Amphibians and Reptiles of Ethiopia—Biogeography, Conservation Priority Areas, and Potentials for “One Plan Approach” Conservation Measures

Abstract

Ethiopia, as one of 36 global hotspots of biodiversity, provides diverse habitats for a variety of species, but little research about the herpetofauna of the country is available. Worldwide, especially amphibians but also reptiles, are threatened by biodiversity and habitat loss due to deforestation and other human actions. Following the One Plan Approach by the International Union for Conservation of Nature, an integrative use of ex situ and in situ measures is recommended to address the ongoing conservation crisis, which is currently largely missing for Ethiopia's 76 amphibian and 191 reptile species. In an effort to close gaps in conservation work, we analyze species richness, protected area coverage, and corrected weighted endemism as well as the threat status and status of ex situ populations for these species. We show the South-Western lowlands as the most important region of high species diversity and threatened and endemic species, having a species richness of 12–14 for amphibians and 50–54 for reptiles. Additional important hotspots are the South-Western Highlands and Arsi-Bale Highlands; the protection of these three regions should be prioritized. For protected areas, the Bale Mountain National Park is particularly important, sheltering 25 amphibians, 7 of which are threatened and 8 endemic, as well as 2 endemic reptiles. The Arsi Mountains National Park also houses a large number of species, with many threatened and endemic ones. Although many priority areas are already under protection, the extension of existing and establishment of new protected areas, for example, in the Southern and South-Western Lowlands, would be advantageous for species conservation. With only 3 amphibian and 43 reptile species in holding, largely with low number of institutions or individuals, representation of Ethiopian species in ex situ facilities is insufficient and priorities should be shifted to endemic and threatened species and the building of breeding programs. For conservation-oriented holding stations and zoos, a list of the 33 most threatened amphibian and reptile species is provided.

Keywords

biodiversity hotspots / corrected weighted endemism / ex situ populations / extinction risk / protected areas / species richness patterns / threat analysis

References

[1] Abebe, F. B., and S. E. Bekele. 2018. “Challenges to National Park Conservation and Management in Ethiopia.” Journal of Agricultural Science 10, no. 5: 52–62. https://doi.org/10.5539/jas.v10n5p52.

[2] Andreone, F., A. Channing, R. Drewes, et al. 2008. “ Amphibians of the Afrotropical Realm.” In Threatened Amphibians of the World, edited by S. N. Stuart, M. Hoffmann, J. S. Chanson, et al., 53–58. Lynx Edicions, IUCN Conservation International.

[3] *Arnold, E. N.2009. “Relationships, Evolution and Biogeography of Semaphore geckos, Pristurus (Squamata, Sphaerodactylidae) Based on Morphology.” Zootaxa 2060: 1–21.

[4] Asefa, M., M. Cao, Y. He, E. Mekonnen, X. Song, and J. Yang. 2020. “Ethiopian Vegetation Types, Climate and Topography.” Plant Diversity 42, no. 4: 302–311.

[5] AZA (Association of Zoos and Aquariums). 2021. Species Survival Plan® (SSP) Program Handbook. Association of Zoos and Aquariums.

[6] *Badiane, A., J. Garcia-Porta, J. Červenka, et al. 2014. “Phylogenetic Relationships of Semaphore Geckos (Squamata: Sphaerodactylidae: Pristurus) With an Assessment of the Taxonomy of Pristurus rupestris.” Zootaxa 3835, no. 1: 33–58.

[7] Bain, R. H., and M. M. Hurley. 2011. “A Biogeographic Synthesis of the Amphibians and Reptiles of Indochina.” Bulletin of the American Museum of Natural History 360: 1–138.

[8] Billi, P.2015a. “ Geomorphological Landscapes of Ethiopia.” In Landscapes and Landforms of Ethiopia, edited by P. Billi, 3–32. Springer.

[9] Billi, P. (ed.). 2015b. Landscapes and Landforms of Ethiopia. Springer.

