Ocurrences in Angola (Map 307): The species occurs in the entire country with exception of the Namibe Province and the Cabinda enclave. Zaire: “Ambrizette” [-7.23333,

12.86667] ( Bocage 1895a:113, 1897a:201;

Boettger 1898:104; Loveridge 1940:68).

Kwanza Norte: “Dondo” [-9.68333,

14.43333] ( Bocage 1872:82, 1895a:113,

1897a:201; Loveridge 1933:252, 1940:68,

1957:280; FitzSimons 1962:235; Auerbach

1987:171, Broadley 1962a:834, 1977b:26,

1990:148, 2002a:118; Wallach et al. 2014:575).

Malanje: “ Malanje ” [-9.55000, 16.35000]

( Peters 1881:149; Bocage 1895a:113,

1897a:201; Loveridge 1940:68); “PungoAndongo” [-9.66667, 15.58333] ( Peters

1877a:620; Bocage 1895a:113, 1897a:201;

Loveridge 1940:68, 1957:280; Broadley MAP 307 Distribution of Psammophis angolensis in Angola.

1977b:26, 2002a:118). Lunda Norte:

“Dundo” [-7.36667, 20.83333] ( Bocage 1895a:113; Laurent 1950a:9, 1954a:59, 1964a:114; Loveridge 1957:280). Moxico: “environs du lac Calundo” [-11.80000, 20.86667] ( Laurent 1964a:114); “Réserve de chasse de Cameia” [-11.83333, 21.00000] ( Laurent 1964a:114). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] ( Bocage 1895a:113; Loveridge 1940:68). Benguela: “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:113, 1897a:201; Loveridge 1940:68). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:113, 1897a:201; Loveridge 1940:68). Cunene: “Humbe” [-16.68333, 14.90000] ( Bocage 1895a:113, 1897a:201; Schmidt 1933:14; Loveridge 1940:68).

Taxonomic and distributional notes: The type locality, Dondo, has been variously misidentified as “Danda, Loanda Distr.” ( Loveridge 1940, Broadley 1962a) “Donda, Loanda Distr.” ( Auerbach 1987), “Donda i.e. Dundo, Loanda distr.” (FitzSimons 1962a; Broadley 1990) or even “Dondo, Luanda ” (Crawford-Cabral and Mesquitela 1989). Psammophis angolensis is a widespread species with several peculiar autapomorphies. It was originally described in the genus Amphiophis Bocage, 1872 , and Brandstätter (1995) suggested that this should be revived as a monotypic subgenus. However, Kelly et al. (2008) did not follow this suggestion in their phylogenetic anaylsis of Psammophis .

Psammophis ansorgii Boulenger, 1905 LINK-MARKED SAND RACER (Endemic)

Psammophis Ansorgii Boulenger 1905:113 , pl. 4, fig. 4. Holotype: BMNH 1946.1 .8.49 (formerly BMNH

1905.5.29.31) (collector W.J. Ansorge). Type locality: “between Benguella and Bihé,” Angola. Psammophis jallae : Loveridge (1940:62). Psammophis ansorgii : Hellmich (1957b:69), Broadley (1977b:26, 2002a:98), Brandstätter (1996:42), Wallach et al. (2014:576).

Global conservation status (IUCN): Not

Evaluated.

Global distribution: The species is known from Angola.

Ocurrences in Angola (Map 308): The species is endemic to the central highlands of

Angola. Huambo: “Bela-Vista” [-12.36667,

16.20000] ( Hellmich 1957b:69). Undetermined locality: “Between Benguella to Bihé”

( Boulenger 1905:113; Loveridge 1940:62;

Broadley 1977b:26, 2002a:98; Wallach et al.

2014:576).

Taxonomic and distributional notes:

Loveridge (1940) placed Psammophis ansorgii in the synonymy of Psammophis jallae Peracca, 1896 , but Broadley (1977b) considered it a

MAP 308. Distribution of Psammophis ansorgii in full species, as have subsequent authors.

Angola.

