Naja haje ( Linnaeus, 1758:225 )

Bauer, Aaron M., DeBoer, Jonathan C. & Taylor, Dylan J., 2017, Atlas of the Reptiles of Libya, Proceedings of the California Academy of Sciences 64 (8), pp. 155-318 : 258-264

Naja haje ( Linnaeus, 1758:225)

1758 Coluber Haje Linnaeus , Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae [Stockholm], Sweden. (4) + 823 + (I) pp.

SYNTYPES.— NRM and Hasselquist collection (see Comments), lost fide Andersson (1899) and Kramer and Schnurrenberger (1963), “AEegypto inferiore” [= Lower Egypt].

Naja haje, Kramer and Schnurrenberger 1963:536 .

Naja haje, Le Berre 1989:256 .

Naja haje, Schleich, Kästle, and Kabisch 1996:526 .

Naja haje, Sindaco, Venchi and Grieco 2013:176 .

DISTRIBUTION.— Across northern Africa exclusive of the Sahara proper, from Senegal to Western Sahara and Morocco to the Nile Valley and Delta ( Baha El Din 2006a). In the Sahel crossing eastwards through northern Nigeria, southern Chad and the Central African Republic into Sudan and the Horn of Africa and as far south as Tanzania in East Africa. Libyan records are chiefly in coastal or near-coastal regions across the country.

Libyan Records (Map 62): TRIPOLITANIA: TRIPOLI: 45: Werner 1909; Scortecci 1939. MURQUB: 60: Werner 1909; Ghigi 1913; Zavattari 1934; Scortecci 1939. 68: SK 448; Kramer and Schnurrenberger 1963. JABAL AL GHARBI: 147: FMNH 83058. “ Tripolitania settentrionale ” Zavattari 1937. SIRTE: 174: MCSN 2688. CYRENAICA: BENGHAZI: 354: Umani 1922. 357: von Martens 1883; Werner 1909; Umani 1922; Zavattari 1922, 1929, 1930, 1934; Calabresi 1923; Ribolla 1923; Scortecci 1939. 376: Zavattari 1922, 1929, 1930, 1934; Calabresi 1923; Scortecci 1939. MARJ: 385: Zavattari 1929, 1937. JABAL AL AKHDAR: 411: FMNH 214914; Resetar 1981. 414: Resetar 1981. 417: Zavattari 1922, 1929, 1930, 1934; Calabresi 1923; Scortecci 1939. 421: Ghigi 1920; Calabresi 1923; Zavattari 1929, 1930. 427: Scortecci 1939. 457: KNP 1981/272, 354, 509; ZSM 145/1983; Resetar 1981; Schleich 1987. 457aj: Schleich 1987. 457an: Schleich 1987. 457ao: Schleich 1987. 457at: Schleich 1987. 457au: Schleich 1987. 457bp: Schleich 1987. 457br: Schleich 1987. 457bs: 179: KNP 1919/13; Schleich 1987. 457bu: Schleich 1987. 457ca: Schleich

1987. 457cc: Schleich 1987. 457cv: Schleich

1987. 163: KNP 1981/311; Schleich 1987.

457i: Schleich 1987. 457r: Schleich 1987.

457u: Schleich 1987. 457w: Schleich 1987.

DARNAH: 469: Kramer and Schnurrenberger

1958, 1963. AL WAHAT: 536: Kramer and

Schnurrenberger 1963. “ Cyrenaica ”: IZUM

20178, IZUM 20179; ZMB 10507; CondorelliFrancaviglia 1896; Kramer and Schnurrenberg-

er 1963. LIBYA: “ Libysche Wüste ”: ZMB

64824.

COMMENTS.— Wüster et al. (2008) have clarified and confirmed the position of the genus Naja within Elapidae . Trape et al. (2009)

revised the N. haje complex. Wallach et al.

(2009) placed the species in the subgenus

Uraeus Wagler, 1830 which Wallach et al.

(2014) regarded as a valid genus. Linnaeus MAP 62. Distribution of Naja haje in Libya.

