Abstract
Syria is home to a wide range of ecosystems and landscapes that support a variety of animal species, including reptiles. Currently, Syria hosts nine species of chelonians (turtles and tortoises) from six different families, making its family-level diversity remarkable for the Mediterranean and the Middle East. We conducted a comprehensive review of existing literature, database information, and museum records, and incorporated new field data collected between 2019 and 2023 to document the distribution and conservation status of chelonians in Syria. As a result, we present an updated species list, accompanied by relevant information and occurrence maps for each turtle and tortoise species found in the country. Our investigation has yielded a total of 321 distribution records for Syrian chelonians (including nests of marine species), 105 of which are new. In addition, we verified 18 sites previously documented in the literature. Among the most frequently recorded species is Testudo graeca terrestris Forskål, 1775. In addition, for the first time, we have systematically evaluated the records of two marine turtle species that use Syrian coasts as nesting sites. This study is the most comprehensive and up-to-date source of information on the distribution and biogeography of chelonians in Syria, encompassing both historical and current geographic records. In addition, our data provide a zoogeographic overview and a substantial dataset of distribution information that can be used for subsequent analysis of macroecological patterns, distribution patterns, and conservation efforts related to this group of reptiles.
References
- Abdulsalam, A. (1990) Geographic Regions of Syria. Damascus University (Faculty of Literature), Damascus, 544 pp. [in Arabic]
- Abi-Said, M.R., Makhlouf, H., Amr, Z.S. & Eid, E. (2018) Illegal trade in wildlife species in Beirut, Lebanon. Vertebrate Zoology, 68, 1–4. https://doi.org/10.3897/vz.68.e32214
- Aidek, E.A. (2010) A Guide to the Biodiversity of Deir ez–Zor Area. Al-Quds House for Sciences, Damascus, 112 pp.
- Aidek, E.A., Ibrahim, A., Amr, Z., Hutterer, R., Kryštufek, B., Serra, G., Benda, P., Shkaky, Y., Barbanera, F. & Al-Sheikhly, F.O. (2024a) Checklist of the mammals of Syria (Chordata: Mammalia). Zootaxa. [in press]
- Aidek, E.A., Jablonski, D., Dufresnes, C.H., Salimeh, M. & Omran, Y. (2024b) The composition, distribution, and endangerment of the amphibian fauna of Syria. In: Heatwole, H. & Das, I. (Eds.), Conservation Biology of Amphibians of Asia. Amphibian Biology. Vol. 11 (6). Decline in the Middle East. Natural History Publications (Borneo) Sdn Bhd., Kota Kinabalu. [in press]
- Aidek, E.A., Omran, Y., Joger, U., Esterbauer, H., Abu Baker, M. & Amr, Z. (2023) Snakes of the Syrian Arab Republic. Edition Chimaira, Frankfurt, 118 pp.
- Amr, Z., Shehab, A. & Abu Baker, M. (2007) Recent observations on the herpetofauna of Syria with notes on trade in reptiles. Herpetozoa, 20, 21–26.
- Atallah, S.I. (1977) Mammals of the Eastern Mediterranean: Their ecology, systematics and zoogeographical relationships. Säugetierkundliche Mitteilungen, 25, 241–320.
- Ayaz, D., Bayrakcı, Y., Çiçek, K., Ihlow, F., Tok, C.V. & Fritz, U. (2021) On the brink of extinction: Results of a 20-year quest for Eiselt’s pond turtle (Emys orbicularis eiselti) in southeastern Turkey. Chelonian Conservation and Biology, 20, 222–230. https://doi.org/10.2744/CCB-1505.1
- Barkoudah, Y., Ali, M., Qadi, I., Nader, S., Haj-Mousa, F., Sanadiqi, N., Sadat, H. & Abd Alatif, K. (2002) Atlas of the Syrian Flora. Ministry of State for Environment Affairs, Damascus, 182 pp. [in Arabic]
- Baur, G. (1888) Dermochelys, Dermatochelys oder Sphargis. Zoologischer Anzeiger, 11, 44–45.
- Berger-Dell’mour, V.H. (1986) Zur Herpetofauna des Golan. Annalen des Naturhistorischen Museums in Wien, 87B, 59–67.
- Berthon, R., Erdal, Y.S., Mashkour, M. & Kozbe, G. (2016) Buried with turtles: The symbolic role of the Euphrates soft-shelled turtle (Rafetus euphraticus) in Mesopotamia. Antiquity, 90, 111–125. https://doi.org/10.15184/aqy.2015.196
- Bischoff, W., in den Bosch, H.A.J. & Schmidtler, J.F. (1998) Mit dem Finger auf der Landkarte entdeckt. Eidechsen auf dem Djebel Druz. Die Eidechse, 9, 1–9.
- Bischoff, W. & Schmidtler, J.F. (1994) Ergebnisse zweier Lacertiden-Exkursionen nach Syrien. Die Eidechse, 5, 4–22.
- Boettger, O. (1880) Die Reptilien und Amphibien von Syrien, Palaestina und Cypern. Berichte der Senckenbergischen Naturforschenden Gesellschaft, 1879–1880, 132–219.
- Böhme, M. & Ilg, A. (2003) fosFARbase. https://www.fosfarbase.org/ (accessed 17 May 2024)
- Bory de Saint-Vincent, J.B. (1836) Expédition Scientifique de Morée. Section des Sciences Physiques. Tome Premier. Relation. F.G. Levrault, Paris, 472 pp.