Psammophis jallae Peracca, 1896 JALLA’S SAND SNAKE

Psammophis jallae Peracca 1896:2 , 2 figs. Syntypes: MZUT 1633 and 1871 (formely MZUT 1823 a-b)

(collector L. Jalla). Type locality: “Strada da Kazungula a Buluwaio” ( Peracca 1896:2), [= road from

Kazungula, Zambia to Bulawayo], Zimbabwe. Psammophis Rohani Angel (1922:116) . Type: MNHP 20-198 (collector J. F. de Rohan-Chabot). Type locality: “Région de la rivière Lumuna, affluent de Loengoué” [= Lumuna River , tributary of the Luina and

Quando Rivers], Angola. Psammophis rohani : Angel (1923:166), Monard (1937b:128). Psammophis jallae : Loveridge (1940:62), FitzSimons (1962:237), Brandstätter (1996:65), Broadley

(1977b:12, 1990:138, 2002a:93), Branch (1998:90), Bates et al. (2014:376), Wallach et al. (2014:576).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from central and southeastern Angola, western Zambia, northeastern Namibia, Botswana, western and central Zimbabwe and Limpopo Province of South Africa. Also from the southern portion of the former Katanga Province, Democratic Republic of Congo (fide Wallach et al. 2014).

Ocurrences in Angola ( Map 309 ): The species occurs in central and southeastern Angola. Bié: “Bigondo” [-12.06667, 17.41667] ( Loveridge 1940:64); “Benguella to Bihé” ( Loveridge 1940:64). Cuando Cubango: “Lumuna regions, Loengoué affluent” ( Angel 1921:118;

1923:168; Monard 1937b:128; Loveridge

1940:62; Broadley 1977b:12, 2002a:93).

Taxonomic and distributional notes:

Angel (1923) provided three views of the head

(figs. 10–12) of the species he described the year previously. The date of description has been given as 1921, but Angel (1923) explicitly stated that the associated issue of the Bulletin de la Société Zoologique de France had appeared only in March 1922. Psammophis rohani was synonymized with P. jallae Perraca ,

1896 by Loveridge (1940).

Psammophis leopardinus (Bocage, 1887) LEOPARD SAND SNAKE

Psammophis sibilans var. leopardinus Bocage 1887a:206 . Lecototype: MBL 1798 (collector J.A. d’Anchieta),

designated by Broadley (1977b:18), destroyed by fire 18 March 1978. Type locality: “Catumbella”

( Bocage 1887b:206) [= Catumbela] Benguela Province, Angola. Psammophis sibilans var. C leopardina: Bocage (1895a:117). Psammophis sibilans var. E brevirostris : Bocage (1895a:118). Psammophis brevirostris : Boulenger (1915:213); Monard (1937b:133), Themido (1941:10). Psammophis brevirostris brevirostris : Brandstätter (1996:45). Psammophis sibilans sibilans : Loveridge (1957:279). Psammophis sibilans leopardinus : Broadley (1977b:18). Psammophis brevirostris leopardinus : Brandstätter (1996:48), Branch (1998:91). Psammophis leopardinus : Broadley (1990:143, 2002a:95), Kelly et al. (2008:1048), Hughes and Wade

(2002:75), Bates et al. (2014:374), Wallach et al. (2014:577).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from dry savanna and semi-desert regions in southwestern Angola and central and northern Namibia.

Ocurrences in Angola (Map 310): This species occurs chiefly in southwestern Angola. Luanda: “Luanda” [-8.83333, 13.26667] (Broadley 2002:111; Hughes and Wade 2002:77). Huambo: “Bella Vista (Missao di Dondi)” [-12.36667, 16.20000] (Broadley 2002:111; Hughes and Wade 2002:77); “Huambo” [-12.76667, 15.73333] ( Themido 1941:10). Benguela: “Lobito bay” [-12.35000, 13.5500] ( Broadley 1977b:19, 2002a:111; Hughes and Wade 2002:77); “Catumbella” [-12.43333, 13.55000] ( Bocage 1895a:117; Loveridge 1957:279; Broadley 1990:143, 2002a:111; Hughes and Wade 2002:77); “Quissange” [-12.43333, 14.05000] ( Bocage 1895a:117); “Catengue” [-13.03333, 13.73333] (Broadley 2002:111; Hughes and Wade 2002:77). Huíla: “Caconda” [-13.73333, 15.06667] (Broadley 2002:111; Hughes and Wade 2002:77); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937b:133; Broadley 2002:111); “Quillengues” [-14.06667, 15.08333] ( Bocage 1895a:114); “Vila da Ponte” [-14.46667, 16.30000] ( Monard 1937b:133; Broadley 2002:111); “Capelongo” [-14.88333, 15.08333] (Hughes and Wade 2002:77). Namibe: “Capangombe” [-15.10000, 13.15000] ( Bocage 1887b:206; Broadley 2002:111); “Mossamedes” [-15.20000, 12.15000] ( Bocage 1887b:206,

