Pelophylax bedriagae (Camerano, 1882)
Levante water frog, Levant waterfrog, Levant Green Frog, Bedriaga's Frog, Jaza' Akhdar, Türkischer Wasserfrosch
© 2015 Martin Pickersgill (1 of 11)
There are two distinct, longitudinal dorsolateral ridges (Baha El Din 2006) on their otherwise smooth to slightly rough dorsal surface. The ventrum is smooth (Escoriza and Ben Hassine 2019).
The fingers are unwebbed, pointed, do not expand into discs, but have prominent subarticular tubercles at the ends (Disi and Amr 2010, Escoriza and Ben Hassine 2019). Their relative finger lengths are II ≈ I < IV < III. During the reproductive season males develop nuptial pads at the base of their first finger (Escoriza and Ben Hassine 2019). The well developed hind limbs are long, with extensive webbing between the toes reaching to about the third phalange on toes I, II, III, and IV and to the base of the third phalange of toe IV (Baha El Din 2006, Disi and Amr 2010). The inner metatarsal tubercle is elongated. The toes are moderately webbed with a formula of I 0.5 - 1.5 II 0.5 - 2 III 1 - 2 IV 2 - 0.5 V and a relative toe length formula of I < II < V < III < IV. Like the fingers, the toes do not have discs (Escoriza and Ben Hassine 2019).
At Gosner stage 36, P. bedriagae tadpoles from south-west Turkey have a total length between 50 – 70 mm, a body length of approximately 20.5 mm, and tail length of approximately 44.4 mm. Larval P. bedriagae have a slightly depressed benthic body form with a rounded snout in both the dorsal and lateral views. The nostrils are closer to the tip of the snout than the eyes and directed anterolaterally. The small eyes are directed dorsolaterally. The oral disc is anteroventral with lateral emargination. The labial tooth row formula is 2(2)/3, with all rows being uniserial. The medium sized jaw sheaths are keratinized with blunt serrations. The spiracle is sinistral and the vent is dextral. The tail is approximately 2 times the body length and ends in a broadly pointed tipped. The dorsal fin is low and extends onto the posterior body. The tail muscular height is approximately 5.9 mm and the maximal tail height is 12.2 mm (Escoriza and Ben Hassine 2019).
Newly metamorphosed frogs have a snout-vent length of about 26 mm (Escoriza and Ben Hassine 2019).
Morphologically, P. bedriagae is difficult to distinguish from other eastern Mediterranean water frogs, which includes P. caralitanus, P. cretensis, P. cypriensis, P. epeiroticus, and P. ridibundus. The species are mainly distinguished by advertisement call, (Schneider and Sinsch 1999) and/or geography (Plötner et al. 2010), however, P. bedriagae has also been introduced to many sites in Europe (Holsbeek et al. 2008), which hampers the ability to properly identify those populations. Pelophylax bedriagae can be differentiated from P. ridibunda, from which it was split, based on call, with P. bedriagae having fewer pulses per pulse group among other features (Schneider and Sinsch 1999), and to a lesser extent by the former having a larger size than the latter. Pelophylax caralitanus can be differentiated from P. bedriagae by the focal species having a smaller snout-vent length and lacking an orange ventrum (Sinsch et al. 2002). Pelophylax cypriensis can be differentiated from P. bedriagae by the focal species having a more narrow distance between nostrils and/or a smaller metatarsal tubercle and a narrower head. Furthermore, the mating call of P. cypriensis has fewer pulse groups per call and a generally shorter pulse group duration of 26 - 27 ms at 20oC (Plötner et al. 2012)
In life, the dorsum of P. bedriagae is typically greenish or brownish, sometimes with a narrow, light-yellowish or green mid-dorsal stripe that extends from the tip of the snout to the urostyle. Large orange, light brown, or dark spots are scattered over the body. The limbs have dark crossbars on the dorsal surface and the inner part of the thighs have black marbling on white or brown background. The ventral sides appear whitish and may be marbled with grey or black. The ventrum is dirty white. The vocal sac in males is darker (dark grey or black) than the rest of the body. The eyes are orange-gold with black reticulations and darker lower two-thirds (Baha El Din 2006, Disi and Amr 2010, Escoriza and Ben Hassine 2019).