- Bottema, S. & Barkoudah, Y. (1979) Modern pollen precipitation in Syria and Lebanon and its relation to vegetation. Pollen et Spores, 21, 427–480.
- Boulenger, G.A. (1923) Etude sur les Batraciens et les Reptiles rapportes par M. Henri Gadeau de Kerville de son voyage zoologique en Syrie (avril-juin 1908). In: Pellegrin, J. (Ed.), Voyage zoologique d’Henri Gadeau de Kerville en Syrie (avril-juin 1908). Tome Quatrième et Dernier. J.-B. Baillière, Paris, pp. 41–55, pl. vi.
- Bour, R. (1989) Caractères diagnostiques offerts par le crâne des tortues terrestres du genre Testudo. Mésogée, 48, 13–19.
- Bour, R. & Dubois, A. (1984) Nomenclatural availability of Testudo coriacea Vandelli, 1761: A case against a rigid application of the rules to old, well-known zoological works. Journal of Herpetology, 17, 356–361. https://doi.org/10.2307/1563588
- Bour, R., Dubois, A. & Webb, R.G. (1995) Types of recent trionychid turtles in the Muséum national d’Histoire naturelle, Paris. Dumerilia, 2, 73–92.
- Çakırlar, C., Koolstra, F.J. & Ikram, S. (2021) Tracking turtles in the past: Zooarchaeological evidence for human-turtle interactions in the ancient Eastern Mediterranean. Antiquity, 95, 125–141. https://doi.org/10.15184/aqy.2020.95
- Casale, P., Broderick, A.C., Camiñas, J.A., Cardona, L., Carreras, C., Demetropoulos, A., Fuller, W.J., Godley, B.J., Hochscheid, S., Kaska, Y., Lazar, B., Margaritoulis, D., Panagopoulou, A., Rees, A.F., Tomás, J. & Türkozan, O. (2018) Mediterranean Sea turtles: Current knowledge and priorities for conservation and research. Endangered Species Research, 36, 229–267. https://doi.org/10.3354/esr00901
- Casale, P. & Mariani, P. (2014) The first ‘lost year’ of Mediterranean sea turtles: Dispersal patterns indicate subregional management units for conservation. Marine Ecology Progress Series, 498, 263–274. https://doi.org/10.3354/meps10640
- Dakdouk, S. (2009) Lebanon – back on the illegal wild animals trade map. Wildlife of the Middle East, 4, 2.
- Daudin, F.M. (1801) Histoire Naturelle, Générale et Particulière des Reptiles. Tome Second. F. Dufart, Paris, 432 pp.
- Disi, A.M. & Böhme, W. (1996) Zoogeography of the amphibians and reptiles of Syria with additional new records. Herpetozoa, 9, 63–70.
- Eid, E., Al Hasani, I., Al Share, T., Abed, O. & Amr, Z. (2010) Animal trade in Amman local Market, Jordan. Jordan Journal of Biological Science, 4, 101–108.
- Esterbauer, H. (1985a) Zur Herpetofauna Südwestsyriens. Herpetofauna, 7 (38), 23–34.
- Esterbauer, H. (1985b) Ökologische und verhaltensbiologische Beobachtungen an der Maurischen Landschildkröte, Testudo graeca terrestris Forskål, 1775 in Südwestsyrien. Aquarien-Terrarien, 32, 389–392.
- Esterbauer, H. (1992) Die Herpetofauna des östlichen Golan und Hermongebietes. Funde und Bemerkungen zur Systematik und Ökologie. Zoology in the Middle East, 7, 21–54. https://doi.org/10.1080/09397140.1992.10637623
- Esterbauer, H. (1994) Lebensweise und Verhalten der Würfelnatter im Masil al Fawwar (Syrien). DATZ (Die Aquarien- und Terrarien-Zeitschrift), 47, 308–311.
- Esterbauer, H. (2022) Die Froschlurche des syrischen Bachsystems Masil al Fawwar. Aqua-Terra Austria, 17, 20–27.
- Forskål, P. (1775) Descriptiones Animalium: Avium, Amphibiorum, Piscium, Insectorum, Vermium; quae in Itinere Orientali Observavit. Post mortem auctoris edidit Carsten Niebuhr. Möller, Copenhagen, 164 pp. https://doi.org/10.5962/bhl.title.2154
- Frétey, T. (2022) The authorship and date of the familial nomen Dermochelyidae (Chelonii). Bionomina, 31, 46–69. https://doi.org/10.11646/bionomina.31.1.3
- Fretey, J. & Bour, R. (1980) Redécouverte du type de Dermochelys coriacea (Vandelli) (Testudinata, Dermochelyidae). Bollettino di Zoologia, 47, 193–205. https://doi.org/10.1080/11250008009440336
- Fritz, U., Ayaz, D., Buschbom, J., Kami, H.G., Mazanaeva, L.F., Aloufi, A.A., Auer, M., Rifai, L., Šilic, T. & Hundsdörfer, A.K. (2008) Go East: Phylogeographies of Mauremys caspica and M. rivulata – Discordance of morphology, mitochondrial and nuclear genomic markers and rare hybridization. Journal of Evolutionary Biology, 21, 527–540. https://doi.org/10.1111/j.1420-9101.2007.01485.x
- Fritz, U., Baran, I., Budak, A. & Amthauer, E. (1998) Some notes on the morphology of Emys orbicularis in Anatolia, especially on E. o. luteofusca and E. o. colchica, with the description of a new subspecies from southeastern Turkey. Mertensiella, 10, 103–121.