1895a:114; Loveridge 1957:279); “Iona”

[-16.28106, 12.22291] (Broadley 2002:111;

Hughes and Wade 2002:77). Cunene:

“Oncocua, 37 km NE on way to Otchinzau”

[-16.65606, 13.42370] (Broadley 2002:111;

Hughes and Wade 2002:77). Cuando Cubango: “Kayundu” [-15.70000, 17.45000]

( Monard 1937b:133). Undetermined Locality: “Cuanza” (Broadley 2002:111).

Taxonomic and distributional notes: In addition to his Catumbella specimen, Bocage

(1887b) based his description on a second specimen from, “l’intérieur de Mossamedes”

collected by Capello and Ivens. Brandstätter

(1996) recognized P. sibilans as occurring no further south than the northern part of Tanzania and proposed the reinstatement of P. brevi-

MAP 310. Distribution of Psammophis leopardinus in Angola.

rostris Peters, 1881 as a full species, rather than a subspecies of P. sibilans , with leopardinus treated as a subspecies of it. Currently P. leopardinus is recognized as a valid species ( Broadley 1977b, 2002a; Kelly et al. 2008; Branch 1998; Wallach et al. 2014). The record from “Calombe” has been assigned to P. leopardinus although the habitat does not correspond to the typical dry savanna and semi-desert regions where the species is commonly found.

Psammophis mossambicus Peters, 1882 OLIVE WHIP SNAKE

Psammophis sibilans var. mossambica Peters 1882c:122 . Lectotype: ZMB 2468 A, designated by Broadley (2002:96). Type locality: “auf der Insel Mossambique und auf dem gegenüberliegenden Festlande, auf der Halbinsel Cabaçeira und Mesuril […] auf den Querimba-Inseln […] und in Boror” (collector W.C.H. Peters), restricted to “der Insel Mossambique” [= Mozambique Island] by lectotype designation.

Psammophis sibilans : Bocage (1866a:48, 1895a:114, 1896a:113), Peters (1877a:615), Ferreira (1904:116), Boulenger (1905:113, 1915:213), Schmidt (1933:14), Monard (1937b:131), Loveridge (1936a:38).

Psammophis elegans : Bocage (1867b:226).

Psammophis sibilans sibilans : Mertens (1938a:441), Bogert (1940:70), Loveridge (1940:30), Laurent (1964a:113), Hellmich (1957b:70), Manaças (1973:196).

Psammophis phillipsii : Broadley (1977b:24), Branch and McCartney (1992:2), Brandstätter (1996:55), Hughes (1999:64), Kelly et al. (2008).

Psammophis mossambicus : Branch (1998:92), Broadley (2002:96), Broadley et al. (2003:167), Broadley and Cotterill (2004:50), Kelly et al. (2008:1048), Wallach et al. (2014:577), Ceríaco et al. (2016b:87), Conradie et al. (2016:22) .

Psammophis cf. mossambicus : Branch and Conradie (2015:200).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from South Sudan to South Africa, west through Swaziland in the east, in the west to northern Namibia (see Notes).

Ocurrences in Angola (Map 311): The species occurs throughout the country, including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] ( Peters 1877a:615; Loveridge 1940:30); “ Cabinda ” [-5.55000, 12.18333] ( Ferreira 1904:116; Loveridge 1940:30). Zaire: “San Antonio” [-6.13333, 12.36667] ( Loveridge 1940:30). Bengo: “Ambriz” [-7.844312, 13.106493]

( Loveridge 1940:30; Broadley 2002:113).

