Contribution to the knowledge of Tunisian amphibians - Herpetology

Introduction

Amphibians have declined dramatically in many areas of the world during the past 25 years (Wake, 1991; Bee-bee, 1996; Beebee and Griffiths, 2005) and the pheno-menon seems to depend mostly on the effect of human activities on natural ecosystems (Collins and Storfer, 2003). In the light of the Global Amphibian Decline (G.A.D.) a sensitive monitoring approach is necessary to determine the distribution and abundance of amphi-bian populations, to assess whether they are declining, and to quantify the extent of declines (Storfer, 2003).

Tunisia is the easternmost country of the Maghreb, the only part of Africa with a clearly Palaearctic amphibian fauna; Maghrebian amphibian fauna is characterized by low species diversity and by a rather high level of endemism (Salvador, 1996; Schleich, Kästle and Kabi-sch, 1996). The landscape diversity of Tunisia is rather high; this influences the zoogeography of the country and makes the study of its fauna of great interest. To date, no extensive data are available on the amphibians of Tunisia, although some biological and faunistic stu-

dies have been conducted in the country since the end of the 18th century (i.e. Boulenger, 1891; Mayet, 1903; Gauthier, 1928; Mertens, 1929; Mosauer, 1934; Blank, 1935; Gallien, 1948; Domergue, 1959; Lanza and Bruz-zone, 1959; Schneider, 1974; Schneider, 1978; Blank and Shane, 1981; Blank and Nouira, 1988; Steinwarz and Schneider, 1991; Salvador, 1996; Schleich, Kästle and Kabisch, 1996; Meddeb and Cheniti, 1998; Noui-ra, 2001; Pasmans and Bogaerts, 2001; Pasmans et al., 2002; Busack, 2006; Meddeb et al., 2007).

The aims of this study are (1) to improve the know-ledge on Tunisian amphibian fauna (2) to analyse the distribution patterns of the studied species (3) to realize a database on the distributional records of the amphibi-ans in Tunisia and compare our data with those availa-ble in literature. We report original data on distribution, breeding phenology and habitat features of the observed species and present the first data on the ecology of the Tunisian populations of the poorly known salamandrid Pleurodeles nebulosus. The scattered and sparse know-ledge on Tunisian amphibians is reviewed and comple-ted with original data in order to better understand the ecology and the distribution of the species. We produ-ced distributional maps and analysed how the climatic factors could influence the distribution of the studied species. The taxonomical status of the studied amphibi-ans was reviewed based on the more recent literature.

The present database represents a first step towards the realization of an extensive census of Tunisian amphibi-

Herpetology Notes, volume 2: 107-132 (published online on 04 August 2009)

Contribution to the knowledge of Tunisian amphibians: notes on distribution, habitat features and breeding phenology

Alessandra Sicilia1*, Federico Marrone1, Roberto Sindaco2, Souad Turki3 and Marco Arculeo1

1 Dipartimento di Biologia Animale, Via Archirafi, 18, 90123 Palermo, Italy.

2 Istituto per le Piante da legno e l’Ambiente S.p.A., Corso Casale, 476, 10132 Torino, Italy.

3 Institut National des Sciences et Technologies de la Mer (INSTM), Port de pêche, 2016 la Goulette, Tunisia.

* Corresponding author; email: [email protected]

Abstract. In order to improve the knowledge on amphibian fauna in Tunisia, about 300 water bodies belonging to different habitat typologies were sampled during the period 2004-2008. At the sites where amphibians were observed, we recorded the UTM coor-dinates, the altitude, and the following environmental features: habitat typology, hydroperiod, electric conductivity, temperature, water turbidity, and macrophyte coverage. The presence of the seven species currently known with certainty for Tunisia has been confirmed: Pleurodeles nebulosus, Discoglossus pictus, Bufo bufo, B. mauritanicus, B. boulengeri, Hyla meridionalis and Pelo-phylax saharicus. Conversely, no evidence on the occurrence of Salamandra algira in Tunisia, currently considered as doubtful, was collected. Furthermore, all the information available in the literature has been summarized and original data on distribution ranges, breeding phenology and habitat features of the observed species are reported; in particular, the first data on the ecology of the poorly known Pleurodeles nebulosus are provided. Present work represents a contribution towards the realization of an extensive atlas of Tunisian amphibian fauna to be used as a management tool for the conservation of Tunisian inland waters and their biota.

Key words. Amphibians, Tunisian fauna, distribution, conservation, checklist

Alessandra Sicilia et al.108

ans to be used as a management tool for the conservati-on of Tunisian wetlands, which are extremely important but threatened ecosystems in this arid country.

Study area

Tunisia is located in the easternmost part of the Maghreb, between Algeria and Libya (Fig. 1). It encompasses an area of about 164000 km² with about 1300 km of coastline. The country is characterised by several geographically and climatically diffe-rent zones. According to the Köppen-Geiger system, the climate of the country belongs to four different categories: (1) temperate region with hot and dry summer (Csa, Mediterranean), (2) arid hot steppe (BSh), (3) arid cold steppe (BSk) and (4) arid hot de-sert (BWh) (Peel, Finlayson and Mcmahon, 2007). The Mediter-ranean area of the country includes the northern part of the eas-ternmost Atlas Mountains (Medjerda Mountains), the Medjerda valley, the alluvial plain of Medjerda river (the plain of Tunis) and Cap Bon Peninsula. The Medjerda Mountains represent theThe Medjerda Mountains represent the wettest area of the country; in the western part of this area the area of the country; in the western part of this area the precipitation value ranges between 800 and 1500 mm y-1 and the average annual temperature is 17-18 °C. Cap Bon Peninsula and the plain of Tunis are characterised by lower annual rainfall (400-600 mm) and an annual average temperature of about 17-19 °C.

South of the Medjerda valley there is a steppic plateau named Tell, extending through the central-western area of Tunisia; close to the Algerian border occurs the highest peak of the country, the Djebel (Mount) Chambi (1544 m). The average annual tempera-tures in this area are comprised between 15 and 19 °C, and the

rainfall ranges from 200 to about 600 mm y-1. The steppic plain in central Tunisia along the Mediterranean coast of the country is named Sahel; it is located between the Gulf of Hammamet and the Gulf of Gabés (annual rainfall of about 200-400 mm; average an-nual temperature of about 19-20 °C). South of the oases of Gafsa and Gabés begins the desert, represented by the desert steppe, the Great Eastern Erg (sand-dune desert), the hamada (rocky desert) and the chotts (salt-lakes depressions). It is characterized by an-nual rainfall values ranging between 75 and 200 mm and by an average temperature comprised between 19 and 22 °C.

Materials and methods

About 300 Tunisian water bodies belonging to different habi-tat typologies were sampled during the period 2004-2008. At the sites where amphibians were observed, the coordinates, the alti-tude, and the reproductive parameters (presence of pairs, male calling activity, eggs, and larvae) were registered. When possible, the following environmental features were recorded: habitat ty-pology, estimated hydroperiod length, turbidity, macrophyte co-verage, water electric conductivity (microSiemens/cm at 20°C), and water temperature (°C). An arbitrary range value from 1 to 3 was established to characterize the water turbidity (1 = crystal clear water; 3 = extremely turbid water or argillotrophic system) and the macrophyte coverage (1 = total absence of vegetation; 3 = absence of open water).