- Fritz, U., Bischoff, W., Martens, H. & Schmidtler, J.F. (1996) Variabilität syrischer Landschildkröten (Testudo graeca) sowie zur Systematik und Zoogeographie im Nahen Osten und in Nordafrika. Herpetofauna, 18 (104), 5–14.
- Fritz, U., Hundsdörfer, A.K., Široký, P., Auer, M., Kami, H., Lehmann, J., Mazanaeva, L.F., Türkozan, O. & Wink, M. (2007) Phenotypic plasticity leads to incongruence between morphology-based taxonomy and genetic differentiation in western Palaearctic tortoises (Testudo graeca complex; Testudines, Testudinidae). Amphibia-Reptilia, 28, 97–121. https://doi.org/10.1163/156853807779799135
- Fritz, U., Tolley, K.A., Vamberger, M. & Ihlow, F. (2022) Phylogeny and phylogeography of chelonians from sub-Saharan Africa—A review of current knowledge in tribute to Margaretha D. Hofmeyr. Vertebrate Zoology, 72, 951–969. https://doi.org/10.3897/vz.72.e95681
- Fritz, U. & Wischuf, T. (1997) Zur Systematik westasiatisch–südosteuropäischer Bachschildkröten (Gattung Mauremys) (Reptilia: Testudines: Bataguridae). Zoologische Abhandlungen, 49, 223–260.
- GBIF (2023a) GBIF Occurrence Download. https://doi.org/10.15468/dl.8mqra8
- GBIF (2023b) GBIF Occurrence Download https://doi.org/10.15468/dl.6cfrg6
- GBIF (2023c) GBIF Occurrence Download. https://doi.org/10.15468/dl.mmcgd6
- GBIF (2023d) GBIF Occurrence Download. https://doi.org/10.15468/dl.e3e8at
- GBIF (2023e) GBIF Occurrence Download. https://doi.org/10.15468/dl.dbzz25
- GBIF (2023f) GBIF Occurrence Download. https://doi.org/10.15468/dl.nav9uj
- GBIF (2023g) GBIF Occurrence Download. https://doi.org/10.15468/dl.ce9gdh
- Georgalis, G.L. & Joyce, W.G. (2017) A review of the fossil record of Old World turtles of the clade Pan-Trionychidae. Bulletin of the Peabody Museum of Natural History, 58, 115–208. https://doi.org/10.3374/014.058.0106
- Georgalis, G.L., Macaluso, L. & Delfino, M. (2021) A review of the fossil record of Afro-Arabian turtles of the clade Testudinoidea. Bulletin of the Peabody Museum of Natural History, 62, 43–78. https://doi.org/10.3374/014.062.0103
- Gmelin, S.G. (1774) Reise durch Russland zur Untersuchung der drey Natur-Reiche. Dritter Theil. Reise durch das nordliche Persien, in den Jahren 1770, 1771, bis im April 1772. Kayserliche Academie der Wissenschaften, St. Petersburg, 508 pp.
- Griggo, C. (2004) Mousterian fauna from Dederiyeh Cave and comparisons with fauna from Umm El Tlel and Douara Cave. Paléorient, 30, 149–162. https://doi.org/10.3406/paleo.2004.4776
- Groombridge, B. (1990) Marine turtles in the Mediterranean: Distribution, population status, conservation. Nature and Environment Series, 48, 1–98.
- Gruvel, A. (1931) Les Etats de Syrie. Richesses marines et fluviales. Exploitation actuelle – Avenir. Société d’Editions Géographiques, Maritimes et Coloniales (Bibliothèque de la faune des Colonies françaises), Paris, 453 pp.