[10] Bishop, P. J., A. Angulo, J. P. Lewis, R. D. Moore, G. B. Rabb, and J. G. Moreno. 2012. “The Amphibian Extinction Crisis - What Will It Take to Put the Action Into the Amphibian Conservation Action Plan?” Surveys and Perspectives Integrating Environment and Society 5, no. 2: 97–111.

[11] Böhm, M., B. Collen, J. E. M. Baillie, et al. 2013. “The Conservation Status of the World's Reptiles.” Biological Conservation 157: 372–385.

[12] Boulenger, G. A.1896. “A List of the Reptiles and Batrachians Collected by Dr. Ragazzi in Shoa and Eritrea.” Annali del Museo Civico di Storia Naturale Giacomo Doria 2, no. 36: 545–554.

[13] *Bour, R.2008. “ Pelusios adansonii (Schweigger 1812) – Adanson's Mud Turtle.” In Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group, edited by A. G. J. Rhodin, P. C. H. Pritchard, P. P. van Dijk, R. A. Saumure, K. A. Buhlmann and J. B. Iverson, Chelonian Research Monographs, 017.1–017.4. Chelonian Research Foundation.

[14] Broadley, D. G.1994. “A Collection of Snakes From Eastern Sudan, With the Description of a New Species of Telescopus Wagler, 1830 (Reptilia: Ophidia).” Journal of African Zoology 108, no. 2: 201–208.

[15] *Broadley, D. G.2002. “A Review of the Species of Psammophis boie Found South of Latitude 12° S (Serpentes: Psammophiinae).” African Journal of Herpetology 51, no. 2: 83–119.

[16] *Broadley, D. G., and R. C. Boycott. 2009. “ Pelusios sinuatus (Smith 1838) – Serrated Hinged Terrapin.” In Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group, edited by A. G. J. Rhodin, P. C. H. Pritchard, P. P. Dijk, et al., 036.1–036.5. Chelonian Research Monographs.

[17] Broadley, D. G., and B. Hughes. 1993. “A Review of the Genus Lycophodion (Serpentes: Colubridae) in Northeastern Africa.” Herpetological Journal 3: 8–18.

[18] *Broadley, D. G., and V. Wallach. 2007a. “A Review of East and Central African Species of Letheobia Cope, Revived From the Synonymy of Rhinotyphlops Fitzinger, With Descriptions of Five New Species (Serpentes: Typhlopidae).” Zootaxa 1515: 31–68.

[19] *Broadley, D. G., and V. Wallach. 2007b. “A Revision of the Genus Leptotyphlops in Northeastern Africa and Southwestern Arabia (Serpentes: Leptotyphlopidae).” Zootaxa 1408: 1–78.

[20] Brown, J. L., A. Cameron, A. D. Yoder, and M. Vences. 2014. “A Necessarily Complex Model to Explain the Biogeography of the Amphibians and Reptiles of Madagascar.” Nature Communications 5: 5046.

[21] Brown, J. L., N. Sillero, F. Glaw, P. Bora, D. R. Vieites, and M. Vences. 2016. “Spatial Biodiversity Patterns of Madagascar's Amphibians and Reptiles.” PLoS One 11, no. 1: e0144076.

[22] Byers, O., C. Lees, J. Wilcken, and C. Schwitzer. 2013. “The One Plan Approach: The Philosophy and Implementation of CBSG's Approach to Integrated Species Conservation Planning.” WAZA Magazine 14: 2–5.

[23] CBD. 2023. Establishment of a New Trust Fund: Global Biodiversity Framework Fund. 64th GEF Council Meeting June 26 – June 29, 2023 Brasilia, Brazil.

[24] Ceballos, G., and P. R. Ehrlich. 2006. “Global Mammal Distributions, Biodiversity Hotspots, and Conservation.” Proceedings of the National Academy of Sciences 103, no. 51: 19374–19379.

[25] Chamberlain, S. 2022. Rredlist. ‘Iucn’ Red List Client. R Package Version 0.7.1. https://cran.R-project.Org/package=rredlist.