Species distribution and ecological data were summarized based from both new and literature data published from 1891 to 2007 (Boulenger, 1891; Mayet, 1903; Gauthier, 1928; Mertens, 1929;

Figure 1. Location of Tunisia in the Mediterranean basin.

Distribution of Tunisian amphibians 109

Mosauer, 1934; Blank, 1935; Gallien, 1948; Lanza and Bruzzone, 1959; Schneider, 1978; Blank and Shane, 1981; Blank and Noui-ra, 1988; Steinwarz and Schneider, 1991; Meddeb and Cheniti, 1998; Plötner, 1998; Nouira, 2001; Plötner and Ohst, 2001; Pas-mans et al., 2002; Harris, Batista and Carretero, 2003; Carranza and Arnold, 2004; Batista et al., 2006; Busack, 2006; Stöck et al., 2006; Zangari, Cimmaruta and Nascetti, 2006; Lymberakis et al., 2007; Meddeb et al., 2007; Recuero et al., 2007). No in-formation on amphibian distribution or ecology was found in the other reviewed studies dealing with Tunisian herpetofauna (e.g. Anderson, 1892; Werner, 1897). Since most bibliographic records are provided without coordinates, we tried to retrieve them based on the locality names reported in the works themselves. Some bi-bliographical data were omitted because of the lack of exact dis-tribution information (e.g. some sites reported by Salvador, 1996; Schleich, Kästle and Kabisch, 1996; Le Berre, 1989), or when they did occur in areas where we already acquired geo-referenced sites for the species involved.

The database was imported into the Geographical Information System (GIS) software ArcGis 9, for mapping process. The dis-tribution data were plotted to the climatic maps (average annual precipitation and average annual temperature) in order to under-stand if and how these physical factors influence the distribution of the species.

Climatic data were kindly provided by the “Institut National de la Metéorologie, Republique Tunisienne” based on the period 1971-2000 (Figures 9 and 10). The coordinates of the sites used in the Appendix I belong to the UTM-WGS84 system.

Results and notes on the observed species

A total of 350 records of amphibians in Tunisia were gathered from our own field data (218 records) and lite-rature records (132). The presence of seven species has been confirmed, one caudate: Pleurodeles nebulosus and six anurans: Discoglossus pictus, Bufo bufo, B. maurita-nicus, B. boulengeri, Hyla meridionalis and Pelophylax saharicus. Conversely, no evidence was collected on the presence of Salamandra algira in Tunisia, reported by some authors for the north-westernmost part of the country (see discussion). Habitat information and distri-bution data are summarized in Appendix I. The distribu-tion maps of Tunisian amphibians are shown in Figures 2-8, the climatic maps in Figures 9-10.

Pleurodeles nebulosus (Guichenot, 1850)Pleurodeles is a salamandrid genus endemic to the

Iberian Peninsula and north-western Africa. The syste-matics and general distribution of the species belonging to this genus were clarified by several recent researches (Carranza and Arnold, 2004; Carranza and Wade, 2004; Veith et al., 2004). The Algerian ribbed newt P. nebulo-sus is distributed across northern Tunisia and Algeria, except in the Edough Peninsula and surrounding low-

land areas, where it is replaced by the endemic P. poireti (Carranza and Wade, 2004).

To date, the ecology of P. nebulosus is almost unknown. Pasmans et al. (2002) collected some field data on ha-bitats and morphological characters of Tunisian Pleuro-deles. They found most of the newts under stones and in clay piles, and assumed that temporary, shallow ditches and ponds in river plains were suitable breeding habitat for the species. Furthermore they observed that animals collected north of the Medjerda Mountain are smaller than those found south of the mountain range.

In the frame of the present study, all the observed reproductive sites (a total of 32) were wetlands with a temporary hydroperiod (swamps, marshes, ponds, floo-ded fields, pools and road ditches) with the exception of a single permanent pond. Larvae were found from the beginning of December (near Tabarka) to the first half of April (in the plain of Tunis). Adults and larvae were observed in water bodies with water temperatures ranging from 10 to 26.8 °C, water electric conductivity values ranging from 221 to 2500 μS cm-1 and turbidity and macrophyte coverage values ranging from 1 to 3. The observed altitudinal distribution ranges from sea level up to 457 m a.s.l. (in the Medjerda Mountains). The highest altitude known for this species in Tunisia is 689 m a.s.l, near Forêt des Chênes (8 km south of Aïn Draham), where Pasmans et al. (2002) found a juvenile specimen. Six reproductive sites of P. nebulosus were found in Cap Bon Peninsula, where the species was not previously known to occur. Pleurodeles nebulosus is exclusively present in the temperate areas (Fig. 2; ave-rage annual precipitation ranging from 400-450 mm in Cap Bon Peninsula and in the vicinity of Tunis, to 1500in the vicinity of Tunis, to 1500, to 1500 mm on the Medjerda Mountain), with an average annual temperature of 17-18 °C. The species’ southern distribu-tion limit in Tunisia seems to be the Medjerda valley.

Discoglossus pictus Otth, 1837The painted frog D. pictus is distributed in the Magh-

reb and in the islands of Sicily, Malta and Gozo. Intro-duced populations, whose source is considered African D. pictus (Zangari, Cimmaruta and Nascetti, 2006), are present in southern France and north-eastern Spain (Castanet and Guyetant, 1989; García-París, Montoni and Herrero, 2004). The taxonomical validity of the sub-species D. pictus auritus Hèron-Royer, 1888 for eastern Maghrebian populations does not seem to be supported by genetic analyses (Fromhage, Vences and Veith, 2004; Zangari, Cimmaruta and Nascetti, 2006).

The species is quite common in northern and central

Figure 2. Distribution map of Pleurodeles nebulosus in Tunisia.



Figure 3. Distribution map of Discoglossus pictus in Tunisia.


Figure 4. Distribution map of Bufo bufo in Tunisia.


Figure 5. Distribution map of Bufo mauritanicus in Tunisia.


Figure 6. Distribution map of Bufo boulengeri in Tunisia.


Figure 7. Distribution map of Hyla meridionalis in Tunisia.


Figure 8. Distribution map of Pelophylax saharicus in Tunisia.


Figure 9. Map of Tunisia with average annual precipitation and amphibian distribution records.


Figure 10. Map of Tunisia with average annual temperature and amphibian distribution records.

Tunisia where it breeds in virtually every habitat typolo-gy: temporary marshes, temporary swamps, temporary and permanent ponds and pools, flooded fields, ditches, puddles, concrete reservoirs, temporary and permanent streams, and pools in the bed of dry streams. The spe-cies seems to become increasingly rare southwards (Fig. 3). The southernmost sites we found where the species occured were irrigation canals and a concrete reservoir in an oasis near Gabés; in the same area it was reported by Busack (2006). Mayet (1903) reports the occurrence of the species in the oases of the Chott El Djerid area. It is reported from the oases of Gafsa (Amor et al., 2007), Tozeur (Boulenger, 1891; Gauthier, 1928; Gallien, 1948) and Nefta (Boulenger, 1891). As far as Tozeur is concer-ned, Gauthier (1928) reports to have found Discoglos-sus in the irrigation channels of the oasis, and Gallien (1948) reports the presence of both the D. pictus forms, the “striped” and the “spotted” one. Both the forms of D. pictus were also observed in the frame of our resear-ches in the north and in the centre of the country. The species is also present on “La Galite” island (Boulenger, 1891; Mayet, 1903; Lanza and Bruzzone, 1959; Zanga-ri, Cimmaruta and Nascetti, 2006). Our data confirms that D. pictus is an opportunistic species which is able to successfully breed in every season of the year (Riggio, 1976; Schleich, Kästle and Kabisch, 1996). We obser-ved tadpoles in water bodies with water temperatures ranging from 6.4 to 29.3 °C, water electric conductivity ranging from 165 to 14480 μS cm-1 and turbidity and macrophyte coverage values ranging from 1 to 3.