- Halpin, P.N., Read, A.J., Fujioka, E., Best, B.D., Donnelly, B., Hazen, L.J., Kot, C., Urian, K., LaBrecque, E., Dimatteo, A., Cleary, J., Good, C., Crowder, L.B. & Hyrenbach, K.D. (2009) OBIS-SEAMAP: The world data center for marine mammal, sea bird, and sea turtle distributions. Oceanography, 22, 104–115. https://doi.org/10.5670/oceanog.2009.42
- Handal, E.N., Amr, Z.S., Basha, W.S. & Qumsiyeh, M.B. (2021) Illegal trade in wildlife vertebrate species in the West Bank, Palestine. Journal of Asia-Pacific Biodiversity, 14, 636–639. https://doi.org/10.1016/j.japb.2021.07.004
- Ihlow, F., Ahmadzadeh, F., Ghaffari, H., Taşkavak, E., Hartmann, T., Etzbauer, C. & Rödder, D. (2014) Assessement of genetic structure, habitat suitability and effectivenes of reserves for future onservation planning of the Euphrates soft-shelled turtle, Rafetus euphraticus (Daudin, 1802). Aquatic Conservation: Marine and Freshwater Ecosystems, 24, 831–840. https://doi.org/10.1002/aqc.2454
- iNaturalist (2023) Available from: https://www.inaturalist.org (accessed 4 December 2023)
- IUCN (2024) The IUCN Red List of Threatened Species. Version 2024-1. Available from: https://www.iucnredlist.org (accessed 5 July 2024)
- Iverson, J.B. (2022) A review of chelonian type specimens (order Testudines). Megataxa, 7, 1–85. https://doi.org/10.11646/megataxa.7.1.1
- Jablonski, D. & Sadek, R.A. (2019) The species identity and biogeography of Blanus (Amphisbaenia: Blanidae) in Lebanon. Zoology in the Middle East, 65, 208–214. https://doi.org/10.1080/09397140.2019.1604471
- Javanbakht, H., Ihlow, F., Jablonski, D., Široký, P., Fritz, U., Rödder, D., Sharifi, M. & Mikulíček, P. (2017) Genetic diversity and Quaternary range dynamics in Iranian and Transcaucasian tortoises. Biological Journal of the Linnean Society, 121, 627–640. https://doi.org/10.1093/biolinnean/blx001
- Jony, M. & Rees, A.F. (2008) Three years of monitoring turtle nesting in Syria (2004–2006): What progress has been made towards protection? In: Rees, A.F., Frick, M., Panagopoulou, A., Williams, K. & Kristina, L. (Eds.), Proceedings of the Twenty-seventh Annual Symposium on Sea Turtle Biology and Conservation. 22–28 February 2007. Myrtle Beach, South Carolina, pp. 238–239.
- Jony, M. & Rees, A.F. (2009) Preliminary finding on the interaction between marine turtle and fisheries in Syria. In: Demetropoulos, A. & Turkozan, O. (Eds.), Proceedings of the Second Mediterranean Conference on Marine Turtles. 4–7 May 2005. Kemer, Antalya, pp. 92–95.
- Jony, M. & Rees, F.A. (2022) Additional records of leatherback turtles (Dermochelys coriacea) from Syria. MedTurtle Bulletin, 2, 11–13.
- Kasparek, M. (1994) Die Nil-Weichschildkröte – eine stark bedrohte Reptilienart im Mittelmeergebiet. Herpetofauna, 16 (89), 8–13.
- Kasparek, M. (1995) The nesting of marine turtles on the coast of Syria. Zoology in the Middle East, 11, 51–62. https://doi.org/10.1080/09397140.1995.10637671
- Kasparek, M. & Kinzelbach, R. (1991) Distribution and bionomics of the Nile soft-shelled turtle, Trionyx triunguis, in the Eastern Mediterranean. Zeitschrift für Angewandte Zoologie, 78, 139–159.
- Kinzelbach, R. (1986) Recent records of the Nile soft-shelled turtle, Trionyx triunguis, and the Euphrates soft-shelled turtle, Trionyx euphraticus, in the Middle East. Zoology in the Middle East, 1, 83–87. https://doi.org/10.1080/09397140.1986.10637519
- Kinzelbach, R. (1988) Die Europäische Sumpfschildkröte (Emys orbicularis) im Einzugsgebiet des Rheins. Zeitschrift für Angewandte Zoologie, 75, 385–419.
- Kornilios, P., Jablonski, D., Sadek, R.A., Kumlutaş, Y., Olgun, K., Avcı, A. & Ilgaz, C. (2020) Multilocus species-delimitation in the Xerotyphlops vermicularis (Reptilia: Typhlopidae) species complex. Molecular Phylogenetics and Evolution, 152, 106922. https://doi.org/10.1016/j.ympev.2020.106922
- Krupp, F. (1992) The karst springs of Ras al-Ain. Aqua geographia, 1, 26–33.
- Krupp, F. & Schneider, W. (1991) Bestandserfassung der rezenten Fauna im Bereich des Nahr al-Habur. In: Kühne, H. (Ed.), Die rezente Umwelt von Tall Seh Hamad und Daten zur Umweltrekonstruktion der assyrischen Stadt Dur-Katlimmu (Berichte der Ausgrabung von Tall Seh Hamad/Dur-Katlimmu 1). D. Reimer, Berlin, pp. 69–85.
- Lortet, L. (1883) Poissons et reptiles du Lac de Tibériade et de quelques autres parties de la Syrie. Archives du Muséum d’histoire naturelle de Lyon, 3, 99–194. https://doi.org/10.3406/mhnly.1883.941
- Lymberakis, R. & Kalionzopoulou, A. (2003) Additions to the herpetofauna of Syria. Zoology in the Middle East, 29, 33–39. https://doi.org/10.1080/09397140.2003.10637967
- Martens, H. (1997) A review of “Zoogeography of amphibians and reptiles of Syria, with additional new records” (Herpetozoa 9 (1/2), 1996). Herpetozoa, 10, 99–106.
- Mertens, R. & Müller, L. (1928) Liste der Amphibien und Reptilien Europas. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 41, 1–62.
- Mhanna, H., AlMasri, A. & Ibrahim, N. (2021) The morphology and size of erythrocytes of six reptiles species [sic] from the Syrian coastal region. Scientific Reports in Life Sciences, 2, 40–46. https://doi.org/10.22034/srls.2021.247948
- Mikulíček, P., Jandzik, D., Fritz, U., Schneider, C. & Široký, P. (2013) AFLP analysis shows high incongruence between genetic differentiation and morphology-based taxonomy in a widely distributed tortoise. Biological Journal of the Linnean Society, 108, 151–160. https://doi.org/10.1111/j.1095-8312.2012.01999.x
- Moravec, J. & Orszagh, I. (1998) Testudo graeca terrestris als Wirt von Leptoconops irritans (Diptera: Ceratopogonidae). Salamandra, 34, 181–182.