[26] *Channing, A., A. Hillers, S. Lötters, et al. 2013. “Taxonomy of the Super-Cryptic Hyperolius nasutus Group of Long Reed Frogs of Africa (Anura: Hyperoliidae), With Descriptions of Six New Species.” Zootaxa 3620: 301–350.

[27] Chanson, J., M. Hoffmann, N. Cox, and S. Stuart. 2008. “ The State of the World's Amphibians.” In Threatened Amphibians of the World, edited by S. N. Stuart, M. Hoffmann and J. S. Chanson, et al., 33–44. Lynx Edicions, IUCN, Conservation International.

[28] *Cherlin, A.1990. “Таксономическая Ревизия Змей Рода Echis (Viperidae). Ii. Анализ Таксономии И Описание Новых Форм В. А. Черлин Зоологический Институт Ан Ссср (Ленинград).” Proceedings of the Zoological Institute Leningrad 207: 1–27.

[29] *Chippaux, J.-P. 1999. Les serpents d'Afrique occidentale et centrale. IRD Éditions. 1 ed.

[30] *Chippaux, J.-P. 2006. Les serpents d'Afrique occidentale et centrale. IRD Éditions. 3rd ed.

[31] *Chirio, L., and I. Ineich. 2006. “Biogeography of the Reptiles of the Central African Republic.” African Journal of Herpetology 55, no. 1: 23–59.

[32] Chirio, L., I. Ineich, A. Schmitz, and J. F. Trape. 2011. “Note sur la systématique de quelques espèces du genre Prosymna Gray, 1849 en Afrique au nord de l'équateur (Serpentes, Prosymnidae).” Revue Suisse de Zoologie 118, no. 1: 157–173.

[33] Clulow, J., V. L. Trudeau, and J. Kouba. 2014. “ Amphibian Declines in the Twenty-First Century: Why We Need Assisted Reproductive Technologies.” In Reproductive Sciences in Animal Conservation: Progress and Prospects, edited by W. V. Holt, J. L. Brown and P. Comizzoli, 275–316. Springer.

[34] Clulow, J., J. Upton, V. L. Trudeau, and S. Clulow. 2019. “ Amphibian Assisted Reproductive Technologies: Moving From Technology to Application.” In Reproductive Sciences in Animal Conservation. Advances in Experimental Medicine and Biology, edited by P. Comizzoli, J. L. Brown and W. V. Holt, 413–464. Springer.

[35] Collins, J. P., and M. L. Crump. 2009. Extinction in Our Times: Global Amphibian Decline. Oxford University Press.

[36] Colston, T. J., R. A. Pyron, and A. M. Bauer. 2020. “A New Species of African Snake-Eyed Skink (Scincidae: Panaspis) From Ethiopia.” Zootaxa 4779, no. 2: 190–200.

[37] Cox, N., S. Stuart, and J. Chanson, et al. 2008. “ An Introduction to the Amphibians.” In Threatened Amphibians of the World, edited by S. N. Stuart, M. Hoffmann and J. S. Chanson, et al., 2–17. Lynx Edicions, IUCN, Conservation International.

[38] Cox, N., B. E. Young, P. Bowles, et al. 2022. “A Global Reptile Assessment Highlights Shared Conservation Needs of Tetrapods.” Nature 605, no. 7909: 285–290.

[39] Crist, E., C. Mora, and R. Engelman. 2017. “The Interaction of Human Population, Food Production, and Biodiversity Protection.” Science 356: 260–264.

[40] Daru, B. H., M. van der Bank, and T. J. Davies. 2015. “Spatial Incongruence Among Hotspots and Complementary Areas of Tree Diversity in Southern Africa.” Diversity and Distributions 21, no. 7: 769–780.

[41] Dessie, G., and J. Kleman. 2007. “Pattern and Magnitude of Deforestation in the South Central Rift Valley Region of Ethiopia.” Mountain Research and Development 27, no. 2: 162–168.