(1758) noted only Hasselquist (1757) as an earlier source for Coluber Haje but as noted by Andersson (1899), he provided scale counts which differ from those in Hasselquist, suggesting that his concept was based on two different specimens. The second set of ventral and subcaudal counts are identical to those of a specimen from the Adolphi Friderici collection published in volume two of the Museum Adolphi Friderici ( Linnaeus 1764). Although not published until after the Systema Naturae, this work had been completed a decade earlier and so Linnaeus used some of the material in 1758. Andersson (1899) noted that neither the specimen from the Museum Adolphi Friderici nor Hasselquist’s specimen appear to be extant. [Note that Schleich (1987) listed specimens KNP 1981/272, 311, 354, and 509 as both Coluber algirus and Naja haje . We have assumed that the associated localities are correct for both species and that specimen numbers were inadvertently duplicated].

UCN Threat Status: Not assessed, but assumed to be Least Concern. However, regardless of the status of the species as a whole, this taxon is certainly threatened in portions of its North African range. Family Viperidae

Cerastes cerastes ( Linnaeus, 1758:217) (FIG. 34)

1758 Coluber Cerastes Linnaeus , Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae [Stockholm], Sweden. (4) + 823 + (I) pp.

HOLOTYPE.— NRM Lin-98, “Oriente” [= Rashid, Nile Delta, Egypt] fide Hasselquist 1762:70-72, 315- 322], restricted to “ Egypt ” by Flower (1933:830) and to “southern Judaea” by Schmidt (1939:88).

Cerastes cerastes, Kramer and Schnurrenberger 1963:539 .

Cerastes cerastes, Le Berre 1989:294 .

Cerastes cerastes, Schleich, Kästle, and Kabisch 1996:534 .

Cerastes cerastes, Trape and Mané 2006:208 .

Cerastes cerastes, Sindaco, Venchi, and Grieco 2013:192 .

DISTRIBUTION.— Mauritania through southern Israel and south to Sudan, although absent from Morocco north of the Atlas Mountains and coastal Algeria (nominate form), also in southwestern Arabia (C. c. hoofieni). In Libya it occurs countrywide.

Libyan Records (Map 63): TRIPOLITA-

NIA: NUQAT AL KHAMS: 1: Frynta et al. 2000.

JAFARA: 34: Werner 1909; Ghigi 1913; Zavattari 1934. TRIPOLI: 39: Kramer and Schnurrenberger 1959. 45: MNHN 1893.614; NMP

35389. 48: Sayers 1964. 51: Schnurrenberger

1959. MURQUB 60: ZMB 18090; Peters 1880, FIGURE 34. Cerastes cerastes from Ghodwa, Murzuq, 1881; Werner 1909; Ghigi 1913; Zavattari Fezzan, Libya. Photo © Adel Ibrahim.

1934. MISRATAH: 72: SMF 89435–40. 78:

MCSN 2971. 88: MCSN 2942. NALUT: 100:

Frynta et al. 2000. 105: AIC no number provid-

ed; MNHN 2004.0078; Ibrahim and Ineich

2005. 108: MCSN 2957 ; Kramer and Schnurrenberger 1963. 109 : BMNH 1960.1.6. 9;

Kramer and Schnurrenberger 1963. 115: Brito et al. 2008. 121: BMNH 1960.1.6. 10,

1960.1.6. 40; MZUF 39090 ; MZUF 39091 ;

Kramer and Schnurrenberger 1963. 127: Zavattari 1937. JABAL AL GHARBI: 131: SK 1751;

Kramer and Schnurrenberger 1963. 142:

BMNH 1960.1.6. 8; Kramer and Schnurrenberger 1963. 154: ZSM 10/1979. 158 : MCSN

2926. SIRTE: 166: MCSN 2914; Kramer and

Schnurrenberger 1963. 197: MCSN 4197.

“ Tripolitania settentrionale ”: Zavattari 1937.

“ Wadi Guerias ”: BMNH 1901.10.28. 10. FEZ- MAP 63. Distribution of Cerstes cerastes in Libya. ZAN: WADI AL SHATII: 210: Essghaier et al .