In Tunisia, the observed altitudinal distribution ranges from sea level up to 871 m a.s.l., near Maktar, in the Tell area. The average annual precipitation of the inhabited area ranges from 100 (desert zone) to 1500 mm (tem-perate zone) and the average annual temperature from 16 to 21 °C.

Bufo bufo (Linnaeus, 1758)The common toad, B. bufo, is distributed in most of

Europe, southern Siberia (east to Lake Baikal), Near East, and in North Africa, where it is a relict species in the moist and cold mountain areas of Morocco, Algeria and Tunisia (Schleich, Kästle and Kabisch, 1996).

In Tunisia it is a rare species and it seems to be present only on the westernmost part of the Mejerda Mountains (Fig. 4). Blank (1935) and Schneider (1974) report the species to occur near Aïn Draham. Our data confirm this report through the finding of an adult male specimen in an oak forest near the reservoir of Beni M’tir, at 631 m a.s.l. The toad was found at the beginning of March. No

other data are available on the Tunisian populations of this species. The average annual precipitation of our site is about 1500 mm (temperate zone), the average annual temperature is 17 °C.

Bufo mauritanicus Schlegel, 1841The Berber toad is distributed throughout the Magh-

reb (Schleich, Kästle and Kabisch, 1996). Reports from Mali, Mauritania and Niger (Salvador, 1996; Schleich, Kästle and Kabisch, 1996; Padial and De La Riva, 2004) are doubtful; an introduced population is present in southern Spain (IUCN, 2006).

We found this species in temporary and permanent streams, concrete reservoirs, agricultural reservoirs, ponds and pools. Tadpoles were observed from the middle of February to the end of November (near Kai-rouan). Schneider (1978) and Salvador (1996) report reproductive activity during spring and summer. The species is reported to occur in the Tunisian oases of Gafsa (Boulenger, 1891; Mosauer, 1934; Amor et al., 2007) and Tozeur (Boulenger, 1891; Mosauer, 1934; Schneider, 1978). According to Schneider (1978) the northern border of the Sahara desert seems to represent the southern limit of the distribution range of B. mauri-tanicus (Fig. 5).

The altitudinal distribution range is from sea level up to 1077 m a.s.l. (near Thala, Tell plateau). The average precipitation range in its distribution area is from about 100 (desertic zone) to about 900 mm y-1 (temperate zone). The average annual temperature ranges from 15 to 21 °C.

Bufo boulengeri Lataste, 1879Stöck et al. (2006) genetically analyzed North African

green toads and demonstrated that the populations from western Morocco to eastern Egypt (previously reported as “Bufo viridis”) differ from the other Palaearctic po-pulations, and that the available name for the North Af-rican clade is Bufo boulengeri Lataste, 1879. Further-more, Stöck et al. (2008B) studied populations from the entire Tunisian territory and main islands, confirming the presence of this species throughout the country, and also on the island of Lampedusa, politically Italian but located on the African shelf.

Bufo boulengeri is widespread in most of Tunisia (Fig. 6). The southernmost breeding site we found was a tem-porary stream in the rocky desert (Hamada) near Tata-ouine. We also found two populations near El Faouar and Kebili, in the sandy desert (erg). The species seems to be quite common in the Kerkennah and Djerba islands


(our data; Mertens, 1929). Tadpoles were observed from the beginning of February (Cap Bon Peninsula) to June (in the hamada). In 2005, B. boulengeri were observed to breed between the end of autumn and the beginning of winter in Kerkennah Islands. According to Salvador (1996) the green toad in Tunisia usually breeds from February until May, though another breeding period, in August and September is also recorded. Mertens (1929) reports late summer reproduction on Djerba Island.

The breeding sites we found belong to the following habitat typologies: temporary swamps, ponds inside salty depressions, saltmarshes, temporary and perma-nent streams, temporary pools and ponds, pools in the bed of dry streams, road ditches, agricultural reservoirs, and concrete reservoirs. Gauthier (1928) reports the presence of the species also in freshwater marshes. We found tadpoles in water bodies with water temperatures ranging from 9.4 to 23.6 °C, water electric conducti-vity ranging from 309 to 16850 μS cm-1 and turbidity and macrophyte coverage values ranging from 1 to 2.5. Adults were observed in a saltmarsh with a water elec-tric conductivity value of 23700 μS cm1.

The altitudinal distribution of the species in Tunisia ranges from sea level up to 1077 m a.s.l. (near Thala, Tell plateau). The average annual precipitation range of its distribution area is from 75 (desert zone) to about 1200 mm (temperate zone), the average annual tempe-rature is from 15 to 21 °C.

Hyla meridionalis Boettger, 1874The Mediterranean tree frog, Hyla meridionalis, is

distributed in the western Mediterranean area: southern France, coastal north-western Italy, north-eastern and south-western Iberian Peninsula, and northern Africa (Morocco, northern Algeria, and northern Tunisia); like-ly introduced populations are present also on the Canary Islands, Madeira and Menorca (Recuero et al., 2007). A recent survey on the phylogeography of H. meridionalis has shown that Tunisian populations are highly diver-gent from the Iberian and Moroccan ones, suggesting an ancient split between the two lineages (Recuero et al., 2007) and, according to Stöck et al. (2008A) Tu-nisian and Algerian populations could represent a new, still undescribed, species. According to our dataset in Tunisia the species is present north of the Medjerda val-ley, in the Medjerda Mountains and in the coastal plain north of the mountains (Fig. 7). We did not record the species in the plain of Tunis, although it was reported to occur near Tunis and in the Bardo garden (Tunis) by Blank (1935). Conversely, we found the Mediterrane-

an tree frog also in Cap Bon Peninsula. Breeding sites are mostly represented by temporary swamps, but also pools, temporary ponds, marshes and agricultural reser-voirs are used by the species. Hyla meridionalis seems to breed from the beginning of February to the end of March. The altitudinal distribution ranges from sea le-vel up to 457 m a.s.l. Tadpoles were found in water bo-dies with water temperatures ranging from 11 to 18 °C, water electric conductivity ranging from 274 to 939 μS cm-1, turbidity and macrophyte coverage values ranging from 1 to 2.5.

The average annual precipitation range of its distribu-tion area ranges from 400 to 450 mm y-1 near Tunis and in Cap Bon Peninsula, to 1500 mm y-1 (temperate area) on the Medjerda Mountain. The average annual tempe-rature ranges from 17 to 18 °C.

