Atlas of amphibians and reptiles in the Moroccan section of the Mediterranean Intercontinental Biosphere Reserve: distribution patterns and habitat use
DOI:
https://doi.org/10.11160/bah.310Keywords:
Conservation, Diversity, Herpetofauna, IBRM, Land-cover, Species RichnessAbstract
This study presents the first comprehensive atlas of the distribution of amphibians and reptiles within the Moroccan section of the Intercontinental Biosphere Reserve of the Mediterranean (IBRM). Data were gathered from literature reviews, museum collections, and extensive field surveys conducted between 2008 and 2019. A Geographical Information System (GIS) was utilized to generate updated distribution maps, assess species richness within a 10x10 km UTM grid, and analyse associations of species with habitat categories. A total of eight amphibian and 25 reptile species, representing five and 17 families, respectively, were recorded. Three distinct distribution patterns emerged for both taxonomic groups: widespread species, species restricted to specific environments, such as high-altitude areas or coastal environments, and species with limited and / or fragmented ranges. Seventeen areas of high species richness (N > 10 species) were identified across the study area. These areas consistently overlapped across both taxonomic groups and were primarily associated with Mediterranean-type habitats. While amphibians exhibited relatively homogeneous habitat preferences, reptiles demonstrated four distinct assemblages: species prefer mixed vegetation mosaics; species occupy forest-agriculture edges; species adaptable to diverse habitats and environments and species privilege open grasslands-rocky areas. The IBRM’s complex topography likely contributes to high levels of environmental diversity, supporting a remarkable proportion of Morocco’s herpetofauna: 60% of amphibian species and 25% of reptile species. These findings underscore the critical importance of the Moroccan IBRM for herpetofaunal conservation at the national and international levels.
References
Argaz, H.; Brito, J.C.; Fahd, S.; Martínez‐Freiría, F.; Boudajbir, C. & Geniez, P. (2020). Atlas des amphibiens et reptiles endémiques du Maroc: répartition et état de conservation. Bulletin de la Société Herpetologique de France 176: 61-94.
Mafalda Barata, M. & Harris, D.J. (2015). Variation cryptique chez le lézard d'altitude marocain Atlantolacerta andreanskyi (Squamata : Lacertidae). African Journal of Herpetology 64: 71370.
Ben Hassine, J.; Gutiérrez-Rodríguez, J.; Escoriza, D. & Martínez-Solano, I. (2016). Inferring the roles of vicariance, climate and topography in population differentiation in Salamandra algira (Caudata, Salamandridae). Journal of Zoological Systematics and Evolutionary Research 54: 116-126.
Beukema, W.; Bok, B.; Tiemann, L. & Speybroeck, J. (2015). Local hybridisation between native Triturus cristatus and introduced Triturus marmoratus (Urodela: Salamandridae) in the Netherlands. Herpetology Notes 8: 549-552.
Beukema, W.; de Pous, P.; Donaire-Barroso, D.; Bogaerts, S.; Garcia-Porta, J.; Escoriza, D.; Arribas, O.J.; El Mouden, E.H. & Carranza, S. (2013). Review of the systematics, distribution, biogeography, and natural history of Moroccan amphibians. Zootaxa 3661: 1-60.
Bingham, H.; Lewis, E.; Belle, E.; Stewart, J.; Klimmek, H.; Wicander, S.; Bhola, N. & Bastin, L. (2021). Protected Planet Report 2020: Tracking Progress Towards Global Targets for Protected and Conserved Areas. UNEP-WCMC & IUCN. International Union for Conservation of Nature, Gland, Switzerland.
Bons, J. & Geniez, P. (1996). Anfibios y Reptiles de Marruecos (Incluido Sáhara Occidental). Atlas Biogeográfico. Asociación Herpetológica Española, Barcelona, Spain.
Bouazza, A.; El Mouden, E.H. & Rihane, A. (2021). Checklist of amphibians and reptiles of Morocco: A taxonomic update and standard Arabic names. Herpetology Notes 14: 1-14.
Brito, J.C.; Fahd, S.; Geniez, P.; Martínez‐Freiría, F.; Pleguezuelos, J.M. & Trape, J.F. (2011). Biogeography and conservation of viperids from North‐West Africa: an application of ecological niche‐based models and GIS. Journal of Arid Environments 75: 1029‐1037.