Luanda: “Loanda” [-8.83333, 13.26667]

( Bocage 1866a:48, 1895a:114; Loveridge

1940:30); “Loanda island” [-8.83333,

13.26667] ( Loveridge 1940:30; Broadley

2002:113); “Quicamba Park” [-9.18333,

13.38333] (Broadley 2002:113). Kwanza

Norte: “Cazengo” [-9.33333, 14.76667] (Ferreria 1904:116; Loveridge 1940:30). Malanje:

“ Duque de Bragança ” [-9.10000, 15.95000]

( Boulenger 1905:113; Loveridge 1940:30;

Broadley 2002:113); “Pungo-Andongo”

[-9.66667, 15.58333] ( Boulenger 1905:113;

Loveridge 1940:30; Broadley 2002:113);

“ Cangandala National Park ” [-9.84606,

16.72233] (Ceríaco et al. 2016b:87); “Mulondo, Luanda Res., Malanje ” ( Broadley

2002:113). Lunda Sul: “Alto Chicapa” MAP 311. Distribution of Psammophis mossambicus in Angola.

[-10.88333, 19.23333] ( Laurent 1964a:113).

Moxico: “ Lago Dilolo” [-11.50000, 22.01667] ( Manaças 1973:196); “evirons du lac Calundo” [-11.80000, 20.86667] ( Laurent 1964a:113); “Calombe, Luso” [-11.83333, 19.93333] ( Manaças 1973:196); “réserve de chasse de Cameia” [-11.83333, 21.00000] ( Laurent 1964a:113); “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:113). Kwanza Sul: “Libolo/Luati” [-9.98333, 14.90000] ( Hellmich 1957b:70); “Chingo” [-11.20000, 13.85000] ( Ferreira 1904:116). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1933:14; Loveridge 1940:30; Broadley 2002:113); “Cubango basin (16b)” [-12.54222, 16.67694] ( Conradie et al. 2016:8,9, 22). Huambo: “Galanga” [-12.06667, 15.15000] ( Bocage 1895a:114; Loveridge 1940:30); “Bela-Vista (Sanguengue)” [-12.36667, 16.20000] ( Hellmich 1957b:70); “Chipipa 13 km N, Hamubo” [-12.55000, 15.73333] (Broadley 2002:113); “Cuma” [-12.85889, 15.06722] ( Loveridge 1936a:38, 1940:30; Broadley 2002:113). Benguela: “Lobito bay” [-12.33333, 13.58333] ( Bogert 1940:79); “Catumbella” [-12.43333, 13.55000] ( Loveridge 1940:30); “Quissange” [-12.43333, 14.05000] ( Loveridge 1940:30); “Benguella” [-12.58333, 13.41667] ( Bocage 1867b:226); “Ebanga” [-12.73333, 14.73333] ( Monard 1937b:131, Loveridge 1940:30); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:441; Loveridge 1940:30; Hellmich 1957b:70; Broadley 2002:113); “Katange” [-13.03333, 13.73333] ( Loveridge 1940:30); “Hanha” [-13.30000, 14.20000] ( Bocage 1896a:113; Loveridge 1940:30; Broadley 2002:113); “Capaia to Cubal” (Broadley 2002:113). Huilla: “ Rio Cuce” [-13.51667, 15.20000] ( Bocage 1895a:114; Loveridge 1940:30); “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:114; Loveridge 1940:30); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937b:131; Loveridge 1940:30); “Quillengues” [-14.06667, 15.08333] ( Loveridge 1940:30); “Kuvangu” = “Vila-da-Ponte” [-14.46667, 16.30000] ( Monard 1937b:131; Loveridge 1940:30); “Capelongo” [-14.91667, 15.08333] ( Bogert 1940:70; Broadley 2002:113); “Huilla” [-15.05000, 13.55000] ( Bocage 1895a:114; Loveridge 1940:30); “Tchiuinguira, Huíla ” [-15.16667, 13.30000] (Broadley 2002:113); “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] ( Laurent 1964a:113), “Negola 6 km S, Huíla ” (Broadley 2002:113). Namibe: “Mossamedes” [-15.20000, 12.15000] ( Loveridge 1940:30; Broadley 2002:113); “ Rio Coroca” [-15.78333, 12.06667] ( Bocage 1895a:114); “Port Alexander” [-15.80000, 11.83333] ( Loveridge 1940:30). Cunene: “fl. Mbalé ” [-15.16667, 16.75000] ( Monard 1937b:131); “Mupa” [-16.18333, 15.75000] ( Monard 1937b:131;