In live larval P. bedriagae, the dorsal and lateral surfaces of the body are uniformly yellowish, greenish, or grayish brown. The belly is white. The tail musculature is speckled with black and white. The fin is transparent in most specimens, but can be spotted with gray or black. The eyes are white-yellow (Escoriza and Ben Hassine 2019).
There is sexual dimorphism in size with females being larger. Males in Jordon have gray or blackish-brown tympanum, while females have lighter tympanum (Disi and Amr 2010). Snout-vent range and advertisement call vary slightly by geography (Escoriza and Ben Hassine 2019). Populations near Ulubey, Turkey have been reported to have larger tympanum than other populations (Sinsch et al. 2002). There is variation in color patterning, with some specimens having a mid-dorsal line, variable number and size of spots on the abdomen, and coloration on the abdomen (Disi and Amr 2010).
Distribution and Habitat
Country distribution from AmphibiaWeb's database: Egypt, Iran, Islamic Republic of, Israel, Jordan, Lebanon, Syrian Arab Republic, Turkey. Introduced: Malta.
Pelophylax bedriagae is widespread in the Eastern Mediterranean. In Turkey, it is commonly found along the Aegean and the Mediterranean coast and in the southern part of the Anatolian highlands. Large numbers can be found in western Syria, with some isolated populations in the north. It is common throughout Lebanon, northeastern Jordan, much of Israel (excluding the extreme south), and Egypt (in the Nile Delta and Valley, vicinity of Cairo, Fayoum basin, on the shore of the Suez Canal, and in northeastern Sinai). The species has been confirmed in western Iran and unconfirmed sightings have been reported from Iraq. In Jordan, the species is found from sea level to 1,500 m asl (Baha El Din 2006, Papenfuss et al. 2016, Pesarakloo et al. 2017, Escoriza and Ben Hassine 2019, H. Schneider per. comm. 2020). The species has also been recorded in Bulgaria around the Vurbitza River (Lukanov et al. 2018).
The species has been introduced to several European sites. In 2000, the first sightings of the species were made on Gozo Island, Malta (Sciberras and Schembri 2006, Escoriza and Ben Hassine 2019). The species has been confirmed in three separate Maltese sites (Sciberras and Schembri 2006). In 2008, the species’ mitochondrial DNA was identified in Belgium indicating an introduction from the Anatolian region (Holsbeek et al. 2008). Anatolian P. bedriagae genes have also been identified in western Switzerland (Dubey et al. 2014). In 2019, the species was reported to have been introduced to northern Sardinia. The species is also known to have been introduced to Cyprus where it is hybridizing with native Pelophylax cypriensis (Bellati et al. 2019).
Pelophylax bedriagae is generally found in swampy habitats like permanent wetlands or aquatic vegetation. It is common in ponds, rain pools, streams, rivers, irrigation channels, reservoirs, marshes, springs and fishponds. It can occasionally also be found in surrounding terrestrial habitats (Baha El Din 2006, Escoriza and Ben Hassine 2019).
Life History, Abundance, Activity, and Special Behaviors
In Egypt, the species can be found year-round (Escoriza and Ben Hassine 2019).
Pelophylax bedriagae breeds throughout early spring to late summer (Baha El Din 2006). In Egypt, the breeding season occurs between late February or March to September (Akef and Schneider 1989, Escoriza and Ben Hassine 2019) while in the Jordan Valley it is between January and February and in the Jordanian highlands it starts between March and May (Disi and Amr 2010, Escoriza and Ben Hassine 2019). Breeding generally occurs in shallow, sheltered water bodies (Baha El Din 2006) with axillary amplexus (Escoriza and Ben Hassine 2019). Larger males appear to breed earlier and smaller males later in the breeding season (Akef and Schneider 1989). Breeding appears dependent on temperature and not dependent on rain as the species lives in permanent water bodies (Disi and Amr 2010).