Brito, J.C.; Godinho, R.; Martínez‐Freiría, F.; Pleguezuelos, J.M.; Rebelo, H.; Santos, X.; Vale, C.G.; Velo-Antón, G.; Boratyński, Z.; Carvalho, S.B.; Ferreira, S.; Gonçalves, D.V.; Silva, T.L.; Tarroso, P.; Campos, J.C.; Leite, J.V.; Nogueira, J.; Álvares, F.; Sillero, N.; Sow, A.S.; Fahd, S.; Crochet, P.-A. & Carranza, S. (2014). Unravelling biodiversity, evolution and threats to conservation in the Sahara‐Sahel. Biological Reviews 89: 215-231.
Buchhorn, M.; Bertels, L.; Smets, B.; De Roo, B.; Lesiv, M.; Tsendbazar, N.E.; Masiliunas, D. & Linlin, L. (2020). Copernicus Global Land Service: Land Cover 100m: Version 3 Globe 2015-2019: Algorithm Theoretical Basis Document. Zenodo, Geneve, Switzerland. Available at https://doi-org.10.5281/zenodo.3938968. Accessed on 20 August 2024.
Challender, D.W.S.; Cremona, P.J.; Malsch, K.; Robinson, J.E.; Pavitt, A.T.; Scott, J.; Hoffmann, R.; Joolia, A.; Oldfield, T.; Jenkins, R.; Conde, D.; Hilton-Taylor, C. & Hoffmann, M. (2023). Identifying species likely threatened by international trade on the IUCN Red List can inform CITES trade measures. Nature Ecology & Evolution 7: 1211-1220.
Chergui, B.; Fahd S.; Santos, X. & Pausas, J.G. (2024). Moroccan cannabis farms threaten biodiversity. Science 385: 941.
Climate Data (2024). Climate Data for Cities Worldwide. Alexander Merkel, Oedheim, Germany. Available at https://en.climate-data.org/africa/morocco-181/. Retrieved on 23 November 2024.
Cox, N.; Chanson, J. & Stuart, S. (2006). The Status and Distribution of Reptiles and Amphibians of the Mediterranean Basin Mediterranean. International Union for Conservation of Nature, Gland, Switzerland.
Cox, R.L. & Underwood, E.C. (2011). The importance of conserving biodiversity outside of protected areas in Mediterranean ecosystems. PLoS ONE 6: e14508.
Crochet, P.A.; Sweet, S.S. & Mateo, J.A. (2004). Recent records of the rare Psammodromus microdactylus (Boettger, 1881) in Morocco. Herpetozoa 17: 184‐186.
de Pous, P.; Beukema, W.; Weterings, M.; Dummer, I. & Geniez, P. (2011). Area prioritization and performance evaluation of the conservation area network for the Moroccan herpetofauna: a preliminary assessment. Biodiversity and Conservation 20: 89‐118.
Dinerstein, E.; Olson, D.; Joshi, A.; Vynne, C.; Burgess, N.D.; Wikramanayake, E.; Hahn, N.; Palminteri, S.; Hedao, P.; Noss, R.; Hansen, M.; Locke, H.; Ellis, E.C..; Jones, B.; Barber, C.V.; Hayes, R.; Kormos, C.; Martin, V.; Crist, E.; Sechrest, W.; Price, L.; Baillie, J.E.M.; Weeden, D.; Suckling, K.; Davis, C.; Sizer, N.; Moore, R.; Thau, D.; Birch, T.; Potapov, P.; Turubanova, S.; Tyukavina, A.; de, Souza, N.; Pintea, L.; Brito, J.C.; Llewellyn, O.A.; Miller, A.G.; Patzelt, A.; Ghazanfar, S.A.; Timberlake, J.; Klöser, H.; Shennan-Farpon, Y.; Kindt, R.; Lilleso, J-P.B.; van, Breugel, P.; Graudal, L.; Voge, M.; Al-Shammari, K.F. & Saleem, M. (2017). An ecoregion-based approach to protecting half the terrestrial realm. BioScience 67: 534-545.