Loveridge 1940:30); “Mupanda” [-17.13333, 15.76667] ( Monard 1937b:131; Loveridge 1940:30); “Tala Kilau, Mossamedes” (Broadley 2002:113). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] ( Monard 1937b:131; Loveridge 1940:30); “Kayundo” [-15.70000, 17.45000] ( Loveridge 1940:30); “vincity of Cuito Cuanavale - approximatley 75 km W of Mavinga” [-15.78333, 19.70000] (Branch and McCartney 1992:2); “Cubango basin (48)” [-14.68136, 17.44530] ( Conradie et al. 2016:9, 12, 22). Undetermined Locality: “without precise location” (Laurent 1954:59, 1964a:113); “Cascalada 6 km NW” (Broadley 2002:113); “Marimo 10 km W of Machado” (Broadley 2002:113).

Taxonomic and distributional notes: The taxonomy of the Psammophis sibilans complex, to which this species belongs, remains confused ( Kelly et al. 2008; Wallach et al. 2014) and we have here treated Angolan specimens previously refered to P. sibilans ( Linnaeus, 1758) (except those attributed to P. zambiensis ) and P. phillipsii (Hallowell, “1844” 1845) as P. mossambicus (e.g., Loveridge 1953; Broadley 1977b; Broadley et al. 2003). Brandstätter (1996) and Hughes (1999) considered P. phillipsii to be restricted to West Africa, and Branch (1998) was the first to use the name P. mossambicus Peters, 1882 to refer to the former southern and East African populations of P. phillipsii . Broadley (2002a) likewise recognized that the Angolan material initially assigned to P. sibilans and P. phillipsii should be attributed to P. mossambicus .

Psammophis namibensis Broadley, 1975 NAMIB SAND SNAKE

Psammophis leightoni namibensis Broadley 1975b:9 , pl. 1. Holotype: TM 37093 (collector W.D. Haacke). Type locality: “Harus in the Uri-Hauchab Mountains , Diamond Area No. 2, South West Africa ” [= Karas Region], Namibia.

Psammophis leightoni namibensis : Broadley (1977b:11, 1990:136), Brandstätter (1996:68), Branch (1998:90).

Psammophis namibensis : Broadley (2002:91), Bates et al. (2014:378), Wallach et al. (2014:577).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from arid regions, including desert and succulent scrubland, occurring along the southwest Africa, from southwestern Angola extending through the Namib Desert into western Namibia, to

Namaqualand, in the Northern Cape Province,

South Africa.

Ocurrences in Angola (Map 312): The species occurs in the southwestern Angola in the Namib Desert. Namibe: “Mossamedes”

[-15.20000, 12.15000] ( Broadley 1975b:11;

2002a:107); “Pico Acezevedo” [-15.55000,

12.51667] ( Broadley 2002:107); “ Rio Coroca”

[-15.78333, 12.06667] ( Broadley 1975b:11;

2002a:107); “ Foz do Cunene ” [-17.28333,

11.80000] ( Broadley 1975b:11; 2002a:107);

“ Cunene Forde, 15 km NE, Iona Res.”

(Broadley 2002:107).

Taxonomic and distributional notes:

Broadley (1975b) initially described this form as a new subspecies of Psammophis leightoni

Boulenger, 1902, but later considered it a full

MAP 312. Distribution of Psammophis namibensis in species (Broadley 2002). Kelly et al. (2008), Angola.

however, noted that Psammophis namibensis and Psammophis leightoni might represent a single species ( Bates et al. 2014; Wallach et al. 2014).