Pelophylax saharicus (Boulenger, 1913)The north African green frog populations were ascri-

bed to “Rana saharica” (i.e. Pelophylax saharicus) by several authors (Buckley et al., 1994; Llorente et al., 1996; Arano et al., 1998; Plötner, 1998; Plötner and Ohst, 2001; Harris, Batista and Carretero, 2003; Lymberakis et al., 2007); however the Maghrebian populations seem to be composed of two distinct clades, one occurring in Morocco and the other one in Algeria and Tunisia (Buckley et al., 1994; Arano et al., 1998). The geogra-phical limit of the two groups is, to date, unknown. Ara-no et al. (1998) suggest that the two clades should be recognized as different subspecies: P. saharicus sahari-cus (eastern clade) and P. saharicus riodeoroi (western clade). Harris, Batista and Carretero (2003) confirm this based on molecular data.

The species is present in Western Sahara, Morocco, Algeria, Libya, Tunisia and Egypt. Introduced popu-lations are also present on the Canary Islands (Lever, 2003).

In Tunisia the species occurs in almost the entire coun-try, reaching also the Sahara (Fig. 8). The north African green frog is relatively common in the irrigation canals of the oases, but it also occurs in the streams of the de-sert area. Active adults were observed throughout the year in all the country, and tadpoles were collected from February (in the central part of the country) to the be-ginning of the summer (throughout the whole country). We found the species in pools in the bed of dry streams, temporary and permanent ponds or pools, tempora-ry marshes, road ditches, temporary swamps, flooded fields, concrete reservoirs, irrigation canals, temporary and permanent streams, and agricultural reservoirs and


ditches. The water temperature of the sites ranged from 10.3 to 24.2 °C, electric conductivity ranged from 223 to 9600 μS cm-1, turbidity values ranged from 1 to 3, and macrophyte coverage from 1 to 2.5.

According to a study carried out by Meddeb and Che-niti (1998) on the feeding habits of two populations in the north of the country (a permanent pond in Radès and a temporary one near Menzel Bourguiba), P. saha-ricus demonstrated a remarkable flexibility in its diet, showing also cannibalism on tadpoles and juveniles. Insects, arachnids, crustaceans, anellids and snails be-long to the diet of the studied populations. Although the Radès population of P. saharicus seems to prefer He-teroptera and Gastropoda prey, the Menzel Bourguiba population seems to have a less specialized diet. Med-deb et al. (2007) studied age structure and growth of the same populations; no males older than 5 years were found, whereas the females, attaining sexual maturity when 3 years old, can reach 7 years of age. It seems that there is not a clear sexual size dimorphism. The “lines of arrested growth” (LAGs) were generally very distinct suggesting a well-defined annual inactivity period. Ac-cording to the authors, P. saharicus hibernates in mud from December to February.

The known altitudinal distribution in Tunisia ranges from sea level up to 1077 m a.s.l. (near Thala, Tell pla-teau). The average annual precipitation range of the spe-cies distribution area is from 75 (desertic zone) to 1500 mm y-1 (temperate zone), the average annual tempera-ture from 15 to 21 °C.

DiscussionWe did not find any evidence of the presence of Sala-

mandra algira in Tunisia. The currently known distribu-tion of the species comprises a few humid or sub-humid forested regions in Morocco and Algeria (Escoriza et al., 2006), and the species’ presence in Tunisia is in fact considered uncertain. This salamandrid was reported to be present in the country by Domergue (1959) later ci-ted by Salvador (1996); conversely, Boulenger (1891), Mayet (1903), Blank (1935), and Schleich, Kästle and Kabisch (1996) did not include it in Tunisian fauna. Although Blank (1935) did not observe the species, he explicitly did not exclude its presence on the Medjerda Mountains. Proper surveys are needed to definitively ex-clude or confirm its presence in the Tunisian territory.

As far as the other species are concerned, this paper re-ports distributional and ecological data which improve the available knowledge on these species. All the spe-cies known to occur in Tunisia are present in the Me-

diterranean part of the country. Pleurodeles nebulosus and Hyla meridionalis seem to be absent in the southern steppic and desert areas, although they are also present in Cap Bon Peninsula and in the southern part of Tu-nis plain, the drier areas of the temperate zone (average annual precipitation value of 400-450 mm y-1). The ob-served southern distribution limit of the Algerian ribbed newt in Tunisia is the Medjerda valley (near Jendouba), while H. meridionalis finds its southern limit along the southern slope of the Medjerda Mountains. Despite the restricted distribution area of the two species in the country, P. nebulosus and H. meridionalis are rat-her common in the wettest area of the temperate zone, where the geomorphology and the nature of the substra-te allow the presence of a large number of suitable wet-lands for the reproduction of these species, mostly tem-porary water bodies like swamps, ponds and marshes. Bufo bufo is present only in the westernmost part of the Medjerda Mountains, close to the Algerian border. All the other species reach the desert zone, where D. pictus is to date not recorded in natural ecosystems, but only in the irrigation canal systems of some oases. Pelophylax saharicus and B. boulengeri were observed in the natu-ral environment of the sandy and rocky deserts, and in the irrigation systems of the oases. Bufo mauritanicus is reported for the oases of Gafsa and Tozeur (Boulen-ger, 1891; Mosauer, 1934; Schneider, 1978; Amor et al., 2007), but no data on the nature of its breeding sites in the desert are available. Altogether, P. saharicus and B. boulengeri seem to be the most widespread species, alt-hough D. pictus is the most common species in the nor-thern and north-central part of the country, being able to breed in every kind of water body, including fairly small and ephemeral pools and puddles.

All the observed species do not seem to be sensibly influenced by the water turbidity or the abundance of macrophytes. The highest value of water electric con-ductivity was recorded for the species B. boulenge-ri (23700 μS cm-1) and D. pictus (14480 μS cm-1). As already reported, these species are able to breed also in brackish water; unfortunately no data on the water electric conductivity of B. mauritanicus breeding sites were recorded, although according to Schneider (1978) it could breed also in brackish waters. Discoglossus pic-tus and P. saharicus show a wide ecological plasticity and we observed them in a wide range of different water bodies.

The studied species show different phenologies and breeding strategies in different parts of the country and when compared with those observed in other regions


of the Maghreb (Salvador, 1996; Schleich, Kästle and Kabisch, 1996); this is likely a result of the climatic differences occurring in North Africa and of the unpre-dictable availability of water and suitable condition for breeding activity, which is a characteristic of the arid regions. In particular, B. boulengeri is an opportunis-tic breeder with a wide margin of variability in annual reproductive cycle patterns, as observed also in other xeric areas of the Mediterranean basin (Kyriakopoulou-Sklavounou, 2000; Sicilia, 2006; Sicilia et al., 2006). Bufo mauritanicus seems to have a long reproductive period as well, breeding from the middle of February to almost the late summer or early autumn, and tadpo-les were observed till the end of November (Kairouan area). Moreover, although P. saharicus LAGs are ge-nerally very distinct suggesting a well-defined winter inactivity period in northern populations, we found active frogs all the year long even in northern Tunisia, which is the coldest part of the country. It may depend on the different characteristics of the studied sites, but it is also possible that during the winter the activity of P. saharicus is remarkably reduced, and that it alternates hibernation and short active phases depending on the climatic conditions.