[42] Evans, B. J., S. M. Bliss, S. A. Mendel, and R. C. Tinsley. 2011. “The Rift Valley Is a Major Barrier to Dispersal of African Clawed Frogs (Xenopus) in Ethiopia: Evolution of African Clawed Frogs in Ethiopia.” Molecular Ecology 20, no. 20: 4216–4230.

[43] Frankham, R., J. D. Ballou, and D. A. Briscoe. 2010. Introduction to Conservation Genetics. Cambridge University Press. 2nd ed.

[44] Freed, P., P. Uetz, R. Aguilar, F. Reyes, and J. Hošek. 2023. The Reptile Database. Electronic Database. http://www.reptile-database.org/.

[45] Freilich, X., J. D. Anadón, J. Bukala, O. Calderon, R. Chakraborty, and S. Boissinot. 2016. “Comparative Phylogeography of Ethiopian Anurans: Impact of the Great Rift Valley and Pleistocene Climate Change.” BMC Evolutionary Biology 16, no. 1: 206.

[46] Freilich, X., M. Tollis, and S. Boissinot. 2014. “Hiding in the Highlands: Evolution of a Frog Species Complex of the Genus Ptychadena in the Ethiopian Highlands.” Molecular Phylogenetics and Evolution 71: 157–169.

[47] Freitas, E. S., A. M. Bauer, C. D. Siler, D. G. Broadley, and T. R. Jackman. 2018. “Phylogenetic and Morphological Investigation of the Mochlus afer-sundevallii Species Complex (Squamata: Scincidae) Across the Arid Corridor of Sub-Saharan Africa.” Molecular Phylogenetics and Evolution 127: 280–287.

[48] Friis, I., S. Demissew, and P. van Breugel. 2010. “Atlas of the Potential Vegetation of Ethiopia.” Royal Danish Academy of Sciences and Letters 58: 1–308.

[49] Frost, D. R. 2023. Amphibian Species of the World: An Online Reference. Version 6.1. American Museum of Natural History. Version 6.1. American Museum of Natural History, New York. https://amphibiansoftheworld.amnh.org/.

[50] *Gans, C.2005. “Checklist and Bibliography of the Amphisbaenia of the World.” Bulletin of the American Museum of Natural History 289: 1–130.

[51] *Goutte, S., J. Reyes-Velasco, and S. Boissinot. 2019. “A New Species of Puddle Frog From an Unexplored Mountain in Southwestern Ethiopia (Anura, Phrynobatrachidae, Phrynobatrachus).” ZooKeys 824: 53–70.

[52] Goutte, S., J. Reyes-Velasco, X. Freilich, A. Kassie, and S. Boissinot. 2021. “Taxonomic Revision of Grass Frogs (Ptychadenidae, Ptychadena) Endemic to the Ethiopian Highlands.” ZooKeys 1016: 77–141.

[53] Goutte, S., J. Reyes-Velasco, A. Kassie, and S. Boissinot. 2022. “Genetic and Morphometric Analyses of Historical Type Specimens Clarify the Taxonomy of the Ethiopian Leptopelis gramineus Species Complex (Anura, Arthroleptidae).” ZooKeys 1128: 63–97.

[54] Gower, D. J., R. K. Aberra, S. Schwaller, et al. 2013. “Long-Term Data for Endemic Frog Genera Reveal Potential Conservation Crisis in the Bale Mountains, Ethiopia.” Oryx 47, no. 1: 59–69.

[55] Gower, D. J., T. M. Doherty-Bone, R. K. Aberra, et al. 2012. “High Prevalence of the Amphibian Chytrid Fungus (Batrachochytrium dendrobatidis) Across Multiple Taxa and Localities in the Highlands of Ethiopia.” Herpetological Journal 22, no. 4: 225–233.

[56] Gower, D. J., E. O. Z. Wade, S. Spawls, et al. 2016. “A New Large Species of Bitis Gray, 1842 (Serpentes: Viperidae) From the Bale Mountains of Ethiopia.” Zootaxa 4093, no. 1: 41–63.

[57] Habel, J. C., L. Rasche, U. A. Schneider, et al. 2019. “Final Countdown for Biodiversity Hotspots.” Conservation Letters 12: e12668.