- Murdoch, D.A. & Aidek, A. (2012) Birding sites of the OSME Region 8 – The birds of the lower Syrian Euphrates. Sandgrouse, 34, 152–176.
- observation.org (2023) Biodiversity citizen science and monitoring. Available from: https://observation.org/ (accessed 4 December 2023)
- Perälä, J. & Bour, R. (2004) Neotype of Testudo terrestris Forsskål, 1775 (Testudines, Testudinidae). Asiatic Herpetological Research, 10, 114–119.
- Plinius, G. Secundus (77–79) Naturalis historia. Reprinted by H.G. Bohn, London (1855–1857).
- Pola, L., Velenská, D., Prondzynska, K.M., Hejduk, V., Zíka, A., Winkelhöfer, T., Abu Baker, M.A., Amr, Z.S. & Šmíd, J. (2023) More geckos, fewer gaps: Diversity and distribution of Hemidactylus geckos (Squamata: Gekkonidae) in Jordan. Systematics and Biodiversity, 21, 2237033. https://doi.org/10.1080/14772000.2023.2237033
- Por, F.D. (1975) An outline of the zoogeography of the Levant. Zoologica Scripta, 4, 5–20. https://doi.org/10.1111/j.1463-6409.1975.tb00713.x
- QGIS Development Team (2023) QGIS Geographic Information System. Open Source Geospatial Foundation Project. Available from: http://qgis.org (accessed 12 July 2024)
- RCoreTeam (2024) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org (accessed 12 July 2024)
- Raza, H.A., Fadhel, O., Ararat, K., Haba, M.K. & Salim, M. (2011) Animal and Bird Trade and Hunting in Iraq. Nature Iraq & Iraqi Ministry of Environment, Sulaimani, 44 pp.
- Rees, A.F., Jony, M., Margaritoulis, D. & Godley, B.J. (2008a) Satellite tracking of a green turtle, Chelonia mydas, from Syria further highlights importance of North Africa for Mediterranean turtles. Zoology in the Middle East, 45, 49–54. https://doi.org/10.1080/09397140.2008.10638306
- Rees, A., Jony, M., Saad, A., Andrea, D. & Phillott, A.D. (2021) Malformations in green turtle embryos and hatchlings in Syria. Marine Turtle Newsletter, 164, 12–15.
- Rees, A.F., Saad, A. & Jony, M. (2004) First record of a leatherback turtle in Syria. Marine Turtle Newsletter, 106, 13.
- Rees, A.F., Saad, A. & Jony, M. (2005) Tagging green turtles (Chelonia mydas) and loggerhead turtles (Caretta caretta) in Syria. Testudo, 6, 51–55.
- Rees, A.F., Saad, A. & Jony, M. (2008b) Discovery of a regionally important green turtle Chelonia mydas rookery in Syria. Oryx, 42, 456–459. https://doi.org/10.1017/S0030605308000926
- Rees, A.F., Saad, A. & Jony, M. (2009) Clutch size and hatching success for green turtle nests in Syria 2004. In: Demetropoulos, A. & Turkozan, O. (Eds.), Proceedings of the Second Mediterranean Conference on Marine Turtles. 4–7 May 2005. Kemer, Antalya, pp. 158–161.
- Rees, A.F., Saad, A. & Jony, M. (2010) Syria. In: Casale, P. & Margaritoulis, D. (Eds.), Sea Turtles in the Mediterranean: Distribution, Threats and Conservation Priorities. IUCN, Gland, pp. 233–243.
- Reshetnikov, A.N., Zibrova, M.G., Ayaz, D., Bhattarai, S., Borodin, O.V., Borzée, A., Brejcha, J., Çiçek, K., Dimaki, M., Doronin, V.I., Drobenkov, S.M., Gichikhanova, U.A., Gladkova, A.Y., Gordeev, D.A., Ioannidis, Y., Ilyukh, M.P., Interesova, E.A., Jadhav, T.D., Karabanov, D.P., Khabibullin, V.F., Khabilov, T.K., Khan, M.H., Kidov, A.A., Klimov, A.S., Kochetkov, D.N., Kolbintsev, V.G., Kuzmin, S.L., Lotiev, K.Y., Louppova, N.E., Lvov, V.D., Lyapkov, S.M., Martynenko, I.M., Maslova, I.V., Masroor, R., Mazanaeva, L.F., Milko, D.A., Milto, K.D., Mozaffari, O., Nguyen, T.Q., Novitsky, R.V., Petrovskiy, A.B., Prelovskiy, V.A., Serbin, V.V., Shi, H-T., Skalon, N.V., Struijk, R.P.J.H., Taniguchi, M., Tarkhnishvili, D., Tsurkan, V.F., Tyutenkov, O.Y., Ushakov, M.V., Vekhov, D.A., Xiao, F., Yakimov, A.V., Yakovleva, T.I., Yang, P., Zeleev, D.F. & Petrosyan, V.G. (2023) Rarely naturalized, but widespread and even invasive: The paradox of a popular pet terrapin expansion in Eurasia. NeoBiota, 81, 91–127. https://doi.org/10.3897/neobiota.81.90473
- Reynaud Savioz, N. (2011) The faunal remains from Nadaouiyeh Aïn Askar (Syria). Preliminary indications of animal acquisition in an Acheulean site. In: Le Tensorer, J.M., Jagher, R. & Otte, M. (Eds.), The Lower and Middle Palaeolithic in the Middle East and Neighbouring Regions. Eraul, Liège, pp. 225–234.
