Testudo graeca nabeulensis Highfield 1990:32

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 : 175-178

Testudo graeca nabeulensis Highfield 1990:32

1990 Furculachelys nabeulensis Highfield, Tortoises of north Africa; taxonomy, nomenclature, phylogeny and evolution with notes of field studies in Tunisia. Journal of Chelonian Herpetology 1(2):1–56. HOLOTYPE.— No specimen number or collection repository stated, “the region of Nabeul, Tunisia.”

Testudo graeca cyrenaica Pieh and Perälä, 2002:8 , figs. 1–5 ( FIG . 9)

2002 Testudo graeca cyrenaica Pieh and Perälä, Variabilität von Testudo graeca Linnaeus, 1758 im östlichen Nordafrika mit Beschreibung eines neuen Taxons von der Cyrenaika (Nordostlibyen). Herpetozoa

15(1/2):3–28. HOLOTYPE.— MTKD 31880 , “ Derna (32°46′N, 22°39′E (= Darnah, Cyrenaika Ostlibyen ).”

Testudo graeca [part], Le Berre. 1989:102.

Testudo graeca [part], Schleich, Kästle, and Kabisch 1996:148.

Testudo graeca, Sindaco and Jeremčenko 2008:84 .

DISTRIBUTION.— Testudo graeca as a whole is widely distributed across North Africa from southern Morocco to Cyrenaica, Libya, though with a gap in the Gulf of Sirte. In the Near and Middle East it ranges from the western Negev Desert of Israel as far as Turkmenistan and eastern Iran. Widespread in Turkey and into the Caucusus, Balkans and Greece. Introduced populations occur in the Canary Islands, France, Spain (including the Balearic Islands), Malta, and Italy, including Sardinia and Sicily.

Schleich et al. (1996) reported Testudo graeca from Egypt, but it does not occur there ( Flower 1933; Buskirk 1996), although it approaches the eastern Sinai just south of the Gaza Strip at Kerem Shalom in Israel ( Buskirk 1996; Baha El Din 2006a; Werner 2016). See Pieh and Perälä (2002) for a discussion of supposed Egyptian specimens.

Testudo g. cyrenaica is endemic to Libya, occurring in Cyrenaica from the region of Jabal al Akhdar [the Green Mountain] to the coast, and from west of Al Marj to as far east as El Adem. Testudo g. nabeulensis occurs in northwestern Tripolitania and in Tunisia.

Libyan Records (Map 5): ( Testudo graeca nabeulensis ): TRIPOLITANIA: JAFARA:

29: MZUT R3682; Andreone et al. 2007.

TRIPOLI: 45: Werner 1909; Ghigi 1913; Zavattari 1934. MURQUB: 58: Peters 1880, 1881;

Anderson 1898; Werner 1909; Ghigi 1913;

Zavattari 1934; Loveridge and Williams 1957.

NALUT: 105: Ibrahim and Ineich 2005. JABAL

AL GHARBI: 140: USNM 146811. 147: MTKD

184072.

( Testudo graeca cyrenaica ): CYRE-

NAICA: BENGHAZI: 346: MTKD 35834; Pieh and Perälä 2002. 357: MZUF 17264; USNM

139091; Ghigi 1913; Calabresi 1923; Zavattari

1929; Loveridge and Williams 1957; Pieh and

Perälä 2002. MARJ: 384: USNM 142155. 385:

MZUT R2024; Calabresi 1923; Zavattari 1929,

1930, 1934, 1937; Loveridge and Williams MAP 5. Distribution of Testudo graeca in Libya. Points in Tripolitania are records of T. g. nabeulensis , those in Cyre- 1957; Pieh and Perälä 2002; Andreone et al. naica are T. g. cyrenaica.