[58] Guedes, J. J. M., H. V. Gomes De Lima, L. R. Mendonça, R. Chen-Zhao, J. A. F. Diniz-Filho, and M. R. Moura. 2024. “Temporal Trends in Global Reptile Species Descriptions Over Three Decades.” Systematics and Biodiversity 22: 2419832.

[59] Harfoot, M. B. J., A. Johnston, A. Balmford, et al. 2021. “Using the IUCN Red List to Map Threats to Terrestrial Vertebrates at Global Scale.” Nature Ecology & Evolution 5, no. 11: 1510–1519.

[60] Hewitt, G. M.2004. “The Structure of Biodiversity - Insights From Molecular Phylogeography.” Frontiers in Zoology 1, no. 1: 4.

[61] Hijmans, R. J., R. Bivand, E. Pebesma, and M. D. Sumner. 2023. terra. e Spatial Data Analysis. R package version 1.6-53. https://cran.r-project.org/web/packages/terra/index.html.

[62] Hijmans, R. J., and J. van Etten. 2022. Raster: Geographic Analysis and Modeling With Raster Data. R Package Version 3.0-15. http://CRAN.R-project.org/package=raster.

[63] IPBES (Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services). 2019. Global Assessment Report of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services, edited by E. S. Brondízio, J. Settele, S. Díaz and H. T. Ngo, IPBES Secretariat. https://zenodo.org/records/6417333.

[64] IUCN (International Union for Conservation of Nature and Natural Resources). 2012. Habitats Classification Scheme, Version 3.1. https://www.iucnredlist.org/resources/habitat-classification-scheme.

[65] IUCN (International Union for Conservation of Nature and Natural Resources). 2023. The IUCN Red List of Threatened Species. Version 2022-2. Electronic Database. https://www.iucnredlist.org.

[66] IUCN/SSC (International Union for Conservation of Nature and Natural Resources/Species Survival Commission). 2014. Guidelines on the Use of Ex Situ Management for Species Conservation. Version 2.0. IUCN Species Survival Commission. Gland.

[67] Jacken, A., D. Rödder, and T. Ziegler. 2020. “Amphibians in Zoos: A Global Approach on Distribution Patterns of Threatened Amphibians in Zoological Collections.” International Zoo Yearbook 54, no. 1: 146–164.

[68] Jetz, W., C. Rahbek, and R. K. Colwell. 2004. “The Coincidence of Rarity and Richness and the Potential Signature of History in Centres of Endemism.” Ecology Letters 7, no. 12: 1180–1191.

[69] Jiren, T. S., J. Leventon, N. W. Jager, et al. 2021. “Governance Challenges at the Interface of Food Security and Biodiversity Conservation: A Multi-Level Case Study From Ethiopia.” Environmental Management 67, no. 4: 717–730. https://doi.org/10.1007/s00267-021-01432-7.

[70] Jung, M., P. R. Dahal, S. H. M. Butchart, et al. 2020. “A Global Map of Terrestrial Habitat Types.” Scientific Data 7, no. 1: 256.

[71] Kafash, A., S. Ashrafi, M. Yousefi, et al. 2020. “Reptile Species Richness Associated to Ecological and Historical Variables in Iran.” Scientific Reports 10, no. 1: 18167.

[72] *Karin, B. R., M. Metallinou, J. L. Weinell, T. R. Jackman, and A. M. Bauer. 2016. “Resolving the Higher-Order Phylogenetic Relationships of the Circumtropical Mabuya Group (Squamata: Scincidae): An Out-of-Asia Diversification.” Molecular Phylogenetics and Evolution 102: 220–232.

[73] Kassie, T. A.2016. “Distribution and Species Richness of Amphibians in Awi Zone and Wombera District, Northwest Ethiopia.” Journal of Zoology Studies 3, no. 4: 1–6.

[74] Kassie, T. A., and S. A. Bekele. 2020. “Diversity, Distribution, and Habitat of Herpetefauna Around Gambella Zuria District, West Ethiopia.” International Journal of Zoology and Applied Biosciences 5, no. 2: 68–78.