Psammophis notostictus Peters, 1867 KAROO SAND SNAKE

Psammophis moniliger var. notostictus Peters 1867a:237 . Syntypes: ZMB 5756 a-b (collector C.H. Hahn). Type locality: “Otjimbingue in Südwestafrica” [= Otjimbingwe, SE Karibib District , Erongo Region], Namibia.

Psammophis sibilans var. stenocephalus Bocage (1887a:205) . Holotype: MBL specimen, number unknown (collectors H.C. Capello and R. Ivens). Type locality: “intérieur de Mossamedes” ( Bocage 1887b:205), corrected to “Rio Coroca” by Bocage (1895a:116), Namibe Province, Angola.

Psammophis sibilans var. B ( var. stenocephala ): Bocage (1895a:116).

Psammophis notostictus : Boulenger (1895a:538, 1896:156, 1915:213), Monard (1937b:128), FitzSimons (1962:223), Frade (1963:253), Broadley (1975b:13, 1977b:10, 2002a:87), Brandstätter (1996:77), Branch (1998:90), Bates et al. (2014:377); Wallach et al. (2014:577); Ceríaco et al. (2014b:41).

Psammophis sibilans notostictus : Loveridge (1940:44).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from Karroid bushveld, Namib Desert and fynbos areas from southwestern Angola, south through Namibia to the Cape Peninsula and eastwards through the Karoo to the Albany District, Eastern Cape Province.

Ocurrences in Angola (Map 313): The species occurs in the southwestern Angola in the Namib Desert. Namibe: “Espinheira”

[-16.78717, 12.35764] (Ceríaco 2016b: 41);

“ Pico Azevedo” [-15.53400, 12.49197] ( Ceríaco 2016b: 41); “São Nicolao River ”

[-14.26667, 12.36667] ( Boulenger 1896:156;

Loveridge 1940:44; Broadley 1975a:14,

2002a:103); “intérieur de Mossamedes (= Rio

Coroca)” [-15.78333, 12.06667] ( Bocage

1887a:205, 1895a:116; Monard 1937b:128;

Loveridge 1940:44; Broadley 1975b:14,

1977b:10, 2002a:87,103).

Taxonomic and distributional notes:

Psammophis sibilans var. stenocephalus

Bocage 1887, was was synonymized with

Psammophis notostictus Peters, 1867 by

Boulenger (1895a). Psammophis notostictus MAP 313. Distribution of Psammophis notostictus in Angola.

was described as a variety of Psammophis moniliger (Daudin, 1803) by Peters (1867a) but was also treated as a variety of P. sibilans ( Linnaeus, 1758) by several authors (see Broadley 1975b). Bocage (1895a) referred to P. sibilans var. B, but in the same account equated this with his P. s. var. stenocephalus Bocage, 1887 . Boulenger (1896) treated P. notostictus as a full species, and was followed by several authors until Loveridge (1940) again considered it as a subspecies of P. sibilans . Mertens (1955) restored notostictus to its current specific rank.

Psammophis subtaeniatus Peters, 1882 STRIPE-BELLIED SAND SNAKE

Psammophis sibilans var. subtaeniata Peters 1882c:121 . Lectoype: ZMB 9992A designated by Broadley (1977b:13). Type locality: “sowohl im Innern des Landes bei Tette, als näher der Küste, wie bei Boror,” restricted to “ Tete, Moçambique ” by Broadley (1966b).

Psammophis bocagii Boulenger (1895a:538) . Syntype: BMNH 67.7.23.22 (collector Donaldson Smith). Type locality: “ Angola ” later given as “Benguella,” Angola ( Boulenger 1896:161).

Psammophis sibilans var. A ( var. subtaeniata ): Bocage (1895a:116).

Psammophis bocagii : Boulenger (1896:161, 1915:213), Monard (1937b:131), Bogert (1940:82).

Psammophis Bocagii : Bocage (1897a:201).

Psammophis subtaeniatus subtaeniatus : Loveridge (1940:55), Broadley (1966b:6, 1977b:13), Brandstätter (1996:99).