- Rhodin, A.G.J., Stanford, C.B., van Dijk, P.P., Eisemberg, C., Luiselli, L., Mittermeier, R.A., Hudson, R., Horne, B.D., Goode, E.V., Kuchling, G., Walde, A., Baard, E.H.W., Berry, K.H., Bertolero, A., Blanck, T.E.G., Bour, R., Buhlmann, K.A., Cayot, L.J., Collett, S., Currylow, A., Das, I., Diagne, T., Ennen, J.R., Forero-Medina, G., Frankel, M.G., Fritz, U., García, G., Gibbons, J.W., Gibbons, P.M., Gong, S., Guntoro, J., Hofmeyr, M.D., Iverson, J.B., Kiester, A.R., Lau, M., Lawson, D.P., Lovich, J.E., Moll, E.O., Páez, V.P., Palomo-Ramos, R., Platt, K., Platt, S.G., Pritchard, P.C.H., Quinn, H.R., Rahman, S.C., Randrianjafizanaka, S.T., Schaffer, J., Selman, W., Shaffer, H.B., Sharma, D.S.K., Shi, H., Singh, S., Spencer, R., Stannard, K., Sutcliffe, S., Thomson, S. & Vogt, R.C. (2018) Global conservation status of turtles and tortoises (Order Testudines). Chelonian Conservation and Biology, 17, 135–161. https://doi.org/10.2744/CCB-1348.1
- Russell, A. (1756) The Natural History of Aleppo, and Parts Adjacent. A. Millar, London, viii + 266 + (10) pp.
- Saad, A. (2012) Importance of Lattakia Beach (Syria) as nesting area for marine turtles: Results of seven years of field survey. Scholarly Journal of Agricultural Science, 2, 108–110.
- Saad, A. & Rees, A. (2020) Syria. In: Casale, P., Hochscheid, S., Kaska, Y. & Panagopoulou, A. (Eds.), Sea Turtles in the Mediterranean region. IUCN-SSC Marine Turtle Specialist Group Regional Report, 2020, 274–286.
- Saad, A. & Rees, A.F. (2022) Syria. In: Casale, P., Hochscheid, S., Jribi, I., Kaska, Y. & Panagopoulou, A. (Eds.), Sea turtles in the Mediterranean region. 2021 IUCN-SSC Marine Turtle Specialist Group Regional Report, 2021, pp. 449–461.
- Saad, A., Ali, M., Badran, M., Aidek, A., Othman, A., Dib, S., Saleh, M., Ahmad, M. & Badour, F. (2019) [Data on Syria]. In: Panigada, S., Boisseau, O., Canadas, A., Lambert, C., Laran, S., McLanaghan, R. & Moscrop, A. (Eds.), ACCOBAMS, 2021. Estimates of Abundance and Distribution of Cetaceans, Marine Mega-fauna and Marine Litter in the Mediterranean Sea from 2018–2019 Surveys. ACCOBAMS Survey Initiative Project, Monaco, pp. 1–177.
- Saad, A., Ali, A. & Darwich, A. (2006) Marine turtle nesting survey, Syria 2005. In: Frick, M., Panagopoulou, A., Rees, A.F. & Williams, K. (Eds.), Book of Abstracts of 26th Annual Symposium on Sea Turtles. Sea Turtle Society, Athens, p. 320.
- Saad, A., Rees, A., Sabour, W., Othman, A., Genaidi, C. & Alkhateeb, N. (2010) Six years of survey (2004–2009) on reproduction of sea turtles on Shkaifat beach, Syrian coast (proposed as protected area). Rapports et procès-verbaux des réunions Commission internationale pour l’exploration scientifique de la Mer Méditerranée, 39, 651.
- Saad, A., Soulaiman, A. & Alkusairy, H. (2020) Marine turtle nesting survey and stranding assessment from Tartus to Syria’s border with Lebanon. Natura Croatica, 29 (Supplment 1), 51–57.
- Schmidt, K.P. (1939) Reptiles and amphibians from southwestern Asia. Zoological Series, Field Museum of Natural History, 24, 49–92. https://doi.org/10.5962/bhl.title.3841
- Schmidt, K.P. (1953) A Check List of North American Amphibians and Reptiles. Sixth Edition. University of Chicago Press, Chicago, Illinois, 280 pp.
- Schoepff, J.D. (1793) Historia Testudinum Iconibus Illustrata. Fasciculus III et IV. Joannes Jacobus Palm, Erlangen, pp. 33–80, pls. XI–XVI + XVIIB–XX.
- Siebenrock, F. (1913) Wissenschaftliche Ergebnisse der Expedition nach Mesopotamien, 1910. Schildkröten aus Syrien und Mesopotamien. Annalen des Naturhistorischen Museums Wien, 27, 171–225.