Finally, the current known distribution of the species has been checked and completed with original records. In particular, P. nebulosus was up to now not known to occur in Cap Bon Peninsula and almost no data were available on the ecology of this species. In the more recent studies and reviews of Northern African amphi-bians, Schleich, Kästle and Kabisch (1996) report D. pictus only for the northern part of Tunisia (with the ex-ception of Cap Bon Peninsula), while Salvador (1996) reports D. pictus in the south of the country, with no precise information on the locality, and do not report the presence of H. meridionalis in Cap Bon Peninsula. Finally Salvador (1996) reported the occurrence of B. bufo in the plain of Tunis, where the presence of the spe-cies is rather unlikely, while it is hereby confirmed for the Medjerda Mountains. Some records (e.g. D. pictus in the southern oases and H. meridionalis in the plain of Tunis) are rather old and need to be confirmed.

To conclude, the present paper summarizes all the in-formation available in the literature on the distribution and ecology of amphibians in Tunisia. Furthermore, it reports an amount of original data with the aim of cons-tituting a basis toward the realization of an extensive at-las of Tunisian amphibian fauna and providing a dataset to be used for the conservation of the amphibians and their habitats in the country.

Acknowledgements. The authors wish to thank Agostino D’Amico, Maurizio Sarà and Ibrahim Turki for the help they gave to the realization of this research.

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Appendix I List of localities and their amphibian fauna. Latitude and longitude are given in Universal Transverse of Mercator system (UTM, map datum: WGS84).

Locality code

LAT. LONG. Locality Habitat Species Source

064M 3949359 32 610523 Near Kairouan Road ditch Discoglossus pictus New datum 064Mb 3949410 32 610470 Near Kairouan - Bufo mauritanicus New datum F001 4075191 32 615342 Near La Goulette, Tunis Road ditch Discoglossus pictus New datum F002 4074972 32 615072 Near La Goulette, Tunis Road ditch Discoglossus pictus New datum F006 4082626 32 616132 La Marsa Temporary marsh Discoglossus pictus

Pelophylax saharicus New data

F011 4113558 32 567500 Tinja Temporary pond Discoglossus pictus New datum F014 4086834 32 478236 Near Tabarka Temporary swamp Pleurodeles nebulosus

Discoglossus pictus New data

F015 4101142 32 520759 Sejenane Temporary swamp with slow flowing water

Pleurodeles nebulosus Discoglossus pictus Hyla meridionalis

New data

F016 3855467 32 706493 El Kraten, Chergui, Kerkennah islands

Countryside Bufo boulengeri New datum

F016b 3842494 32 701560 Remla, Kerkennah islands

Countryside Bufo boulengeri New datum

F021 3742610 32 550768 Between Gabés and Kebili

Pool in the bed of a dry stream

Pelophylax saharicus New datum

F032 3919170 32 549202 Hadjeb el Aioun Temporary swamp Discoglossus pictus New datum F033 3961136 32 603322 Near Kairouan Temporary pond Discoglossus pictus


F034 3957179 32 600955 Near Kairouan Road ditch Discoglossus pictus Pelophylax saharicus

New data

F037 4090434 32 609633 Raoued Temporary pond Pleurodeles nebulosus New datum F041 4090709 32 606982 Raoued Temporary swamp Pleurodeles nebulosus New datum F045 4089622 32 598455 Bou Hnach Temporary pond Pelophylax saharicus New datum F053 4107186 32 591150 Near Dawwar al

Huwayd Astatic swamp Pelophylax saharicus New datum

F054 4113914 32 567377 Tinja Temporary pond Discoglossus pictus New datum F061 4064095 32 605740 Fouchana Temporary marsh Discoglossus pictus New datum F063 4066248 32 612934 Near Ben Arous Road ditch Discoglossus pictus New datum F068 4062655 32 630509 Soliman Temporary swamp Discoglossus pictus New datum F072 3958411 32 588773 El Baten Pool in the bed of

a dry stream Discoglossus pictus New datum

F073 3989026 32 595385 Sidi Naji Temporary pool Discoglossus pictus New datum F077 4071953 32 613846 Rades Temporary swamp Discoglossus pictus New datum F088 4059842 32 633394 Near Soliman Flooded field Discoglossus pictus New datum F089 4060389 32 646843 El Itha Country side Discoglossus pictus New datum F090 4059971 32 650914 Near Menzel Bou Zelfa Pools Discoglossus pictus



F091 4060483 32 651516 Near Menzel Bou Zelfa Flooded field Pleurodeles nebulosus Discoglossus pictus Pelophylax saharicus

New data

F092 4065080 32 654317 Near Ad Damus Marsh close to a permanent stream

Pleurodeles nebulosus Discoglossus pictus

New data

F093 4066287 32 659919 Near Oum Dhouil Pool in the bed of a dry stream


New data

F094 4066148 32 660158 Near Oum Dhouil Marsh in the bed of a dry stream


New data

F097 4078735 32 686828 Kelibia Temporary swamp Discoglossus pictus Pelophylax saharicus

New data

F099 4099366 32 680480 Near El Haouaria Flooded field Discoglossus pictus Pelophylax saharicus

New data

F100 4103550 32 678315 Near El Haouaria Temporary stream Discoglossus pictus New datum F101 4101609 32 678691 El Haouaria Pool Discoglossus pictus New datum F102 4088131 32 665525 Near Enchaa Pool Discoglossus pictus New datum F103 4085385 32 664820 Enchaa Temporary swamp Pleurodeles nebulosus

Discoglossus pictus Hyla meridionalis

New data

F104 4066537 32 672509 Near Manzil Hurr Temporary swamp Discoglossus pictus Bufo boulengeri Pelophylax saharicus

New data

F107 4094739 32 594465 El Kantara Permanent pond Pelophylax saharicus New datum F108 4107422 32 562865 Ichkeul Park, near

MateurPools Pelophylax saharicus New datum

F110 4091002 32 582792 Near Dawwar al Mabtusah

Permanent pond Discoglossus pictus New datum

F116 3825650 32 643108 Near Nuqtah Pools in the bed of a dry stream

Bufo boulengeri New datum

F117 3820910 32 636617 Mahares Pools in the bed of a dry stream


F120 3743629 32 605313 Gabès Irrigation canals and concrete reservoir

Discoglossus pictus Pelophylax saharicus

New data

F126 3700659 32 501458 Douz Irrigation canals Pelophylax saharicus New datum F127 3719201 32 500880 Near Sidi Messaïd Stream Bufo boulengeri New datum F131 3797595 32 494042 Chott El Guettar, Al

QattarSwamp in a salty depression


F132 3798638 32 496635 Al Qattar Concrete reservoir Pelophylax saharicus New datum F134 3803465 32 484510 Oued Melah, near Lala Pool in the bed of

a dry stream Bufo boulengeri New datum

F136 3906710 32 527846 Near Sbeitla Temporary pool Pelophylax saharicus New datum F137 3908039 32 533270 Near Sbeitla Country side Discoglossus pictus New datum F138 3935401 32 568516 Near Funduq al ‘Uqbi Reservoir Pelophylax saharicus New datum F139 3935691 32 569269 Near Funduq al ‘Uqbi Temporary pool Bufo boulengeri New datum F140 4043168 32 588325 Near Bi’r al Mashariqah Country side Bufo mauritanicus New datum F141 4048058 32 597374 Near Djebel Oust Road ditch Discoglossus pictus New datum F142 4015008 32 572251 Near Pont du Fahs Pool Discoglossus pictus New datum F144 3968066 32 547684 Dechret Jenoua Stream Discoglossus pictus New datum F146 3962267 32 572002 ‘Ayn Jalulah Stream Discoglossus pictus