2015. JUFRA: 229: SK 3567; Kramer and Schnurrenberger 1963. GHAT: 233: Kramer and Schnurrenberger 1959. 235: Frynta et al. 2000. 264: MSNM [no specific number given]; Kramer and Schnurrenberger 1963. WADI AL HAYAA: 265: ZFMK.63668; Wagner and Wilms 2010. 271: ZMB 37943. SABHA: 281: ZCT 2005.46; Ibrahim 2008a. 284: Kramer and Schnurrenberger 1963; Schnurrenberger 1963. 286: Essghaier et al. 2015. 290: SK 1311; Schnurrenberger 1962; Kramer and Schnurrenberger 1963. MURZUQ: 298: Essghaier et al. 2015. 300: ZCT 2005.03; Ibrahim 2008a. 302: ZCT 2005.17, 2005.19; Ibrahim 2008a. 303: Essghaier et al. 2015; Zavattari 1937; Scortecci 1939. 308: ZCT 2005.18, 2005.33; Ibrahim 2008a. 313: ZCT 2005.34–35, 2006.53; Ibrahim 2008a. 321: MCSN 2964; Scortecci 1935b. “ Fezzan ”: Scortecci 1939. “ Sahara tripolitain ” Angel and Lhote 1938. CYRENAICA: BENGHAZI: 351: BMNH 1974.5300. 357: BMNH 1945.11.9. 1; Ghigi 1920; Umani 1922; Calabresi 1923; Zavattari 1929, 1930, 1934. 366: NHMC 80.3.5.1. 367: Calabresi 1923; Zavattari 1929, 1930, 1934. 377: Kramer and Schnurrenberger 1959. 379: SK 422, 456, 986; Kramer and Schnurrenberger 1958, 1963. 381: Kramer and Schnurrenberger 1959. MARJ: 385: Zavattari 1937. 389: Zavattari 1922, 1929, 1930, 1934; Calabresi 1923. 409: Kramer and Schnurrenberger 1959. JABAL AL AKHDAR: 455: SMF 54174. 457: Schleich 1987. DARNAH: 466: USNM 146794; ZSM 8/1979. 475: MCZ R 46849. 484: Zavattari 1929, 1930, 1934. BUTNAN: 519: SK 98, 149–151, 377; Schnurrenberger 1958a; Kramer and Schnurrenberger 1959, 1963; SMF 55436. AL WAHAT: 529: NMP 34938. 536: SK 324, 556; Kramer and Schnurrenberger 1963. 540: Schnurrenberger 1959. 549: SK 336, 369, 421, 425–26; Kramer and Schnurrenberger 1963. 550: SK 457; SMF 54733; Schnurrenberger 1958a; Kramer and Schnurrenberger 1963. 554: Schnurrenberger 1958a. KUFRAH: 571: Zavattari 1937. 575: Peters 1880, 1881; Werner 1909; Ghigi 1913; Zavattari 1929, 1930, 1934. “ Cyrenaica ”: SMF 54732; Werner 1909. LIBYA: RMNH RENA 13928.

COMMENTS.— Two subspecies of C. cerastes are currently recognized, C. c. cerastes from North Africa and into Sinai and southwestern Israel and C. c. hoofienii Werner and Sivan in Wern- er, Sivan, Kushnir and Motro, 1999 which is found in the southwestern Arabian Peninsula (Wagn- er and Wilms 2010). The original description cites Hasselquist (1750a, 1757) and Bellonius (1553).

Schnurrenberger (1963) and Kramer and Schnurrenberger (1963) noted that the localities provided for the two species of Cerastes by Schnurrenberger (1959) were reversed. Schleich (1987) noted a sight record of what was presumably this species near the northwest corner of watershed area in Kouf (El Kouf) National Park. Zavattari (1934) credited Brezzi (1930) with a Kufrah area record of this species, but we were unable to locate this in Brezzi’s book.

UCN Threat Status: Not assessed, but assumed to be Least Concern.

Cerastes vipera ( Linnaeus, 1758:216) (FIG. 35)

1758 Coluber vipera Linnaeus , Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae [Stockholm], Sweden. (4) + 823 + (I) pp.

HOLOTYPE.— NRM Lin-99, “Aegypto” [= Egypt], [= Rashid , Nile Delta, Egypt fide Hasselquist 1762: 70-72, 315-322].

1842 Cerastes Richiei Gray, Monographic synopsis of the vipers, or the Family Viperidae , pp. 68–71 in The Zoological Miscellany, part 2. Treuttel, Wurtz and Co., London, United Kingdom.

Syntypes: BMNH 1946.1.20. 85, 1946.1.20. 89, 1946.1.20. 94 (formerly BMNH iii.3.2a–c), “ Tripoli ” [ Libya] .