Dinis, M.; Merabet, K.; Martínez-Freiría, F.; Steinfartz, S.; Vences, M.; Burgon, J. D. & Velo-Antón, G. (2019). Allopatric diversification and evolutionary melting pot in a North African Palearctic relict: The biogeographic history of Salamandra algira. Molecular Phylogenetics and Evolution 130: 81-91.
Donaire‐Barroso, D. & Bogaerts, S. (2003). A new subspecies of Salamandra algira Bedriaga, 1883 from northern Morocco. Podarcis 4: 84‐100.
Doniol-Valcroze, P.; Rancilhac, L.; Brito, J.C.; Miralles, A.; Geniez, P.; Benoit, L.; Loiseau, A.; Leblois, R.; Dufresnes, C. & Crochet, P.A. (2024). Combining RADseq and contact zone analysis to decipher cryptic diversification in reptiles: Insights from the Spiny-footed Lizard (Reptilia, Lacertidae). Zoologica Scripta 53: 1-15.
El Kourchi, Y.; Santos, X.; Fahd, S. & Chergui, B. (2025). Occupancy analysis uncovers environmental factors contributing to road mortality of amphibians. European Journal of Wildlife Research 71: 53.
Enriquez-Urzelai, U.; Martínez-Freiría, F.; Freitas, I.; Perera, A.; Martínez-Solano, Í.; Salvi, D.; Velo-Antón, G., & Kaliontzopoulou, A. (2022). Allopatric speciation, niche conservatism and gradual phenotypic change in the evolution of European green lizards. Journal of Biogeography 49: 2193-2205.
Fahd, S. & Mediani, M. (2007). Herpétofaune du Bassin Versant d’Oued Laou. Wadi 6 FP, INCO‐CT2005‐015226, Tétouan, Morocco.
Fahd, S. & Pleguezuelos, J.M. (1996). Los reptiles del Rif (norte de Marruecos), I: Quelonios, Saurios. Revista Española de Herpetología 10: 55-89.
Fahd, S. & Pleguezuelos, J.M. (2001). Los reptiles del Rif (Norte de Marruecos), II: Anfisbenios y Ofidios. Comentarios sobre la biogeografía del grupo. Revista Española de Herpetología 15: 13-36.
Fahd, S. (2001). Biogéographie, Morphologie et Ecologie des Ophidiens du Rif (Nord du Maroc). Thèse d’Etat, Université Abdelmalek Essaâdi, Tétouan, Morocco.
Fahd, S.; Mediani, M.; Other, A.M.; Denys, C. & Santos, X. (2015). Diversity and conservation of amphibians in the Oued Laou Catchent Rif (Rif, northwestern Morocco). Travaux de l’Institut Scientifique 8: 69-84.
Foden, W.B.; Butchart, S.H.M.; Stuart, S.N.; Vié, J.-C.; Akçakaya, H.R.; Angulo, A.; DeVantier, L.M.; Gutsche, A.; Turak, E., Cao, L., Donner, S.D., Katariya, V., Bernard, R., Holland, R.A.; Hughes, A.F.; O’Hanlon, S.E.; Garnett, S.T.; Şekercioğlu, C.H. & Mace, G.M. (2013). Identifying the World’s most climate change vulnerable species: A systematic trait-based assessment of all birds, amphibians and corals. PLoS ONE 8: e65427.
GBIF.org (2025). GBIF Home Page. Global Biodiversity Information Facility, Copenhagen, Denmark. Available at https://www.gbif.org/. Retrieved on 04 May 2025.
Geniez, P.; Mateo, J.A. & Geniez, M. (2004). The Amphibians and Reptiles of the Western Sahara and Adjacent Regions. Edition Chimaira, Frankfurt am Main, Germany.
Gregory, A.; Spence, E.; Beier, P. & Garding, E. (2021). Toward best management practices for ecological corridors. Land 10: 140.
Harris, D.J.; Carretero, M.A.; Brito, J.C.; Kaliontzopoulou, A.; Pinho, C.; Perera, A.; Vasconcelos, R.; Barata, M.; Barbosa, D.; Carvalho, S.; Fonseca, M.M.; Pérez-Lanuza, G. & Rato, C. (2008). Data on the distribution of the terrestrial herpetofauna of Morocco: records from 2001‒2006. Herpetological Bulletin 103: 19-28.