Bufo mauritanicus New data

F147 3956443 32 584977 Near Kairouan Temporary swamp Discoglossus pictus Bufo boulengeri

New data

F148 3952987 32 588672 Near Kairouan Pool Bufo boulengeri New datum F150 3949673 32 596589 Near Kairouan Road ditch Discoglossus pictus New datum F151 3933755 32 594284 Oued Zeroud, near

Fadden al Baqar Pools in the bed of a dry stream


F152 3942575 32 585613 Near Kairouan Road ditch Discoglossus pictus Bufo boulengeri

New data

F153 3944067 32 564884 Near Haffur Reservoir Discoglossus pictus New datum F155 3971781 32 519481 Near Maktar Pool Discoglossus pictus New datum F156 3975590 32 517953 Near Maktar Pool Discoglossus pictus New datum

Appendix I (continued)List of localities and their amphibian fauna. Latitude and longitude are given in Universal Transverse of Mercator system (UTM, map datum: WGS84).


F158 4036497 32 480314 Near Jendouba Road ditch Discoglossus pictus New datum F159 4052275 32 471884 Oued Ghrib, near Al

Firnanah Stream and nearby pools

Pelophylax saharicus New datum

F160 4055097 32 472917 Near Al Firnanah Flooded field Pleurodeles nebulosus Pelophylax saharicus

New data

F161 4060690 32 471024 Near Al Firnanah Permanent pool Pleurodeles nebulosus Discoglossus pictus Pelophylax saharicus

New data

F162 4060916 32 470964 Near Al Firnanah Temporary pond Discoglossus pictus New datum F163 4061859 32 471145 Near Al Firnanah Spring and pools Pleurodeles nebulosus

Discoglossus pictus Hyla meridionalis Pelophylax saharicus

New data

F164 4066450 32 471351 Near Aïn Draham Pools Discoglossus pictus New datum F165 4064410 32 473435 Near Beni M’tir Oak wood, near a

reservoirBufo bufo New datum

F166 4064634 32 473698 Near Beni M’tir Temporary pool Discoglossus pictus New datum F167 4066608 32 474546 Near Beni M’tir Temporary pool Discoglossus pictus


F168 4083594 32 477339 Jaballah Temporary pond Pleurodeles nebulosus Hyla meridionalis

New data

F169 4086941 32 478252 Near Tabarka Temporary swamp Pleurodeles nebulosus Hyla meridionalis

New data

F170 4087261 32 478237 Tabarka Temporary swamp Pleurodeles nebulosus Hyla meridionalis

New data

F171 4087169 32 478184 Tabarka Temporary swamp Pleurodeles nebulosus Discoglossus pictus Hyla meridionalis

New data

F173 4090682 32 491422 Guelta el Essafra Marsh with flowing water

Discoglossus pictus Hyla meridionalis

New data

F175 4093076 32 507250 Near Djebel Abiod Pool Pleurodeles nebulosus Discoglossus pictus

New data

F176 4103542 32 512725 Near Sejenane Temporary swamp Discoglossus pictus Hyla meridionalis

New data

F177 4110210 32 523607 Near Dawwar Mraf Concrete reservoir Bufo mauritanicus New datum F178 4116486 32 529302 Near Sidi Ferdjani Temporary pool Discoglossus pictus New datum F179 4111379 32 532924 Near Dawwar Mraf Pool Discoglossus pictus


F180 4110341 32 533167 Near Dawwar Mraf Pool Discoglossus pictus New datum F181 4105312 32 538786 Near Dawwar Bechouk Pond Pleurodeles nebulosus Bufo

mauritanicusPelophylax saharicus

New data

F182 4103969 32 541082 Near Dawwar Bechouk Agricultural reservoir

Hyla meridionalis Pelophylax saharicus

New data

F183 4103264 32 542479 Near Dawwar Bechouk Country side Bufo mauritanicus New datum F184 4103725 32 551577 Near Dawwar Bechouk Road ditch Discoglossus pictus New datum F185 4101204 32 555456 Near Mateur Road ditch Discoglossus pictus New datum F186 4065257 32 657135 Near Ad Damus Country side Bufo mauritanicus New datum F191 4104062 32 602396 Near Qal’at Andalus Temporary swamp Bufo mauritanicus


F192 4099127 32 600974 Near Qal’at Andalus Flooded field Discoglossus pictus New datum F193 4097541 32 601639 Near Qal’at Andalus Agricultural ditch Discoglossus pictus


F194 4089648 32 491678 Strada Meteur-Sejenane Pool Bufo mauritanicus Pelophylax saharicus

New data

F195 4056635 32 568592 Near Montarnaud Saltmarsh Bufo boulengeri New datum F196 4061233 32 537252 Near Wadi az Zarqa’ Road ditch Discoglossus pictus


F197 4059282 32 473914 Near Al Firnanah Reservoir Pelophylax saharicus New datum F198 4045789 32 478096 Bulla Regia Road ditch Pleurodeles nebulosus




F199 4092786 32 601370 Bou Hnach Temporary swamp Pleurodeles nebulosus Pelophylax saharicus

New data

FA199 4052244 32 498551 Bou Salem Road ditch Pleurodeles nebulosus Bufo boulengeri

New data

F200 4092833 32 601510 Bou Hnach Temporary swamp Pleurodeles nebulosus Pelophylax saharicus

New data

F206 3933530 32 471396 Thala Agricultural reservoir

Bufo mauritanicus Bufo boulengeri Pelophylax saharicus

New data

F209 4118378 32 519854 Cap Serrat, near Sidi Ferdjani

Pools Discoglossus pictus Bufo mauritanicus Pelophylax saharicus

New data

F210 4108286 32 559797 Ichkeul, near Mateur Pools Pelophylax saharicus New datum F211 3707101 32 595930 Matmata City Bufo boulengeri New datum F212 4003661 32 474093 El Kef Concrete reservoir

and City garden Discoglossus pictus Bufo boulengeri Pelophylax saharicus

New data

F 213 3752703 32 420042 Tozeur Concrete reservoir Pelophylax saharicus New datum F214 3643762 32 634483 Oued near Tataouine Pools in the bed of

a dry stream Bufo boulengeri New datum

F215 3753235 32 420328 Tozeur Irrigation canals Pelophylax saharicus New datum F217 3881372 32 526588 Near Sidi Bu Zayd Temporary pool Discoglossus pictus

Bufo mauritanicus Pelophylax saharicus

New data

F218 3935669 32 569247 Near Funduq al ‘Uqbi Temporary pools Pelophylax saharicus New datum F220 4084287 32 599275 El Bokkri Permanent stream Discoglossus pictus

Bufo boulengeri Pelophylax saharicus

New data

F221 3986200 32 618801 Oued Amer, near Burj al Bay

Temporary swamp Discoglossus pictus Bufo boulengeri

New data

F222 3962281 32 603710 Metbassta Temporary swamp Discoglossus pictus New datum F224 4110191 32 533330 Near Dawwar Mraf Temporary

agricultural channel

Discoglossus pictus New datum

F225 4085700 32 664867 Enchaa Road ditch Pleurodeles nebulosus New datum F228 4092577 32 607587 Raoued Temporary swamp Discoglossus pictus New datum F230 4120702 32 574024 Menzel Abderahmen Flooded field Discoglossus pictus New datum F231 4107339 32 562000 Parco dell'Ichkeul Temporary pool Pleurodeles nebulosus New datum F232 4117865 32 546215 Teskraia Temporary pools Pleurodeles nebulosus New datum F233 4110256 32 533377 Barrage Sejenane Road ditch Discoglossus pictus New datum F234 4110331 32 530076 Barrage Sejenane Temporary swamp Pleurodeles nebulosus