Cerastes vipera, Kramer and Schnurrenberger 1963:541 .

Cerastes vipera, Le Berre 1989:296 .

Cerastes vipera, Trape and Mané 2006:210 .

Cerastes vipera, Schleich, Kästle, and Kabisch 1996:537 .

Cerastes vipera, Sindaco, Venchi, and Grieco 2013:193 . JABAL AL GHARBI: 131: SK 1750, 2458; Kramer and Schnurrenberger 1963. 158: MCSN 2927.

SIRTE: 185: MCSN 2929; Kramer and Schnurrenberger 1963. “ Tripolitania ”: Scortecci

1939. “ Tripolitania settentrionale ” Zavattari

1937. FEZZAN: GHAT: 233: Schnurrenberger

1959. 239: Scortecci 1937a. 245: Scortecci

1937a. 249: Scortecci 1939. MURZUQ: 298:

MCSN 2946, 2958. 305: ZCT 2005.36;

Ibrahim 2008a. 316: ZCT 2006.06; Ibrahim

2008a. “ Ghat-Murzuq-Mesak-Mellet-Uadi

Tilemsin ”: MCSN 2933. CYRENAICA:

BENGHAZI: 358: MZUF 989. 372: Frynta et al.

2000; Kratochvíl et al. 2002. MARJ: 384: CM

91617. BUTNAN: 491: MZUF 992. 521: MSNG

37503; Zavattari 1929, 1930, 1934, 1937. AL

WAHAT: 536: SK 323, 423–24, 287; Kramer and Schnurrenberger 1963. 540: Schnurren- MAP 64. Distribution of Cerestes vipera in Libya. berger 1959. 544: BMNH 1960.1.6. 11–12; CM S7209; Kramer and Schnurrenberger 1963; BMNH 1985.1181. 547: Kramer and Schnurrenberger 1958. 548: SK 301–05; Kramer and Schnurrenberg- er 1963. 549: SK 405–08, 630; Kramer and Schnurrenberger 1963. 550: Schnurrenberger 1958a. 553: MCSN 4198, Zavattari 1937. 554: MCSN 2965; Scortecci 1935b; Kramer and Schnurrenberger 1963. 562: MSNG 31581*; Vinciguerra 1931; Zavattari 1934. 564: MSNG 31581*; Vinciguerra 1931; Zavattari 1934, 1937. KUFRAH: 575: SMF 32710; ZMB 18091; Scortecci 1939. “ Cyrenaica ”: MCSN 2959; MZUF 12153–54; Gestro and Vinciguerra 1931; Scortecci 1939. “ Soluch to Hatiet el Rtém ”: Schnurrenberger 1959.

COMMENTS.— Wagner and Wilms (2010) commented on the conflicting reports of sympatry and syntopy in the literature regarding the spatial relationships between C. cerastes and C. vipera . Our records support that the two occur in sympatry, although C. vipera tends to occupy more xeric environments than C. cerastes , as was concluded by Wagner and Wilms (2010). Schnurrenberger (1963) and Kramer and Schnurrenberger (1963) noted that the localities provided for the two species of Cerastes by Schnurrenberger (1959) were reversed. Scortecci (1937a) listed several localities from near Ghat including Tafilellelt, Uadi Iseien near Tin Caraden, Uadi Inelegghi, and Uadi Iseien near Tin Alcun. Kramer and Schnuerrenberger (1963) treated the first two of these as being in Algerian territory, but considered the other two to be Libyan. However, all four of these localities are actually in the Tassili n’Ajjer of Algeria. Likewise, Scortecci’s (1939) mention of a specimen from the Tassili certainly refers to an Algerian snake.

IUCN Threat Status: Least Concern.

Echis pyramidum (I. Geoffroy Saint-Hilaire, 1827:151, pl. 8, fig. 1) (FIG. 36) 1827 Scythale pyramidum Geoffroy Saint-Hilaire , Description des reptiles qui se trouvent en Égypte, pp.

121–160 in M.J.-C.L. de Savigny, (ed.), Description de l’Égypte, ou Recueil des Observations et des Recherches qui ont été Faites en Égypte Pendant l’Expédition de l’Armée Française. Histoire Naturelle. Tome premier. Partie premier. Imprimerie Impériale, Paris, France.