Pelophylax bedriagae have three types of vocalizations: advertisement, territorial, and transitional calls (H. Schneider per. comm.). Vocalizations occur in the water (Escoriza and Ben Hassine 2019). The advertisement call consists of a series of 30 calls separated by intercall intervals that start with shorter calls and lengthen in duration until the middle of the series. The longer call lengths are maintained until the end of the call or just shortly before the end (H. Schneider pers. comm.). Individual calls have a duration between 530 – 943 ms, with 5.89 – 10.50 pulse groups/call, 12.36 pulses/group, and a dominant frequency range of 2000 – 3000 Hz (Baha El Din 2006, Escoriza and Ben Hassine 2019, H. Schneider per. comm.). The pulse groups begin at a low amplitude and gradually increases until shortly before the end or at the end (H. Schneider pers. comm.). Call characteristics differ by population, geography, and ambient temperature (Schneider and Sinsch 1999). Pulse group duration, interval between pulse groups, and period of pulse groups decrease with increasing water temperature, whereas the number of pulse groups per second increases. Call duration, intercall interval, call period and number of pulse groups per call are temperature independent (H. Schneider pers. comm.).
At water temperatures of 20oC, the advertisement calls of male Pelophylax bedriagae at the type locality have the following mean values: call duration 700.7 ms, intercall interval 1017.0 ms, call period 1722.0 ms, number of pulse groups per call 10.3, pulse group duration 45.5 ms, interval between pulse groups 22.23 ms, period of pulse groups 67.8 ms, pulse groups per second 16.0 Hertz. The frequency spectrum of the advertisement calls is broad. It shows a low-frequency component at 450 Hertz, the dominant frequency is between 2 and 3.2 Kilohertz, and another energy component between 4.5 and 5.2 Kilohertz (H. Schneider pers. comm.).
To hear the call, go to the Tree of Life webpage for Pelophylax bedriagae.
In Egypt, females lay up to 2344 eggs. Individual eggs are 1.3 – 1.8 mm in diameter and are laid in a transparent gelatinous sheath (Escoriza and Ben Hassine 2019). At one site in Israel, tadpoles were found from June to August (Degani 1982). The species takes 2 – 3 years to reach sexual maturity with males having a lifespan of four years and females having a lifespan of five years (Escoriza and Ben Hassine 2019).
Like most frogs, adult P. bedriagae have a generalist diet, but most commonly consume Hymenoptera and Coleoptera. A study in Iran found the diet is composed of 75% terrestrial species and 25% of aquatic species. Ecological factors, habitat, and food abundance also play roles in the diet of the species (Bam-e-Zar et al. 2019). However, there have also been reports of P. bedriagae adults consuming terrestrial vertebrates, such as lizards and birds (Nicolaou et al. 2014).
Larval P. bedriagae feed on plants, algae, and protozoa (Degani 1982, 1986).
Adult P. bedriagae from Izmir and Manisa Provinces, Turkey were found to host eight species of helminth parasites including: Rhabdias bufonis, Cosmocerca ornata, Oswaldocruzia filiformis (members of Nematoda), Diplodiscus subclavatus, Haematoloechus breviansa, Gorgoderina vitelliloba (Digenea), Acanthocephalus ranae (Acanthocephala), and Hirudo medicinalis (Hirudinea) (Demir et al. 2015).
Almost a third of larval P. bedriagae in Karpuzcay Creek, Antalya, Turkey suffered from Lernaea cyprinacea parasitism (Innal et al. 2019).
Trends and Threats
In Turkey and Egypt, the species is collected and exported for human consumption in Europe (Papenfuss et al. 2016). It is unknown how this harvest affects the population (Escoriza and Ben Hassine 2019).
In the western Aegean Region of Turkey, the species is also threatened by the introduction of the Wels Catfish (Siluris glanis), and crayfish of the species Astacus leptodactylus (Baskale and Kaya 2012).
The species is protected in many countries. In Israel, the species is protected by legislation. In Jordan, it is found in several protected areas, such as the Azraq Wetland Reserve, the Dana Wildlife Reserve, and the Al Maujib Wildlife Reserve. In Lebanon, there are no known threats to the species and it occurs within several protected areas, such as Arz El-Shouf, Horj Ehden, the Ammiq marshes and the Sandy Beach of Sour (Baha El Din 2006, Papenfuss et al. 2016). It can also be found in several protected areas in Turkey (Papenfuss et al. 2016).