F235 4091234 32 486307 Tabarka aeroporto Temporary swamp Pleurodeles nebulosus Discoglossus pictus

New data

F236 4121055 32 575599 vicino Bizerte Temporary pond Pleurodeles nebulosus Discoglossus pictus

New data

F239 4078730 32 664615 Sidi Hassoun Flooded field Pleurodeles nebulosus New datum F240 4079501 32 664854 Garaa El Khala Temporary swamp Pleurodeles nebulosus

Discoglossus pictus Pelophylax saharicus

New data

F241 3951967 32 600087 Near Kairouan Pool in the bed of a dry stream

Discoglossus pictus New datum

F243 4109734 32 548959 Sidi Salem Road ditch Pleurodeles nebulosus New datum

F244 4116360 32 551370 Teskraia Temporary pool Pleurodeles nebulosus Discoglossus pictus

New data

FE 3998826 32 471934 Near El Kef Stream Discoglossus pictus Bufo mauritanicus Pelophylax saharicus

New data

PEK 4002596 32 474172 El Kef Country side Bufo boulengeri New datum Map 4094896 32 507253 Near Djebel Abiod Reservoir Bufo mauritanicus New datum Tam 3805606 32 403115 Tamerza Stream Pelophylax saharicus New datum



Rob2 4103008 32 677723 El Haouaria Swimming pool Discoglossus pictus New datum Rob6 4070369 32 475639 Near Ain Draham Puddle Discoglossus pictus New datum Rob20 4093386 32 497063 Sidi Berrak barrage Small stream on

sandy soil Discoglossus pictus Pelophylax saharicus

New data

Rob29 4112745 32 524052 Near Sidi Ferdjani Temporary stream with pools

Bufo mauritanicus Pelophylax saharicus

New data

Rob33 4117762 32 520914 Near Sidi Ferdjani Pool along a small stream in a pasture

Discoglossus pictus Bufo mauritanicus Pelophylax saharicus

New data

Rob54 3807914 32 410493 Midès Small pools and temporary stream in a canyon

Bufo boulengeri Pelophylax saharicus

New data

Rob66 3691042 32 470712 El Faouar Oasis Bufo boulengeri New datum Rob74 3706888 32 596206 Matmata Temporary stream

with pools Bufo boulengeri Pelophylax saharicus

New data

- - - Nefta - Discoglossus pictus Boulenger, 1891 - - - Tozeur - Discoglossus pictus Boulenger, 1891 - - - La Galite - Discoglossus pictus Boulenger,

1891;Lanza and Buzzone, 1959; Zangari, Cimmaruta and Nascetti, 2006

- - - Nean Tunis* - Bufo mauritanicus Boulenger, 1891 - - - Tozeur - Bufo mauritanicus Boulenger,

1891;Mosauer, 1934; Schneider, 1978

- - - Gafsa - Bufo mauritanicus Boulenger, 1891;Mosauer, 1934

- - - Hammam El Lif - Pleurodeles nebulosus Mayet, 1903 Staz. 207 - - km 109 between Mateur

and Tabarka* Swamp (garaa) Pleurodeles nebulosus Gauthier, 1928

Staz. 211 - - Near Oued Tindja Swamp (garaa) Hyla meridionalis Pelophylax saharicus

Gauthier, 1928

Staz. 215 - - km 8.35 between Sidi=Athman and Protville

Swamp (garaa) Pleurodeles nebulosus Discoglossus pictus

Gauthier, 1928

Staz. 217 - - km 10.9 between Sidi=Athman and Protville

Swamp (garaa) Discoglossus pictus Pelophylax saharicus

Gauthier, 1928

Staz. 223 - - 1.500 m from Kairouan to the roat to Enfida

Marsh (R’dir) Bufo boulengeri Gauthier, 1928

Staz 224 - - 34 km 5 from Sousse to the road to Kairouan


Staz. 228 - - km 243,4 between sfax and El-Djem


- - - Tozeur Irrigation channel Discoglossus pictus Gauthier, 1928; Gallien, 1948

- - - Ischkeul - Pelophylax saharicus Gauthier, 1928; Lymberakis et al., 2007

1 - - Belvedere-Park bei Tunis

City park Discoglossus pictus Bufo boulengeri Pelophylax saharicus

Mertens, 1929

2 - - Karthago - Bufo mauritanicus Mertens, 1929; Mosauer, 1934

2 - - Karthago - Bufo boulengeri Mertens, 1929 3 - - Gabés Oasis Bufo boulengeri Mertens, 1929



3 - - Gabés Irrigation canals of the oasis

Pelophylax saharicus Mertens, 1929

4 - - Houmt Souk, Jerba Island

Irrigation canals Bufo boulengeri Mertens, 1929

4 - - Meboubine, Jerba Island Ponds Bufo boulengeri Mertens, 1929 - - - Near Maxula Rades Puddles and

irrigation ditches Discoglossus pictus Mosauer, 1934

- - - Near Maxula Rades - Bufo mauritanicus Bufo boulengeri Pelophylax saharicus

Mosauer, 1934

- - - Gafsa Oasis Pelophylax saharicus Mosauer, 1934 - - - Near Sidi Bou Ali - Bufo boulengeri Mosauer, 1934- - - Between Kairouan and

Ain Ghrasesia - Pelophylax saharicus Mosauer, 1934

- - - Gabés Oasis Pelophylax saharicus Mosauer, 1934 - - - Tozeur Oasis Pelophylax saharicus Mosauer, 1934 - - - Grombalia - Pleurodeles nebulosus Blank, 1935 - - - Oued Medjerda, near

Medjez El Bab - Pleurodeles nebulosus Blank, 1935

- - - Oued Miliane, St. Germain

- Pleurodeles nebulosus Blank, 1935

- - - Zaonia de Melassina - Pleurodeles nebulosus Discoglossus pictus Hyla meridionalis

Blank, 1935

- - - Djbel Raças* - Discoglossus pictus Blank, 1935 - - - St. Germain - Discoglossus pictus

Hyla meridionalis Blank, 1935

- - - Near Tunis* - Hyla meridionalis Blank, 1935 - - - Bardò garden, Tunis - Hyla meridionalis Blank, 1935 - - - Cherichera* - Bufo mauritanicus Schneider, 1978 - - - Bou Hedma - Discoglossus pictus

Bufo mauritanicus Bufo boulengeri Pelophylax saharicus

Blank and Shane, 1981

- - - El Kantara (Kerkennah) - Bufo boulengeri Blank andNouira, 1988

- - - Sidi Fredj (Kerkennah) - Bufo boulengeri Blank andNouira, 1988

- - - Qued Nabtour, next to the road linking Bizerte and Tunis*

- Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Qued Jourmine, on the outskirts of Mateur

Stream Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Qued near the village of Ghezala