[75] Kassie, T. A., S. A. Bekele, and B. Afework Bogale. 2022. “Species Composition and Distribution of Endemic Frog Species of Keffa, Southwest Ethiopia.” Global Ecology and Conservation 38: e02211.

[76] Kielgast, J., D. Rödder, M. Veith, and S. Lötters. 2010. “Widespread Occurrence of the Amphibian Chytrid Fungus in Kenya.” Animal Conservation 13: 36–43.

[77] Kier, G., and W. Barthlott. 2001. “Measuring and Mapping Endemism and Species Richness: A New Methodological Approach and Its Application on the Flora of Africa.” Biodiversity & Conservation 10: 1513–1529.

[78] Kilpatrick, A. M., C. J. Briggs, and P. Daszak. 2010. “The Ecology and Impact of Chytridiomycosis: An Emerging Disease of Amphibians.” Trends in Ecology & Evolution 25, no. 2: 109–118.

[79] *Kilunda, F. K., W. Conradie, D. V. Wasonga, et al. 2019. “Revalidation and Resurrection of Panaspis massaiensis (Angel, 1924) and the Description of a New Species of Panaspis Cope (Squamata: Scincidae) From South-Eastern Kenya.” Zootaxa 4706, no. 2: 255–274.

[80] *Koppetsch, T.2020. “A New Species of Trachylepis (Squamata: Scincidae) From the Amhara Region, Ethiopia, and a Key to the Ethiopian Trachylepis.” Zootaxa 4859, no. 1: 113–126.

[81] Koppetsch, T., P. Nečas, and B. Wipfler. 2021. “A New Chameleon of the Trioceros affinis Species Complex (Squamata, Chamaeleonidae) From Ethiopia.” Zoosystematics and Evolution 97, no. 1: 161–179.

[82] Krzikowski, M., T. Q. Nguyen, C. T. Pham, et al. 2022. “Assessment of the Threat Status of the Amphibians in Vietnam - Implementation of the One Plan Approach.” Nature Conservation 49: 77–116.

[83] Lacy, R. C.2013. “Achieving True Sustainability of Zoo Populations.” Zoo Biology 32, no. 1: 19–26.

[84] *Lanza, B.1990. “Amphibians and Reptiles of the Somali Democratic Republic: Check List and Biogeography.” Biogeographia – The Journal of Integrative Biogeography 14: 407–467.

[85] *Largen, M. J., and J. B. Rasmussen. 1993. “Catalogue of the Snakes of Ethiopia (Reptilia Serpentes), Including Identification Keys.” Tropical Zoology 6, no. 2: 313–434.

[86] Largen, M. J., and S. Spawls. 2001. Amphibians and Reptiles Recorded from the Bale Mountains. Walia, Special Edition on the Bale Mountains, 89–93.

[87] Largen, M. J., and S. Spawls. 2006. “Lizards of Ethiopia (Reptilia Sauria): An Annotated Checklist, Bibliography, Gazetteer and Identification Key.” Tropical Zoology 19: 21–109.

[88] Largen, M. J., and S. Spawls. 2010. Amphibians and Reptiles of Ethiopia and Eritrea. Edition Chimaira.

[89] Lewin, A., A. Feldman, A. M. Bauer, et al. 2016. “Patterns of Species Richness, Endemism and Environmental Gradients of African Reptiles.” Journal of Biogeography 43, no. 12: 2380–2390.

[90] Loader, S. P., D. J. Gower, W. Ngalason, and M. Menegon. 2010. “Three New Species of Callulina (Amphibia: Anura: Brevicipitidae) Highlight Local Endemism and Conservation Plight of Africa's Eastern Arc forests: THREE NEW SPECIES OF CALLULINA.” Zoological Journal of the Linnean Society 160, no. 3: 496–514.

[91] *Loveridge, A.1947. “Revision of the African Lizards of the Family Gekkonidae.” Bulletin of the Museum of Comparative Zoology at Harvard College 98, no. 1: 1–469.