The species has been introduced to several European sites where it may be affecting local species through competition or hybridization. In Malta, the effects of P. bedriagae on the only native amphibian species, Discoglossus pictus, are unknown (Sciberras and Schembri 2006). In Cyprus, P. bedriagae is reported to hybridize with the native amphibian, Pelophylax cypriensis (Bellati et al. 2019).
Göçmen et al. (2013) found one B. dendrobatidis positive juvenile individual in Turkey but did not elaborate on the health of the individual.
Relation to Humans
Possible reasons for amphibian decline
General habitat alteration and loss
Pelophylax bedriagae was split from P. ridibundus by Schneider and Sinsch (1992), but its relationship and placement with other closely related Pelophylax is unresolved as multiple molecular analyses indicate that P. bedriagae may be composed of several cryptic species (Plötner et al. 2002, Lymberakis et al. 2007, Plötner et al. 2012, Toli et al. 2018). Based on phylogenetic analyses (Bayesian Inference, Maximum Parsimony, Maximum Likelihood, and Neighbor joining) of Cyt B and 16S rRNA, a clade composed of P. bedriagae (from Syria and Turkey), P. cerigensis and P. cypriensis is sister to the clade composed of P. kurtmuelleri and R. ridibunda. The Syrian samples of P. bedriagae were sister to P. cypriensis while Turkish samples of P. bedriagae were sister to Greek Pelophylax from Rhoades Island (Lymberakis et al. 2007). However, while also supporting the relationship of Syrian P. bedriagae and P. cypriensis, analysis of CytB and AFLPs by Toli et al. (2018) found that the Rhoades Island Pelohylax were more closely related to Pelophylax from Karpathos. Based on Maximum Likelihood analyses of two mitochondrial genes (ND2 and and ND3) and the nuclear intron SAI-1, the close relationship between P. bedriagae and P. cypriensis was supported in Plötner et al. (2012) with the caveats of P. cretensis being sister to P. cypriensis and the clade composed of P. cretensis and P. cypriensis being sister to P. bedriagae from Jordan rather than Syria. Plötner et al. (2012) also found P. bedriagae to be polyphyletic.
The high genetic but low morphological diversity of the eastern Mediterranean Pelophylax is attributed to Messinian salinity crisis (c. 5.5 – 5.3 Ma) followed by isolation events during the Pleistocene (c. 2.85 – 0.012 Ma) and climatic changes during the Pliocene (Akin et al. 2010, Toli et al 2018).
The genus, “Pelophylax” is derived from the Greek words “pēlos”, meaning “mud” and “phulax”, meaning “sentinel” (Escoriza and Ben Hassine 2019).
The species epithet, “bedriagae” is in honor of Jacques Vladimir von Bedriaga, a Russian herpetologist (Escoriza and Ben Hassine 2019).
Populations in Israel were formerly considered to be Rana ridibunda. These populations were also briefly classified as a new species, Rana levantina, based on bioacoustics data (Schneider and Sinsch 1992). However, Dubois suggested the new species name was a junior synonym of P. bedriagae, which was later confirmed by bioacoustics analyses (Schneider and Sinsch 1999).
Archeological studies from the Early-Middle Pleistocene (0.8 - 0.7 Ma) in northern Israel recovered a large number of P. bedriagae fossils along with Hyla cf. savignyi, Discoglossus (cf.) nigriventer, and Bufo cf. viridis. Holocene sites in the Lower Jordan Valley at Gilgal I and Netiv Hagdud also recovered P. bedriagae fossils (Rabinovich and Biton 2011).
Known, as voracious predators, there is a report of an adult P. bedriagae choking to death while attempting to eat a bird that was over half of its size (Nicolaou et al. 2014).
Akef, M. S. A., Schneider, H. (1989). ''The eastern form of Rana ridibunda(Anura: Ranidae) inhabits the Nile delta.'' Zoologischer Anzeiger, 223, 129-138.
Akin, C., Bilgin, C.C., Beerli, P., Westaway, R., Ohst, T., Litvinchuk, S.N., Uzzell, T., Bilgin, M., Hotz, H., Guex, G.-D., Plötner, J. (2010). ''Phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs were determined by geological processes and climate change in the Late Cenozoic.'' Journal of Biogeography, 37(11), 2111–2124 . [link]
Baha El Din, S. (2006). A Guide to the Reptiles and Amphibians of Egypt. The American University in Cairo Press, Cairo.