2007. 387: SMF 37598. 389: MZUT R2031; Calabresi 1923; Loveridge and Williams 1957; Zavattari 1929, 1930, 1934; Andreone et al. 2007. 400: Frynta et al. 2000. 457aa: Schleich 1987. 457ab: Schleich 1987. 457ac: Schleich 1987. 457ad: Schleich 1987. 457ae: Schleich 1987. 457b: Schleich 1987. JABAL AL AKHDAR: 410: Resetar 1981. 419: FMNH 214920; Resetar 1981; Frynta et al. 2000; Pieh and Perälä 2002. 423: USNM 140247; Pieh and Perälä 2002. 434: USNM 146810. 438: FMNH 214919; Resetar 1981; Pieh and Perälä 2002. 440: MTKD 103734. 446: ZSM 106193; Resetar 1981. 447: CUP R 141; Frynta et al. 2000. 448: USNM 146811. 457: KNP 1981/184, 192, 216, 290, 316, 317, 463, 466, 488, 497, 510, 1983/[2 specimens, no number given]; ZSM 108– 109/1983, 110/1997, 125/1997, 127/1997; Schleich 1987, 1989; Pieh and Perälä 2002; Schneider and Schneider 2006a, b. 457ar: Schleich 1987. 457at: Schleich 1987. 457au: Schleich 1987. 457ba: Schleich 1987. 457bb: Schleich 1987. 457bc: Schleich 1987. 457bm: Schleich 1987. 457bn: Schleich 1987. 457bo: Schleich 1987. 457bs: Schleich 1987. 457bv: Schleich 1987. 457bx: Schleich 1987. 457c: Schleich 1987. 457ce: Schleich 1987. 457cf: Schleich 1987. 457ch: Schleich 1987. 457ci: Schleich 1987. 457cn: Schleich 1987. 457cr: Schleich 1987. 457cs: Schleich 1987. 457cv: Schleich 1987. 457e: Schleich 1987). DARNAH: 466: MTKD 31880, SMF 36127; Pieh and Perälä 2002. 475: MTKD 184073. BUTNAN: 490: SMF 39035; Zavattari 1922, 1929, 1930, 1934; Calabresi 1923; Loveridge and Williams 1957; Pieh and Perälä 2002. 495: Loveridge and Williams 1957. Cyrenaica: “ Marmarica ”: Zavattari 1937. Eastern Libya: MTKD 28774, 31879, 34852, 34853; Pieh and Perälä 2002.

( Testudo graeca ssp.): LIBYA: MTKD 12488–89, 14554; NHMW 34351; Andreone et al. 2007.

COMMENTS.— Many subspecific entities have been recognized within Testudo graeca at various times. Van der Kuyl et al. (2002, 2005) recognized only four subspecies, but their taxonomy was criticized by Perälä (2004). Fritz and Havaš (2007) listed 17 subspecies. Van Dijk et al. (2014) recognized 10 subspecies, two of which occur in Libya. The nominate form, T. g. marokkensis Pieh and Perälä, 2004, and T. g. soussensis Pieh, 2001 occupy portions of western North Africa. Testudo g. cyrenaica Pieh and Perälä, 2002 occurs in Cyrenaica and T. g. nabeulensis (Highfiled, 1990) is in Tripolitania and Tunisia (Fritz and Havaš 2006, 2007; Van Dijk et al. 2014).

Cyrenaican populations were previously included in the nominotypical form (e.g., Iverson 1992), or assigned to T. g. terrestris (e.g., Wermuth and Mertens 1961). Graciá et al. (2017) found that T. g. nabeulensis is sister to all remaining subspecies. They also identified a new lineage of T. graeca based on a single individual obtained from a Cyrenaican market.

Schleich et al. (1996) suggested that the distribution of T. graeca in Libya was limited to the Mediterranean of Cyrenaica, with a gap westwards to the Tunisian border. However, Iverson (1986, 1992) plotted records from Tripolitania, showing a gap only across the Gulf of Sirte, and Le Berre (1989) implied a more or less continuous range across the Mediterranean coast of Libya. Schnei- der and Schneider (2008) mapped 4 localities for T. g. nabeulensis and 11 for T. g. cyrenaica, but did not specify place names.

Several historical records of Libyan tortoises have caused particular confusion. Peters (1880, 1881) referred to T. campanulata [= T. marginata ] from Bir-Milrha. Werner (1909) suggested that this record might actually refer to T. ibera [= T. graeca ] but Schneider and Schneider (2008) argued that the likely species intended was T. kleinmanni , based on the similarity of plastron pattern between adult T. kleinmanni and juvenile T. marginata . Another record reported by Peters (1880, 1881) from “Uadi Tessiua” was for many years placed in Kufra, deep in the desert and more than 700 km south of the nearest legitimate Libyan record of any Testudo . Schneider and Schneider (2008) convincingly argued that the Kufra error stems from miscitations of Werner (1909) who mentioned “Uadi Tessina [sic] (Rohlfs, Kufra),” referring to Rohlfs’ book, entitled “ Kufra ” (1881) in which Peters’ records appeared. The locality is, in fact, near Gharyan, Tripolitania, and the specimen (now lost) is likely assignable to T. g. nabeulensis .

IUCN THREAT STATUS.— Vulnerable A1cd.

Figures

FIGURE 9. Testudo graeca cyrenaica from the archaeological site of Cyrene, Jabal al Akhdar, Cyrenaica, Libya (32.8239, 21.8519), 1998. Photo courtesy of Natural History Museum of Crete © A. Trichas.