[92] Luedtke, J. A., J. Chanson, K. Neam, et al. 2023. “Ongoing Declines for the World's Amphibians in the Face of Emerging Threats.” Nature 622, no. 7982: 308–314.

[93] Marchese, C.2015. “Biodiversity Hotspots: A Shortcut for a More Complicated Concept.” Global Ecology and Conservation 3: 297–309.

[94] *Malonza, P. K., and A. M. Bauer. 2022. “Resurrection of the African Gecko Genus Ancylodactylus Müller, 1907 (Squamata: Gekkonidae) and Description of Six New Species From Kenya.” Zootaxa 5141, no. 2: 101–139.

[95] Mason, N., M. Ward, J. E. M. Watson, O. Venter, and R. K. Runting. 2020. “Global Opportunities and Challenges for Transboundary Conservation.” Nature Ecology & Evolution 4, no. 5: 694–701.

[96] *Mazuch, T., J. Šmíd, and A. M. Bauer. 2016. “Rediscovery and a New Record of Hemidactylus laevis (Reptilia: Gekkonidae) From Somaliland, With Notes on and Resurrection of Hemidactylus fragilis.” Zootaxa 4117, no. 4: 529–542.

[97] *Medina, M. F., A. M. Bauer, W. R. Branch, et al. 2016. “Molecular Phylogeny of Panaspis and Afroablepharus Skinks (Squamata: Scincidae) in the Savannas of Sub-Saharan Africa.” Molecular Phylogenetics and Evolution 100: 409–423.

[98] Meiri, S., A. M. Bauer, A. Allison, et al. 2018. “Extinct, Obscure or Imaginary: The Lizard Species With the Smallest Ranges.” Diversity and Distributions 24, no. 2: 262–273.

[99] Mengistu, A. A., P. Nagel, A. Getahun, S. A. Saber, and S. P. Loader. 2013. “Updated Review of Amphibian Diversity, Distribution and Conservation in Ethiopia.” Ethiopian Journal of Biological Sciences 12, no. 1: 81–116.

[100] Mengist, W.2020. “Challenges of Protected Area Management and Conservation Strategies in Ethiopia: A Review Paper.” Advances in Environmental Studies 4, no. 1: 277–285.

[101] Mestanza-Ramón, C., S. M. Henkanaththegedara, P. Vásconez Duchicela, et al. 2020. “In-Situ and Ex-Situ Biodiversity Conservation in Ecuador: A Review of Policies.” Actions and Challenges Diversity 12, no. 8: 1–18.

[102] *Metallinou, M., J. Červenka, P. A. Crochet, et al. 2015. “Species on the Rocks: Systematics and Biogeography of the Rock-Dwelling Ptyodactylus geckos (Squamata: Phyllodactylidae) in North Africa and Arabia.” Molecular Phylogenetics and Evolution 85: 208–220.

[103] Mittermeier, R. A., W. R. Turner, F. W. Larsen, T. M. Brooks, and C. Gascon. 2011. “ Global Biodiversity Conservation: The Critical Role of Hotspots.” In Biodiversity Hotspots, edited by F. Zachos and J. Habel, 3–22. Springer.

[104] Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. da Fonseca, and J. Kent. 2000. “Biodiversity Hotspots for Conservation Priorities.” Nature 403: 853–858.

[105] * NHM (Natural History Museum London). 2023. Natural History Museum London Data Portal. https://data.nhm.ac.uk/search.

[106] Oljirra, A.2019. “The Causes, Consequences and Remedies of Deforestation in Ethiopia.” Journal of Degraded and Mining Lands Management 6, no. 3: 1747–1754.

[107] Pabijan, M., G. Palomar, B. Antunes, W. Antoł, P. Zieliński, and W. Babik. 2020. “Evolutionary Principles Guiding Amphibian Conservation.” Evolutionary Applications 13, no. 5: 857–878.

[108] *Parker, H. W.1942. “The Lizards of British Somaliland.” Bulletin of the Museum of Comparative Zoology, Harvard 91, no. 1: 1–101.