Psammophis subtaeniatus : Branch (1998:91), Broadley (1990:342, 2002b:93), Bates et al. (2014:379), Wallach et al. (2014:580), Conradie et al. (2016:22) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from southern Angola and northern Namibia, east through Botswana to southern Zambia, Zimbabwe, parts of western Mozambique, the northeastern provinces of South Africa and eastern Swaziland.

Ocurrences in Angola (Map 314): The species occurs in southern Angola. Benguela:

“Catumbella” [-12.43333, 13.55000] ( Bocage

1895a:115, 1897a:201; Loveridge 1940:55;

Broadley 2002:108); “Benguella” [-12.58333,

13.41667] ( Boulenger 1896:161; Loveridge

1940:55; Broadley 1966b:8, 1977b:13,

2002a:108). Huíla: “Molundo” [-15.01667,

13.20000] ( Monard 1937b:131; Loveridge

1940:55). Namibe: “Biballa” [-14.76667,

13.36667] ( Bocage 1895a:115, 1897a:201;

Loveridge 1940:55); “Chao de Chella”

[-14.88944, 13.27417] (Broadley 2002:108);

“Maconjo” [-15.01667, 13.20000] ( Bocage

1895a:115, 1897a:201; Loveridge 1940:55;

Broadley 2002:108). Cunene: “Mupa”

[-16.18333, 15.75000] ( Bocage 1897a:201;

Loveridge 1940:55; Broadley 2002:108); MAP 314. Distribution of Psammophis subtaeniatus in Angola.

“Cahama 5 km SE” [-16.28333, 14.30000]

(Broadley 2002:108); “Humbe” [-16.68333, 14.90000] ( Bocage 1895a:115, 1897a:201; Loveridge 1940:55; Broadley 2002:108); “Forte Roçadas” [-16.71667, 15.01667] ( Monard 1937b:131; Loveridge 1940:55; Broadley 2002:108); “Dongwenna, Mossamedes” [-17.01667, 14.71667] (Broadley 2002:108). “Mupanda” [-17.13333, 15.76667] ( Monard 1937b:131; Loveridge 1940:55; Broadley 2002:108); “ Cunene ” ( Bocage 1895a:114; Loveridge 1940:55; Broadley 2002:108). Cuando Cubango: “Cuito drainage” [-17.57333, 23.26000] ( Conradie et al. 2016:9-10, 22). Undetermined Locality: “Cuanza” ( Bocage 1895a:115); “Rio Bengo ” ( Bocage 1895a:115, 1897a:201; Loveridge 1940:55; Broadley 2002:108); “Pen Pen” (Broadley 2002:108). Taxonomic and distributional notes: Psammophis subtaeniatus has been known by a variety of names, Boulenger (1895a) restricted the name subtaeniatus to the eastern form and erected a new name, Psammophis bocagii (Boulenger 1895) , for the western race based on specimens from Angola ( Broadley 1977b). The description appears as part of a key and no precise locality or number of specimens is given, however, the fact that a range of scale counts is provided demonstrates that there was originally a syntype series. Boulenger (1896) indicated that the single BMNH specimen had come from Bocage in Lisbon, so it is likely that at least one syntype remained in the MBL collection, perhaps not marked as such. Bocage (1895a) referred to P. sibilans var. A, but in the same account equated this with P. s. var. subtaeniatus Peters, 1882 . Loveridge (1940) used P. subtaeniatus subtaeniatus for the western form and placed P. bocagii in its synonymy.

Psammophis trigrammus Günther, 1865 WESTERN SAND SNAKE

Psammophis trigrammus Günther 1865b:95 , pl. 2, fig. e. Holotype: BMNH 1946.1 .8.12 (formerly BMNH 64.7.11.4) (collector J. Monteiro). Type locality: “on the banks of river S. Nicolao (Little Fish Bay, West Africa)” [= Rio São Nicolau ], Namibe Province, Angola.