- - - Near the village of Nefza, in the Qued Melah


- - - On the outskirts of Tabarka, near the broad mouth of the Qued el Kebir


- - - Ben Metir reservoir: in small ponds below the side of the dam away from the reservoir

Ponds near the reservoir

Pelophylax saharicus Steinwarz and Schneider, 1991

- - - At two sites along the road between Tabarka and Jendouba, near Fernaria

Small, slowly flowing streams




- - - In the immediate surroudings of the city El Kef and along the road between El Kef and Sakiet Sidi Youssef

- Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Barrage Mellegue, near the village of Nebeur

Several ponds near a reservoir


- - - On the outskirts of Haffouz

Large pool created as a watering place


- - - On the southern outskirts of Sfax

Small stream Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Oasis of Gafsa Large water reservoir


- - - Tozeur around the oasis garden "Paradise"

Irrigation canals Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Nefta Irrigation canals Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Bechri Man-madewatering place


- - - El Hamma Distribution basin Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Qued El Hamma* Stream Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Qued Gabès Stream Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Oasis of Kettana (Qued el Ferch)


- - - Oasis of Teboulbou Small stream Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Qued Smar (Medenine), under the Medenine-Foum Tatahouine road, and near the underpass

Swamp in the bed of a stream


- - - Two Km southern the previous site

Small stream Pelophylax saharicus Steinwarz and Schneider, 1991

- - - Radès Permanent pond Pelophylax saharicus Meddeb and Cheniti, 1998; Meddeb et al., 2007

- - - Menzel Bourguiba Temporary pond Pelophylax saharicus Meddeb and Cheniti, 1998; Meddeb et al., 2007

- 4040001 32 601199 Surroundings of Tunis - Pelophylax saharicus Plötner, 1998 - 4006267 32 548546 El Fahs - Pelophylax saharicus Plötner, 1998 - - - Port Prince - Forêt et

Oued El Abid Stream and marsh Discoglossus pictus Nouira, 2001

- - - Port Prince - Bufo boulengeri Nouira, 2001 - - - Port Prince - Forêt et

Oued El Abid Stream Pelophylax saharicus Nouira, 2001

- - - Lagune de Korba (Cap Bon)

- Bufo boulengeri Nouira, 2001

- - - Lagune de Korba (Cap Bon)

- Pelophylax saharicus Nouira, 2001

- - - Dar Chichou (Cap Bon) - Pelophylax saharicus Nouira, 2001 - 3678590 32 616456 Medenine - Pelophylax saharicus Plötner and

Ohst, 2001 A1 4055164 32 473033 South of Fernana - Pleurodeles nebulosus Pasmans et al.,

2002A2 4038670 32 506269 Near Jendouba - Pleurodeles nebulosus Pasmans et al.,

2002



A3 4058676 32 473342 East of Fernana - Pleurodeles nebulosus Pasmans et al., 2002

B1 4086950 32 478180 South of Tabarka - Pleurodeles nebulosus Pasmans et al., 2002

B2 4090990 32 501187 West of Nefza - Pleurodeles nebulosus Pasmans et al., 2002

B3 4095802 32 509341 East of Nefza - Pleurodeles nebulosus Pasmans et al., 2002

B4 4089159 32 482193 East of Tabarka - Pleurodeles nebulosus Pasmans et al., 2002

C 4065148 32 473214 Forêt des Chênes, 8 km south of Aïn Draham

- Pleurodeles nebulosus Pasmans et al., 2002

RS7 - - Bou Ghanem* - Pelophylax saharicus Harris, Batista and Carretero, 2003

RS8 - - Kesra - Pelophylax saharicus Harris, Batista and Carretero, 2003

22 4090246 32 477751 Tabarka - Pleurodeles nebulosus Carranza and Arnold, 2004

25 4106839 32 508153 5 Km West of Nefza - Pleurodeles nebulosus Carranza and Arnold, 2004

26 4084671 32 477686 5 km south of Tabarka - Pleurodeles nebulosus Carranza and Arnold, 2004

35 4038817 32 479293 Souk-el Arba (Jendouba)

- Pleurodeles nebulosus Carranza and Arnold, 2004

36 4055097 32 472917 Near Al Firnanah - Pleurodeles nebulosus Carranza and Arnold, 2004

37 4058725 32 478362 Dam Bou Heurtma - Pleurodeles nebulosus Carranza and Arnold, 2004

39 4062654 32 471972 Ain Draham - Pleurodeles nebulosus Carranza and Arnold, 2004

Tun2 - - Bou Chebka - Bufo boulengeri Batista et al., 2006

Tun3 - - Jbel Bour* - Bufo boulengeri Batista et al., 2006

Tun5 - - Dam Bou* - Bufo boulengeri Batista et al., 2006

Tun6 - - Ain Draham - Bufo boulengeri Batista et al., 2006

3737246 32 616022 Oued El Ferd, near Gabès

Stream Discoglossus pictus Busack, 2006

- 3737246 32 616022 Oued El Ferd, near Gabès

Stream Pelophylax saharicus Busack, 2006

- - - Nefta - Bufo boulengeri Stoeck et al., 2006

- - - Jerba - Bufo boulengeri Stoeck et al., 2006

- 3806496 32 481234 Gafsa Oasis - Discoglossus pictus Amor et al., 2007

- 3948177 32 599123 Kairouan - Discoglossus pictus Amor et al., 2007

- 3891601 32 484341 Kasrine - Discoglossus pictus Amor et al., 2007

- 4058383 32 478975 Beja - Discoglossus pictus Amor et al., 2007

- 3806496 32 481234 Gafsa - Bufo mauritanicus Amor et al., 2007

- 3806583 32 484170 Oued El Maleh Stream Bufo mauritanicus Amor et al., 2007



- 4026209 32 637973 Hammamet - Bufo mauritanicus Amor et al., 2007

- 3802357 32 488199 Orbata - Bufo boulengeri Amor et al., 2007

- 3891601 32 484341 Kassrine - Bufo boulengeri Amor et al., 2007

- 3806496 32 481234 Gafsa Oasis - Pelophylax saharicus Amor et al., 2007

- 3805168 32 483478 Ain Soltane - Pelophylax saharicus Amor et al., 2007

- 3806583 32 484170 Oued El Maleh Stream Pelophylax saharicus Amor et al., 2007

- 3805233 32 481359 Oued Bayash Stream Pelophylax saharicus Amor et al., 2007

- 3753754 32 419068 Ras El Aïn - Pelophylax saharicus Amor et al., 2007

- - - Sahline - Pelophylax saharicus Amor et al., 2007

- 4081380 32 605317 El Menzeh - Pelophylax saharicus Amor et al., 2007

- 3948177 32 599123 Kairouan - Pelophylax saharicus Amor et al., 2007

- - - 4 km before Sedjenan - Pelophylax saharicus Lymberakis et al., 2007

- - - Bizerte - Pelophylax saharicus Lymberakis et al., 2007

35 4072556 32 677353 Lebna, Nabeul - Hyla meridionalis Recuero et al., 2007

36 4089846 32 478459 Tabarka, Jendouba - Hyla meridionalis Recuero et al., 2007

* Sites not included in the distribution maps because of the lack of clear geographic information


Accepted by Angelica Crottini; Managing Editor: Philip de Pous

Contribution to the knowledge of Tunisian amphibians - Herpetology

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