- Sindaco, R. & Jeremčenko, V.K. (2008) The Reptiles of the Western Palearctic 1: Annotated Checklist and Distributional Atlas of the Turtles, Crocodiles, Amphisbaenians and Lizards of Europe, North Africa, Middle East and Central Asia. Belvedere, Latina, 580 pp.
- Sindaco, R., Serra, G. & Menegon, M. (2006) New data on the Syrian herpetofauna, with a newly recorded species of snake. Zoology in the Middle East, 37, 29–38. https://doi.org/10.1080/09397140.2006.10638146
- Široký, P., Jandzík, D., Mikulíček, P., Moravec, J. & Országh, I. (2007) Leptoconops bezzii (Diptera: Ceratopogonidae) parasitizing tortoises Testudo graeca (Testudines: Testudinidae) in mountain ranges of Lebanon and western Syria. Parasitology Research, 101, 485–489. https://doi.org/10.1007/s00436-007-0490-2
- Sivan, N. & Werner, Y.L. (1992) Survey of the reptiles of the Golan plateau and Mt. Hermon. Israel Journal of Ecology and Evolution, 37, 193–211. https://doi.org/10.1080/00212210.1992.10688655
- Smith, H.M. & Taylor, E.H. (1950a) Type localities of Mexican reptiles and amphibians. University of Kansas Science Bulletin, 33, 313–380.
- Smith, H.M. & Taylor, E.H. (1950b) An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. United States National Museum Bulletin, 199, 1–253. https://doi.org/10.5479/si.03629236.199
- Sönmez, B., Türkecan, O. & Jded, A. (2017) Long distance movement between nesting sites for two green turtles in the eastern Mediterranean. Marine Turtle Newsletter, 153, 7–8.
- Stanford, C.B., Iverson, J.B., Rhodin, A.G.J., van Dijk, P.P., Mittermeier, R.A., Kuchling, G., Berry, K.H., Bertolero, A., Bjorndal, K.A., Blanck, T.E.G., Buhlmann, K.A., Burke, R.L., Congdon, J.D., Diagne, T., Edwards, T., Eisemberg, C.C., Ennen, J.R., Forero-Medina, G., Frankel, M., Fritz, U., Gallego-García, N., Georges, A., Gibbons, J.W., Gong, S., Goode, E.V., Shi, H.T., Hoang, H., Hofmeyr, M.D., Horne, B.D., Hudson, R., Juvik, J.O., Kiester, R.A., Koval, P., Le, M., Lindeman, P.V., Lovich, J.E., Luiselli, L., McCormack, T.E.M., Meyer, G.A., Páez, V.P., Platt, K., Platt, S.G., Pritchard, P.C.H., Quinn, H.R., Roosenburg, W.M., Seminoff, J.A., Shaffer, H.B., Spencer, R., Van Dyke, J.U., Vogt, R.C. & Walde, A.D. (2020) Turtles and tortoises are in trouble. Current Biology, 30, R721–R735. https://doi.org/10.1016/j.cub.2020.04.088
- Stümpel, N., Rajabizadeh, M., Avcı, A., Wüster, W. & Joger, U. (2016) Phylogeny and diversification of mountain vipers (Montivipera, Nilson et al., 2001) triggered by multiple Plio–Pleistocene refugia and high-mountain topography in the Near and Middle East. Molecular Phylogenetics and Evolution, 101, 336–351. https://doi.org/10.1016/j.ympev.2016.04.025
- Tamar, K., Carranza, S., in den Bosch, H., Sindaco, R., Moravec, J. & Meiri, S. (2015) Hidden relationships and genetic diversity: Molecular phylogeny and phylogeography of the Levantine lizards of the genus Phoenicolacerta (Squamata: Lacertidae). Molecular Phylogenetics and Evolution, 91, 86–97. https://doi.org/10.1016/j.ympev.2015.05.002
- Taşkavak, E. & Akçınar, S.C. (2009) Marine records of the Nile soft-shelled turtle, Trionyx triunguis from Turkey. Marine Biodiversity Records, 2, e9.
- Taşkavak, E., Atatür, M.K., Ghaffari, H. & Meylan, P.A. (2016) Rafetus euphraticus (Daudin 1801) – Euphrates softshell turtle. In: Rhodin, A.G.J., Iverson, J.B., van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Pritchard, P.C.H. & Mittermeier, R.A. (Eds.), Conservation biology of freshwater turtles and tortoises: A compilation project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs, 5, 98.1–98.11. https://doi.org/10.3854/crm.5.098.euphraticus.v1.2016
- Taşkavak, E., Reimann, M.J. & Polder, W.N (1999) First record of the Nile soft-shelled turtle, Trionyx triunguis, from Kos Island, Greece, with comments on its occurrence in the eastern Mediterranean. Chelonian Conservation and Biology, 3, 510–512.