Bam-E-Zar, F., Fathinia, B., Shafaei-Pour, A. (2019). ''Trophology of Levant Green Frog, Pelophylax bedriagae (Amphibia: Anura: Ranidae) in Choram Township, Iran.'' North-Western Journal of Zoology, 15(2), 168-174. [link]
Baskale, E., Kaya, U. (2012). ''Decline of the Levantine Frog, Pelophylax bedriagae Camerano, 1882, in the western Aegean Region of Turkey: changes in population size and implications for conservation (Amphibia: Ranidae).'' Zoology in the Middle East, 57(1), 69-76. [link]
Bellati, A., Bassu, L., Nulchis, V., Corti, C. (2019). ''Detection of alien Pelophylax species in Sardinia (western Mediterranean, Italy).'' BioInvasions Records, 8(1), 8–25. [link]
Degani, G. (1982). ''Amphibian tadpole interaction in a winter pond.'' Hydrobiologia , 96, 3-7. [link]
Demir, S., Yakar, O., Yildirimhan, H.S., Birlik, S. (2015). ''Helminth parasites of the levantine frog (Pelophylax bedriagae Camerano, 1882) from the western part of Turkey.'' Helminthologia , 52(1), 71-76. [link]
Disi, A.M., Amr, Z.S. (2010). ''Morphometrics, distribution and ecology of the amphibians in Jordan.'' Vertebrate Zoology, 60(2), 147-162. [link]
Dubey, S., Leuenberger, J., Perrin, N. (2014). ''Multiple origins of invasive and ‘native’ water frogs (Pelophylax spp.) in Switzerland.'' Biological Journal of the Linnean Society, 112, 442–449. [link]
Escoriza, D., Ben Hassine, J. (2019). ''14 Bufotes boulengeri (Lataste, 1879) African green toad.'' Amphibians of North Africa. Academic Press , San Diego, CA, 159 - 168. [link]
Göçmen, B., Veith, M., Iğci, N., Akman, B., Godmann, O., Wagner, N. (2013). ''No detection of the amphibian pathogen Batrachochytrium dendrobatidis in terrestrial Turkish salamanders (Lyciasalamandra) despite its occurrence in syntopic frogs (Pelophylax bedriagae).'' Salamandra, 49(1), 51-55.
Holsbeek, G., Hotz, H., Plötner, J., Volckaert, F.A.M., De Meester, L. (2008). ''A cryptic invasion within an invasion and widespread introgression in the European water frog complex: consequences of uncontrolled commercial trade and weak international legislation.'' Molecular Ecology , 17, 5023–5035. [link]
Innal, D., Sozbilen, D., Ozmen, O. (2019). ''Prevalence and pathology of Lernaea cyprinacea (Crustacea: Lernaeidae) parasitizing tadpoles of Pelophylax bedriagae (Anura: Ranidae) in Antalya, Turkey.'' Phyllomedusa , 18(2), 177-183. [link]
Lukanov, S., Popgeorgiev, G., Tzankov, N. (2018). ''First bioacoustic and morphological data for the presence of Pelophylax bedriagae in Bulgaria.'' Acta Scientifica Naturalis, 5(1), 54-63. [link]
Lymberakis, P., Poulakakis, N., Manthalou, G., Tsigenopoulos, C.S., Magoulas, A., Mylonas, M. (2007). ''Mitochondrial phylogeography of Rana (Pelophylax) populations.'' Molecular Phylogenetics and Evolution, 44, 115-125. [link]
Nicolaou, H., Zogaris, S., Pafilis, P. (2014). ''Frog vs. lizard: an unusual feeding behavior in the Levantine Marsh Frog, Pelophylax bedriagae from Cyprus.'' North-Western Journal of Zoology , 10(1), 221-222. [link]
Papenfuss, T., Kuzmin, S., Disi, A.M., Degani, G., Ugurtas, I.H., Sparreboom, M., Anderson, S., Sadek, R., Hraoui-Bloquet, S., Gasith, A., Elron, E., Gafny, S., Lymberakis, P., Böhme, W., Baha El Din, S. 2009. ''Pelophylax bedriagae (errata version published in 2016)''. The IUCN Red List of Threatened Species 2009: e.T58559A86622844. https://dx.doi.org/10.2305/IUCN.UK.2009.RLTS.T58559A11803274.en. Downloaded on 05 February 2020. https://www.iucnredlist.org/species/58559/86622844