Psammophis trigrammus : Bocage (1887b:206), Loveridge (1940:23), FitzSimons (1962:221), Broadley (1977b:9, 1990:133, 2002a:87), Brandstätter (1996:104), Branch (1998:89), Bates et al. (2014:379), Wallach et al. (2014:581).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known from southern Angola and western Namibia, reaching its southern limit in the extreme north of the Richtersveld, South Africa.

Ocurrences in Angola (Map 315): The species is restricted to Namibe Province.

Namibe: “ Catara River ” [-13.55000,

12.55000] (Broadley 2002:103); “on the banks of river S. Nicolao ( Little Fish bay , West Afrika)” [-14.26667, 12.36667] ( Günther

1865b:95); “ Rio de São Nicolau, Mossamedes

(Rio San Nicolau)” [-14.26667, 12.36667]

( Bocage 1887b:206; Loveridge 1940:23;

FitzSimons 1962:221; Broadley 1977b:9,

1990:133, 2002ba:86, 103; Wallach et al.

2014:581); “ Rio Coroca ” [-15.78333,

12.06667] (Broadley 2002:103); “Iona

Reserve, 7 km to Oncócua” [-16.85831,

12.61275] (Broadley 2002:103).

Taxonomic and distributional notes:

Bocage (1887b) corrected the type locality to

MAP 315. Distribution of Psammophis trigrammus in “Mossamedes” [= Rio São Nicolau, Namibe]. Angola.

Psammophis zambiensis Hughes and Wade, 2002 ZAMBIAN WHIP SNAKE

Psammophis zambiensis Hughes and Wade 2002:75 . Holotype: BMNH 1959.1 .1.81 (collector H.J. Bredo). Type locality: “supposedly from “Abercorn” (= Mbala) area of Zambia … but likely to be from Mweru-Wantipa”.

Psammophis sibilans sibilans : Laurent (1950a:9, 1954a:59), Thys van den Audenaerde (1966:34).

Philothamnus irregularis irregularis : Manaças (1973:191).

Psammophis zambiensis : Broadley and Cotterill (2004:49), Wallach et al. (2014:582).

Global conservation status (IUCN): Not Evaluated.

Global distribution: The species is known with certainty from the southern Democratic Republic of Congo and northern Zambia. Literature records of Psammophis sibilans that may be referable to P. zambiensis exist from the Caprivi Strip of Namibia and adjacent northern Botswana, northern Zimbabwe, Malawi, Zambezia Province of Mozambique and northestern Angola.

Ocurrences in Angola (Map 316): None confirmed but Psammophis zambiensis is expected to occur in northeastern regions of the country. The following records are P. ‘ sibilans ’ records that may be referable to P. zambiensis . Moxico:

“Calombe” [-11.83333, 19.93333] ( Manaças

1973:191). Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1950a:9, 1954a:59,

1964a:113; Thys van den Audenaerde

1966:34); “Barrage de la Luachimo”

[-7.38333, 20.85000] (Thys van den Audenaer-

de 1966:34); “Dundo, R. Mussungue, aff.

Luachimo” [-7.41667, 20.83333] ( Thys van den Audenaerde 1966:34); “Muita (Luembe

E)” [-7.80000, 21.45000] ( Laurent 1954a:59).

Taxonomic and distributional notes:

Hughes and Wade (2002) found significant differences between specimens of “ leopardinus ”

from Angola and from Zambia and described

P. zambiensis to accommodate the latter. Some eastern records of P. leopardinus could corre-

MAP 316 Distribution of Psammophis zambiensis in spond to this new species. Hughes and Wade Angola.

(2002) presented information from Desmond

Vesey-FitzGerald via Donald G. Broadley that substantiated the likely origin of the holotype from Mweru-Wantipa in the Mporokoso District [ Zambia]. Hughes and Wade (2002) plotted localities of Psammophis ‘ sibilans ’ from the literature, which could include material assignable to P. zambiensis . Two such Angolan records correspond to literature references from Laurent (1950, 1954a, 1964a) and Thys van den Audenaerde (1966). Wallach et al. (2014) included only the former province of Katanga ( Democratic Republic of Congo) and northern Zambia in the distribution of the species, corresponding to the verified records given in the description.

Figures

MAP 309. Distribution of Psammophis jallae in Angola.