BAUER, DEBOER, & TAYLOR: ATLAS OF THE REPTILES OF LIBYA 
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2007. 387: SMF 37598. 389: MZUTR2031; Calabresi 1923; Loveridge and Williams 1957; Zavat- 
tari 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: Schle¬ 
ich 1987. Jabal al Akhdar : 410: Resetar 1981. 419: FMNH 214920; Resetar 1981; Frynta et al. 
2000; Pieh and Perala 2002. 423: USNM 140247; Pieh and Perala 2002. 434: USNM 146810. 438: 
FMNH 214919; Resetar 1981; Pieh and Perala 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 Perala 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 
Perala 2002. 475: MTKD 184073. Butnan : 490: SMF 39035; Zavattari 1922, 1929, 1930, 1934; 
Calabresi 1923; Loveridge and Williams 1957; Pieh and Perala 2002. 495: Loveridge and Williams 
1957. Cyrenaica: “Marmarica”: Zavattari 1937. Eastern Libya: MTKD 28774, 31879, 34852, 
34853; Pieh and Perala 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 var¬ 
ious times. Van der Kuyl et al. (2002, 2005) recognized only four subspecies, but their taxonomy 
was criticized by Perala (2004). Fritz and Havas (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 Perala, 2004, and T. g. soussensis Pieh, 2001 occupy portions of western North Africa. Testu¬ 
do g. cyrenaica Pieh and Perala, 2002 occurs in Cyrenaica and T. g. nabeulensis (Highfiled, 1990) 
is in Tripolitania and Tunisia (Fritz and Havas 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). Gracia 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) 