Pesarakloo, A., Rastegar-Pouyani, E., Rastegar-Pouyani, N., Kami, H., Najibzadeh, M., Khosravani, A. (2017). ''The first taxonomic revaluation of the Iranian water frogs of the genus Pelophylax (Anura: Ranidae) using sequences of the mitochondrial genome.'' Mitochondrial DNA PartA, 28, 392-398. [link]
Plötner J., Uzzell, T., Beerli, P., Akin, C., Bilgin, C.C., Haefeli, C., Ohst, T., Köhler, F., Schreiber, R., Guex, G.-D., Litvinchuk, S.N., Westaway, R., Reyer, H.-U., Pruvost, N., Hotz, H. (2010). ''Genetic divergence and evolution of reproductive isolation in eastern Mediterranean Water Frogs.'' Evolution in Action. Glaubrecht, M. , eds., Springer, Berlin, Heidelberg.
Plötner, J., Baier, F., Akin, C., Mazepa, G., Schreiber, R., Beerli, P., Litvinchuk, S.N., Bilgin, C.C., Borkin, L., Uzzell, T. (2012). ''Genetic data reveal that water frogs of Cyprus (genus Pelphylax) are an endemic species of Messinian origin.'' Zoosystematic and Evolution, 88(2), 261-283. [link]
Plötner, J., Ohst, T., Böhme, W., Schreiber, R. (2002). ''Divergence in mitochondrial DNA of near Eastern water frogs with special reference to the systematic status of Cypriote and Anatolian populations (Anura, Ranidae).'' Amphibia-Reptilia, 22(4), 397-412. [link]
Rabinovich, R., Biton, R. (2011). ''The early middle Pleistocene faunal assemblages of Gesher Benot Ya‘aqov: Inter-site variability.'' Journal of Human Evolution, 60(4), 357-374. [link]
Schneider, H., Sinsch, U. (1999). ''Taxonomic reassessment of Middle Eastern water frogs: Bioacoustic variation among populations considered as Rana ridibunda, R. bedriagae or R. levantina.'' Journal of Zoological Systematics and Evolutionary Research, 37(2), 57 - 66. [link]
Schneider, H., Sinsch, U. (1992). ''Mating call variation in lake frogs referred to as Rana ridibunda Pallas, 1771. Taxonomic implications.'' Journal of Zoological Systematics and Evolutionary Research, 30(4), 297–315. [link]
Sciberras, A, Schembri, P.J. (2006). ''Occurrence of the alien Bedriaga's frog (Rana bedriagae) Camerano, 1882 in the Maltese Islands, and implications for conservation.'' Herpetological Bulletin, 95, 2-5. [link]
Sinsch, U., Schneider, H., Kaya, U., Arikan, H. (2002). ''The water frogs (Anura, Ranidae) of Turkey: a morphometric view on systematics.'' Herpetological Journal, 12(4), 141-153. [link]
Toli, E.A., Siarabi, S., Bounas, A., Pafilis, P., Lymberakis, P., Sotiropoulos, K. (2018). ''New insights on the phylogenetic position and population genetic structure of the Critically Endangered Karpathos marsh frog Pelophylax cerigensis (Amphibia: Anura: Ranidae).'' Acta Herpetologica , 13(2), 117-123. [link]
Originally submitted by: Ann T. Chang (first posted 2009-03-19)
Edited by: Ann T. Chang (2020-03-29)
Species Account Citation: AmphibiaWeb 2020 Pelophylax bedriagae: Levante water frog <https://amphibiaweb.org/species/5078> University of California, Berkeley, CA, USA. Accessed Dec 2, 2022.
Feedback or comments about this page.
Citation: AmphibiaWeb. 2022. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 2 Dec 2022.
AmphibiaWeb's policy on data use.