Harris, D.J.; Dimitra, S. & Faria, J.F. (2024). New data on the diversity and distribution of lineages of the Acanthodactylus erythrurus species complex in North Africa derived from mitochondrial DNA markers. Herpetozoa 37: 281-286.
Hidalgo, P.J.; Hernández, H.; Sánchez-Almendro, A.J.; López-Tirado, J.; Vessella, F. & Porras, R. (2021). Fragmentation and connectivity of island forests in agricultural Mediterranean environments: A comparative study between the Guadalquivir Valley (Spain) and the Apulia Region (Italy). Forests 12: 1201.
Hilty, J.; Worboys, G.L.; Keeley, A.; Woodley, S.; Lausche, B.; Locke, H.; Carr, M.; Pulsford I.; Pittock, J.; White, J.W.; Theobald, D.M.; Levine, J.; Reuling, M.; Watson, J.E.M.; Ament, R. & Tabor, G.M. (2020). Guidelines for Conserving Connectivity Through Ecological Networks and Corridors. Best Practice Protected Area Guidelines Series No. 30. International Union for Conservation of Nature, Gland, Switzerland.
IUCN (2024). IUCN Red List of Threatened Species. Version 2024-2. International Union for Conservation of Nature, Gland, Switzerland. Available at https://www.iucnredlist.org. Retrieved on 27 August 2024.
Margules, C.R. & Pressey, R.L. (2000). Systematic conservation planning. Nature 405: 243‐253.
Margules, C.R. & Sarkar, S. (2007). Systematic Conservation Planning. Cambridge University Press, Cambridge, UK.
Martínez del Mármol, G.; Harris, D.J.; Geniez, P.; de Pous, P. & Salvi, D. (2019). Amphibians and Reptiles of Morocco. Edition Chimaira. Frankfurt am Main, Germany.
Martínez-Freiría, F.; Argaz, H.; Fahd, S. & Brito, J.C. (2013). Climate change is predicted to negatively influence Moroccan endemic reptile richness. Implications for conservation in protected areas. Naturwissenschaften 100: 877-889.
Martínez-Freiría, F.; Bouazza, A.; Buldain, J.; Freitas, I.; Avella, I.; Scaramuzzi, A.; Sioumpoura, K.; Oliveira, D. & Fahd, S. (2023). Fieldwork campaigns and citizen science data increase the distributional range of the elusive Vipera monticola in Morocco. Basic and Applied Herpetology 37: 95-105.
Martínez-Freiría, F.; Fahd, S.; Bouazza, A.; Crochet, P.-A. & Beddek, M. (2024). Vipera monticola. The IUCN Red List of Threatened Species 2024: e.T221202628A3105206. Available at https://dx.doi.org/10.2305/IUCN.UK.2024-2.RLTS.T221202628A3105206.en. Retrieved on 17 May 2025.
Martínez-Freiría, F.; Freitas, I.; Velo-Antón, G.; Lucchini, N.; Fahd, S.; Larbes, S.; Pleguezuelos, J.M.; Santos, X. & Brito, J.C. (2021). Integrative taxonomy reveals two species and intraspecific differentiation in the Vipera latastei–monticola complex. Journal of Zoological Systematics and Evolutionary Research 59: 2278-2306.
Martínez‐Freiría, F.; Freitas, I.; Zuffi, M.A.; Golay, P.; Ursenbacher, S. & Velo‐Antón, G. (2020). Climatic refugia boosted allopatric diversification in western Mediterranean vipers. Journal of Biogeography 47: 1698-1713.
Mediani, M.; Brito, J.C. & Fahd, S. (2015). Atlas of the amphibians and reptiles of northern Morocco: Updated distribution and patterns of habitat selection. Basic and Applied Herpetology 29: 1-27.
Miralles, A.; Geniez, P.; Beddek, M.; Aranda, D.M.; Brito, J.C.; Leblois, R. & Crochet, P.-A. (2020). Morphology and multilocus phylogeny of the Spiny-footed Lizard (Acanthodactylus erythrurus) complex reveal two new mountain species from the Moroccan Atlas. Zootaxa 4747: 302-326.
