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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 64, No. 8 
on fishing and hunting large aquatic vertebrates (Smith 1968; Sereno et al. 2008). About 4000 years 
ago the modem cycle of desertification began, resulting in the modem landscape and faunal pat¬ 
terns of the country (Kropelin et al. 2008). 
Crocodylus niloticus was once widespread in the Sahara. Historical records exist from areas of 
Tunisia, Algeria, northern Niger and the Tibesti of Chad, but records from the territory of Libya 
(Djebel Akhdar — 32.19618°N, 21.41950°E, In Habiter — 26.49082°N, 12.69354°E, Wadi Math- 
endous — 25.76267°N, 12.16647°E) are Holocene in age (Peters and von den Driesch 2003; de 
Smet 1999; Joleaud 1933; Brito et al. 2011). Kratochvil et al. (2002) provided a photo of a croco¬ 
dile carving on stone in Wadi Mathendous in the Akakus region of Fezzan, suggesting that croco¬ 
diles were present in Libya approximately 8000 ybp, when more mesic conditions prevailed in 
southern Libya. Libya was rich in crocodilian diversity in earlier periods, such as the lower 
Miocene (Llinas Agrasar 2004) and numerous fossil squamates have also been described from 
Libya (Arambourg and Magnier 1961; Hoffstetter 1960, 1961). Tortoise fossils are also known 
from Fezzan (Peters and von den Driesch 2003), where no chelonians occur today. 
History of Libyan Herpetology. — Zoological research in Libya has been reviewed by 
Massa (2009), Chiozzi (2013) and Latella (2013) among many others, and publications specific to 
the herpetology of Libya were reviewed by Bshaena (2011). Early herpetological explorations in 
the area were prevented by the dangerous conditions in North Africa. During the late 1700s and 
early 1800s, the “Barbary States,” including Tripoli engaged extensively in piracy in the southern 
Mediterranean, often capturing European and American ships and enslaving their crews. This 
activity ceased only after the First (1801-1805) and Second (1815-1816) Barbary Wars, and the 
eventual return to Ottoman control over the Tripoli in 1835. As a consequence, in the first half of 
the 19 th century, Libyan species of reptiles were mentioned only in isolated descriptions of species 
or mentions of specimens from “Tripoli” (Daudin 1802; Gray 1838, 1842, 1845, 1849). 
Late in the century the first focused herpetological research resulted from the study of collec¬ 
tions made by a series of expeditions to the region. Peters (1880, 1881) described material in the 
Zoological Museum of Berlin (now Museum fur Naturkunde) collected on the expedition of Ger¬ 
hard Rohlfs and A. Stecker to the oasis of Kufra in southeastern Cyrenaica in 1879. This was fol¬ 
lowed by Emilio Comalia’s list of taxa obtained by Guiseppe Haimann in Cyrenaica (Comalia in 
Haimann 1882) [also repeated in a second edition in 1886], and a summary of material collected 
around Benghazi by Gustav Ferdinand Ruhmer and described by von Martens (1883). Incidental 
to his ornithological work Alexander Konig (1888) reported on his own collections from North 
Africa, which included some reptiles from Tripoli in addition to a more comprehensive collection 
from Tunis and its surroundings. Boulenger (1891), likewise focused on the areas from Tunisia 
west to Morocco in his study of the Barbary Coast fauna, but did include some material from the 
Tripoli area. Some of the earliest Italian research on reptiles was published by Condorelli-Fran- 
caviglia (1896) and Rizzardi (1896), both dealing with Tripolitanian reptiles collected the year 
before by Antonio Balboni and Luigi Bricchetti-Robecchi, respectively. Werner (1909) critically 
analyzed some of the earlier papers and added a discussion of material obtained from the sur¬ 
roundings of Tripoli by Dr. Bruno Klaptocz in 1906, rounding out the herpetological contributions 
made during the Ottoman period in Libya. 
The era of Italian colonialism in Libya began in 1912, following the 1911-1912 Italo-Turkish 
War. Arnolfo Andreucci (1913) reported on his brother Augusto’s collections from Tripolitania and 
Boulenger (1914) discussed the 1912-1913 collections of Alfredo Andreini from Misrata and 
Homs to the east of Tripoli. Alessandro Ghigi (1913) summarized the Libyan fauna as a whole and 
subsequently focused on the vertebrates of Cyrenaica (Ghigi 1920, 1922). In the 1920s, major con¬ 
tributors to knowledge of the Libyan herpetofauna were Edoardo Zavattari, who reported on Cyre- 