LECTOTYPE.— MNHN 4031 , designated by Stemmler and Sochurek (1969:90), “Kaire, Egypte” [= Cairo, Egypt].

1972 Echis carinatus leucogaster Roman, Deux sous-especes de la vipere Echis carinatus (Schneider) dans les territoires de Haute-Volta et du Niger: Echis carinatus ocellatus Stemmler — Echis carinatus leucogaster n. ssp. Notes et Documents Voltaïques, 5(4):1–11, 2 maps, 1 pl. [6, pl. figs. 3–8].

HOLOTYPE.— RN 191 , “ Boubon , à 20 km au Nord de Niamey ” [ Niger].

1990 Echis pyramidum lucidus Cherlin , Taxonomic revision of the snake genus Echis ( Viperidae ). II. An analysis of taxonomy and description of new forms [in Russian]. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR, 207:193–223, pls. IV–VI. [206, pl. V, fig. 3].

HOLOTYPE.— USNM 134975, “Египет, Фаюм, Синнурис” [ Egypt, Faiyum, Sinnuris ].

Echis carinatus pyramidum, Kramer and Schnurrenberger 1963:544 .

Echis leucogaster, Le Berre 1989:298 .

Echis pyramidum, Le Berre 1989:300 .

Echis pyramidum lucidus, Schleich, Kästle, and Kabisch 1996:542 .

Echis arenicola leucogaster, Schleich, Kästle, and Kabisch 1996:540 .

Echis pyramidum leucogaster, Sindaco, Venchi, and Grieco 2013:200 . FIGURE 36. Echis pyramidum from between Al Khums and Tripoli, Tripolitania, Libya (32.6694, 14.1242), 2007. Photo courtesy of Natural History Museum of Crete © V. Paravas.

DISTRIBUTION.— From Senegal and Guinea-Bissau across North Africa to Egypt and south to Kenya and the Horn of Africa. In both Egypt and Libya there are scattered localities, none from Kufrah and the southeastern portion of the country, nor from the Egyptian border region.

Libyan Records (Map 65): TRIPOLITANIA: TRIPOLI: 45: BMNH 1960.1.6. 13; Kramer and Schnurrenberger 1963. MURQUB: 65: NHMC 80.3.4.1. 68: Boulenger 1914; Zavattari 1934; Scortecci 1939. “Tripolitania settentrionale”: Zavattari 1937. FEZZAN: SABHA: 286: NMBA-REPT 18127. CYRENAICA: BUTNAN: 490: Ribolla 1923; Zavattari 1934; Scortecci 1939. “ Cyrenaica ”: Condorelli-Francaviglia 1896; Schleich et al. 1996; Sindaco et al. 2013; Werner 1909; Calabresi 1923. “ Marmarica ”: Zavattari 1937.

COMMENTS.— The complicated dating of the natural history portions of Description de l’Égypte has been elucidated by Sherborn

(1897) and Tollitt (1986). Cherlin (1990) and

Cherlin and Borkin (1990) recognized a large number of species and subspecies in the

E. pyramidum complex, including E. arenicola leucogaster Roman, 1972 (type locality:

Boubon, Niger) reaching its easternmost extent in west central Libya and E. pyramidum lucidus

Cherlin, 1990 (type locality: Faiyum, Egypt) ,

which they considered to be distributed across northern Libya and Egypt west of the Nile.

Golay et al. (1993) synonymized E. arenicola with E. pyramidum . Despite obvious color differences ( Hughes 1976), Echis leucogaster was found to be minimally genetically distinct from

E. pyramidum ( Arnold et al. 2009) and has not subsequently been recognized at the specific level by some authors (e.g., Sindaco et al. MAP 65. Distribution of Eryx pyramidum in Libya.

2013), although it has by others (e.g. Wallach et al. 2014). Following Hughes (1976) and Sindaco et al. (2013), the nominotypical form is regarded as extending from Egypt southwards to Kenya, whereas E. p. leucogaster occupies the western portion of the species distribution. These authors did not explicitly indicate into which subspecies Libyan populations should be placed. Alternative interpretations have been reached by McDiarmid et al. (1999), David and Ineich (1999), Pook et al. (2009), and Wallach et al. (2014). Pending further study of the group we consider all Libyan populations as assignable to E. pyramidum pyramidum .

UCN Threat Status: Least Concern.