Abstract
Continental Africa contains more than 800 species of amphibians that correspond to the distinct and largely non-overlapping faunas of North Africa and sub-Saharan Africa (Channing and Rödel 2019; Escoriza and Ben Hassine 2019). The amphibian fauna of North Africa is dominated by salamander and frog species that colonized this region from western Europe in the Late Miocene (e.g., Carranza and Arnold 2004; Escoriza et al. 2006; Busack and Lawson 2008), possibly during the Messinian Salinity Crisis when there was contiguous land between these two regions (Krijgsman et al. 1999; Roveri et al. 2014). In comparison to other countries in North Africa, the amphibian fauna of Libya has received little attention, especially during the past forty years. The meager Libyan amphibian fauna is particularly interesting because the distinct North African and sub-Saharan faunas interdigitate and possibly overlap in Libya. There are many documented populations of frogs in small, isolated water bodies scattered across the large arid regions of the country. Taken together, these indicate that Libyan amphibians provide a unique opportunity for understanding the impact of dramatic climatic changes over the past 10,000 years on the water-dependent faunas in what were once one extensive lake and paleodrainage systems (Drake and Bristow 2006; Drake et al. 2008, 2011). With just five species, Libya has the most species-poor amphibian fauna of the 49 countries of continental Africa (Channing and Rödel 2019). Because its reptile fauna is also among the poorest in Africa (only 63 terrestrial species), Libya has received little attention from those interested in the herpetology of North Africa (Bauer et al. 2017; Escoriza and Ben Hassine 2019). Obviously, the lack of amphibian species diversity is not surprising given the vast majority of Libya’s landscape is dominated by the Sahara Desert with few perennial inland water bodies across most of the country. Of the five anuran species recorded in Libya, most records are of two species that are widespread across northern Africa: Pelophylaxsaharicusand Bufotesboulengeri. Pelophylaxsaharicus likely represents a colonization event of North Africa via the western Mediterranan (Beerli et al. 1996) but the direction from which Bufotes colonized this region is more ambiguous (Stöck et al. 2006). The other three species are restricted to southwestern Libya (Hoplobatrachus occipitalis, Ptychadena sp., Sclerophrysxeros) and represent isolated northern extensions of the fauna found in the Sahel along the southern fringes of the Sahara (Channing and Rödel 2019). While salamanders are known from neighboring Tunisia and Algeria, there are no records of salamanders from Libya (Schleich et al. 1996; Escoriza and Ben Hassine 2019). The earliest published record of frogs from Libya is that of Günther (1858a) who noted a record of what is now Bufotesboulengeri collected by J. Ritchie in Tripoli. However, the recently published journals of Hugh Clapperton (1788–1827) contain what is likely the earliest written European record of frogs in Libya (May 1822) from the springs near “Traghan” (now Taraghin) in the Fezzan: “there were a number of frogs hopping in and out and the sides were covered with their spawn” (Bruce-Lockhart and Wright 2000). These records likely correspond to the toad Bufotes boulengeri, which remains the only anuran reported from this area (Ibrahim 2008; Scortecci 1934, 1936). Half of the published research reporting amphibians from Libya comes from a 50-year period extending from the late 19th-century to early 20th-century (1880–1931). Most of these short publications are based on collections of amphibians and reptiles in the mesic regions near the Mediterranean coast (Peters 1880, 1881; Haimann 1886; Rizzardi 1896; Werner 1909; Andreucci 1913; Ghigi 1913, 1919–1920, 1922; Hartert 1913, Boulenger 1914; Zavattari 1922; Calabresi 1923; Vincinguerra 1927; Gestro and Vincinguerra 1931; see Bauer et al. 2017 for a more extensive summary). Scortecci (1936, 1937a) provided the first extensive documentation of frogs from the interior deserts of Libya (i.e., the Fezzan), especially Ghat in extreme southwestern Libya. During the middle and late 20th-century, the reports of Libyan frogs are restricted to the few records from Schnurrenberger (1963) and Sayers (1964) and Dumont’s (1982, 1987) reports of several species from Ghat. The most significant works from this period are by Resetar (1981) and Schleich (1987) on the amphibians and reptiles from in or near what is now El-Kouf National Park near the coast in eastern Libya. The last 20 years has seen a slow resurgence of interest in Libyan frogs, including records from new localities in the interior deserts (Frynta et al. 2000; Doria et al. 2001–2002; Ibrahim and Ineich 2005; Stöck et al. 2006; Jdeidi 2008; Essghaier et al. 2015). Both Ibrahim (2013) and the recent book on North African amphibians by Escoriza and Ben Hassine (2019) that provide summaries on the biology of several anuran species in Libya. The availability of standing water in the Sahara has changed dramatically during the late Pleistocene. River systems and large paleolakes, such as Megachad and Megafazzan, dominated parts of the central Sahara during 100,000–500,000 years ago (Drake and Bristow 2006; Drake et al. 2008, 2011), likely with repeated expansions and contractions following regional climatic changes. At least some of these water bodies persisted into the early Holocene when human communities living along lake margins fished for Nile perch, catfish, hippos, turtles, and crocodiles (Sereno et al. 2008). Whereas a few relict populations of crocodiles persisted into the 19th or early 20th century (Brito et al. 2011), most large-bodied vertebrates associated with rivers and lakes no longer occur in the central Sahara. Yet many invertebrates, such as dragonflies and snails, and small vertebrates, including fish in the genus Barbus and some frogs, persist at isolated oases across the Sahara (Pellegrin 1911, 1919, 1934; Monod 1921; Seurat 1935; Dumont 1978, 1982; Lévêque 1989, 1990). Phylogeographic studies of these species in an explicit paleohydrological context could provide a unique opportunity to understand how the dramatic climatic changes over the past 10,000 years impacted these now-isolated populations. Based on our complete survey of the literature, we provide a gazetteer of localities with frogs from XX sites in Libya. In addition, surveys of data from museum records provide a small handful of additional records from localities that are otherwise not represented in the literature. Significant to the conservation of frogs at isolated water bodies across Libya, we note that there are at least ## localities for which there are no published records in the past ## years. We intend for this gazetteer of Libyan frogs to complement the recent work on North African amphibians by Padial et al (2004, 2013), Baha El Din (2006), Mateo et al. (2013), Beukema et al. (2013), Bouazza et al. (2021), and the excellent recent summary by Escoriza and Ben Hassine (2019). We hope that our work inspires a new generation of Libyan scientists eager to study frogs.