- TTWG [Turtle Taxonomy Working Group] (2021) Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status (9th Ed.). Chelonian Research Monographs, 8, 1–472. https://doi.org/10.3854/crm.8.checklist.atlas.v9.2021
- Türkozan, O., Javanbakht, H., Mazanaeva, L., Meiri, S., Kornilev, Y.V., Tzoras, E., Popgeorgiev, G., Shanas, U. & Escoriza, D. (2023) Testudo graeca Linnaeus 1758 (eastern subspecies clades: Testudo g. armeniaca, Testudo g. buxtoni, Testudo g. ibera, Testudo g. terrestris, Testudo g. zarudnyi) – Armenian tortoise, Zagros tortoise, Anatolian tortoise, Levantine tortoise, Kerman tortoise. In: Rhodin, A.G.J., Iverson, J.B., van Dijk, P.P., Stanford, C.B., Goode, E.V., Buhlmann, K.A. & Mittermeier, R.A. (Eds.), Conservation biology of freshwater turtles and tortoises: A compilation project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs, 5, 120.1–120.33. https://doi.org/10.3854/crm.5.120.eastern.graeca.v1.2023
- Türkozan, O., Özdemir, A. & Kiremit, F. (2008) International Testudo trade. Chelonian Conservation and Biology, 7, 269–272. https://doi.org/10.2744/CCB-0724.1
- Vamberger, M., Stuckas, H., Ayaz, D., Graciá, E., Aloufi, A.A., Els, J., Mazanaeva, L.F., Kami, H.G. & Fritz, U. (2013) Conservation genetics and phylogeography of the poorly known Middle Eastern terrapin Mauremys caspica (Testudines: Geoemydidae). Organisms, Diversity & Evolution, 13, 77–85. https://doi.org/10.1007/s13127-012-0102-6
- Vamberger, M., Stuckas, H., Ayaz, D., Lymberakis, P., Široký, P. & Fritz, U. (2014) Massive transoceanic gene flow in a freshwater turtle (Testudines: Geoemydidae: Mauremys rivulata). Zoologica Scripta, 43, 313–322. https://doi.org/10.1111/zsc.12055
- Vamberger, M., Stuckas, H., Vargas-Ramírez, M., Kehlmaier, C., Ayaz, D., Aloufi, A.A., Lymberakis, P., Široký, P. & Fritz, U. (2017) Unexpected hybridization patterns in Near Eastern terrapins (Mauremys caspica, M. rivulata) indicate ancient gene flow across the Fertile Crescent. Zoologica Scripta, 46, 401–413. https://doi.org/10.1111/zsc.12227
- VertNet (2023) Available from: http://vertnet.org/ (accessed 4 December 2023)
- Veselý, M. (1991) O herpetofaune Syrie. Akvárium-Terárium, 34, 24–25.
- Vigna Taglianti, A., Audisio, P., Biondi, M., Bologna, M., Carpaneto, G., De Biase, A., Fattorini, S., Piattella, E., Sindaco, R., Venchi, A. & Zapparoli, M. (1999) A proposal for a chorotype classification of the Near East fauna, in the framework of the Western Palearctic region. Biogeographia, 20, 31–59. https://doi.org/10.21426/B6110172
- Wallace, B.P., Posnik, Z.A., Hurley, B.J., DiMatteo, A.D., Bandimere, A., Rodriguez, I., Maxwell, S.M., Meyer, L., Brenner, H., Jensen, M.P., LaCasella, E., Shamblin, B.M., Abreu-Grobois, F.A., Stewart, K.R., Dutton, P.H., Barrios-Garrido, H., Dalleau, M., Dell’Amico, F., Eckert, K.L., FitzSimmons, N.N., Garcia-Cruz, M., Hays, G.C., Kelez, S., Lagueux, C.J., Madden Hof, C.A., Marco, A., Martins, S.L.T., Mobaraki, A., Mortimer, J.A., Nel, R., Phillott, A.D., Pilcher, N.J., Putman, N.F., Rees, A.F., Rguez-Baron, J.M., Seminoff, J.A., Swaminathan, A., Turkozan, O., Vargas, S.M., Vernet, P.D., Vilaça, S., Whiting, S.D., Hutchinson, B.J., Casale, P. & Mast, R.B. (2023) Marine turtle regional management units 2.0: An updated framework for conservation and research of wide-ranging megafauna species. Endangered Species Research, 52, 209–223. https://doi.org/10.3354/ESR01243
- Wermuth, H. (1956) Versuch der Deutung einiger bisher übersehener Schildkröten-Namen. Zoologische Beiträge, Neue Folge, 2, 399–423.
- Wermuth, H. (1958) Status und Nomenklatur der Maurischen Landschildkröte, Testudo graeca, in SW-Asien und NO-Afrika. Senckenbergiana biologica, 39, 149–153.
- Werner, F. (1919) Ergebnisse einer mit Unterstützung der Kais. Akademie der Wissenschaften in Wien ausgeführten Forschungsreise von weiland Prof. Dr. Franz Tölg. I. Reptilien und Amphibien aus dem Amanus-Gebirge. Archiv für Naturgeschichte Berlin, 85A/8, 130–141.
- Werner, F. (1935) Im Auto durch Syrien. Blätter für Aquarien- und Terrarienkunde, 46, 225–227.
- Werner, F. (1939) Die Amphibien und Reptilien von Syrien. Abhandlungen und Berichte des Museums für Naturkunde und Vorgeschichte Magdeburg, 7, 211–223.
- Werner, Y.L. & Avital, E. (1980) The herpetofauna of Mt. Hermon and its altitudinal distribution. Israel Journal of Zoology, 29, 192–193.
- Zohary, M. (1973) Geobotanical Foundations of the Middle East. Vol. 1. Gustav Fischer, Swets & Zeitlinger, Stuttgart and Amsterdam, 340 pp.