Nori, J.; Lemes, P.; Urbina-Cardona, N.; Baldo, D.; Lescano, J. & Loyola, R. (2015). Amphibian conservation, land-use changes and protected areas: A global overview. Biological Conservation 191: 367-374.
Nori, J.; Villalobos, F. & Loyola, R. (2018). Global priority areas for amphibian research. Journal of Biogeography 45: 2588-2594.
Pough, F.H. (1980). The advantages of ectothermy for tetrapods. American Naturalist 115: 92‐112.
Schleich, H.H.; Kästle, W. & Kabisch, K. (1996). Amphibians and Reptiles of North Africa. Koeltz Scientific Books, Koenigstein, Germany.
Sinervo, B.; Méndez-de-la-Cruz, F.; Miles, D.B.; Heulin, B.; Bastiaans, E.; Villagrán-Santa-Cruz, M.; Lara-Resendiz, R.; Martínez-Méndez, N.; Calderón-Espinosa, M.L.; Meza-Lázaro, R.N.; Gadsden, H.; Avila, L.J.; Morando, M.; De, la, Riva, I.J.; Victoriano-Sepulveda, P.; Rocha, C.F.D.; Ibargüengoytía, N.; AguilarPuntriano, C.; Massot, M.; Lepetz, V.; Oksanen, T.A.; Chapple, D.G.; Bauer, A.M.; Branch, W.R.; Clobert, J. & Sites, J.W. (2010). Erosion of lizard diversity by climate change and altered thermal niches. Science 328: 894-899.
Steininger, F.F.; Rabeder, G. & Rögl, F. (1985). Land mammal distribution in the Mediterranean Neogene: A consequence of geokinematic and climatic events, In D.J. Stanly & F.C. Wezel (eds.) The Geological Evolution of the Eastern Mediterranean. Springer, New York, USA, pp. 559-571.
Stuckas, H.; Velo-Antón, G.; Fahd, S.; Kalboussi, M.; Rouag, R.; Arculeo, M., Marrone, F.; Sacco, F.; Vamberger, M & Fritz, U. (2014). Where are you from, stranger? The enigmatic biogeography of North African pond turtles (Emys orbicularis). Organisms Diversity & Evolution 14: 295-306.
Uetz, P.; Freed, P.; Aguilar, R.; Reyes, F.; Kudera, J. & Hošek, J. (2025). The Reptile Database. Richmond, Virginia, USA. Available at https://www.reptile-database.org/. Retrieved on 24 April 2025 .
UNEP-WCMC (2024). CITES Trade Database: Amphibians and Reptiles of Morocco. United Nations Environment Programme & World Conservation Monitoring Centre, Cambridge, UK. Available at https://www.trade.cites.org/. Retrieved on 06 May 2025 .
UNESCO (2006). Intercontinental BR of the Mediterranean. United Nations Educational, Scientific and Cultural Organization, Paris, France. Available at https://www.unesco.org/en/mab/intercontinental-br-mediterranean-0. Retrieved on 12 January 2025 .
Valdés, B.; Rejdali, M.; Achhal el Kadmiri, A.; Jury, J.L. & Montserrat, J.M. (2002). Catalogue des Plantes Vasculaires du Nord du Maroc, Incluant des Clés d’Identification. Consejo Superior de Investigaciones Científicas, Madrid, Spain.
Veith, M.; Fromhage, L.; Kosuch, J. & Vences, M. (2006). Historical biogeography of Western Palaearctic pelobatid and pelodytid frogs: A molecular phylogenetic perspective. Contributions to Zoology 75: 109-120.
Veith, M.; Mayer, C.; Samraoui, B.; Barroso, D.D. & Bogaerts, S. (2004). From Europe to Africa and vice versa: Evidence for multiple intercontinental dispersal in ribbed salamanders (genus Pleurodeles). Journal of Biogeography 31: 159-171.
Velo-Antón, G.; García-Cardenete, L.; Jiménez Cazalla, F. & Martínez-Freiría, F. (2014). New record of Salamandra algira isolated on the north-western Tingitana peninsula, with some notes on the reproductive modes within the species. Boletín de la Asociación Herpetológica Española 25: 46-50.
Downloads
Published
Issue
Section
License
Copyright (c) 2025 See B&AH copyright notice
This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License.