[109] Pineda, E., and J. M. Lobo. 2009. “Assessing the Accuracy of Species Distribution Models to Predict Amphibian Species Richness Patterns.” Journal of Animal Ecology 78, no. 1: 182–190.

[110] Powney, G. D., R. Grenyer, C. D. L. Orme, I. P. F. Owens, and S. Meiri. 2010. “Hot, Dry and Different: Australian Lizard Richness Is Unlike That of Mammals, Amphibians and Birds.” Global Ecology and Biogeography 19, no. 3: 386–396.

[111] Poynton, J. C., S. P. Loader, E. Sherratt, and B. T. Clarke. 2007. “Amphibian Diversity in East African Biodiversity Hotspots: Altitudinal and Latitudinal Patterns.” Biodiversity and Conservation 16, no. 4: 1103–1118.

[112] Pressey, R. L., C. J. Humphries, C. R. Margules, R. I. Vane-Wright, and P. H. Williams. 1993. “Beyond Opportunism: Key Principles for Systematic Reserve Selection.” Trends in Ecology & Evolution 8, no. 4: 124–128.

[113] Rech, I., P. Ginal, A. Rauhaus, T. Ziegler, and D. Rödder. 2023. “Geckos in Zoos: A Global Approach on Distribution Patterns of Threatened Geckos (Gekkota) in Zoological Institutions.” Journal for Nature Conservation 75: 126467.

[114] *Rocha, S., M. A. Carretero, and D. J. Harris. 2010. “On the Diversity, Colonization Patterns and Status of Hemidactylus spp. (Reptilia: Gekkonidae) From the Western Indian Ocean Islands.” Herpetological Journal 20: 83–89.

[115] de Sá, R. O., S. P. Loader, and A. Channing. 2004. “A New Species of Callulina (Anura: Microhylidae) From the West Usambara Mountains, Tanzania.” Journal of Herpetology 38, no. 2: 219–224.

[116] Saber, S. A., and M. F. Masood. 2021. “Contribution of Reptile's Diversity at Lake Tana Basin, Ethiopia.” Life Science Journal 18, no. 5: 42–49.

[117] SCBD (Secretariat of the Convention on Biological Diversity). 2005. Handbook of the Convention on Biological Diversity Including Its Cartagena Protocol on Biosafety. Friesen. 3rd ed.

[118] *Sindaco, R., and V. K. Jeremčenko. 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 (Vol. 1). Edizioni Belvedere.

[119] *Sindaco, R., R. Nincheri, and B. Lanza. 2014. “Catalogue of Arabian Reptiles in the Collections of the “La Specola” Museum, Florence.” Scripta Herpetologica: 137–164.

[120] * SMF (Senckenberg Museum Frankfurt). 2023. Senckenberg Aquila Datenbank. https://search.senckenberg.de/aquila-public-search/search.

[121] *Šmíd, J., T. Mazuch, L. Nováková, et al. 2020. “Phylogeny and Systematic Revision of the Gecko Genus Hemidactylus From the Horn of Africa (Squamata: Gekkonidae).” Herpetological Monographs 33, no. 1: 26.

[122] *Šmíd, J., J. Moravec, L. Kratochvíl, et al. 2015. “Multilocus Phylogeny and Taxonomic Revision of the Hemidactylus robustus Species Group (Reptilia, Gekkonidae) With Descriptions of Three New Species From Yemen and Ethiopia.” Systematics and Biodiversity 13, no. 4: 346–368.

[123] Šmíd, J., M. Uvizl, M. Shobrak, et al. 2022. “Diversification of Hemidactylus geckos (Squamata: Gekkonidae) in Coastal Plains and Islands of Southwestern Arabia With Descriptions and Complete Mitochondrial Genomes of Two Endemic Species to Saudi Arabia.” Organisms Diversity & Evolution 23, no. 1: 185–207.

[124] Smith, M. L., B. P. Noonan, and T. J. Colston. 2017. “The Role of Climatic and Geological Events in Generating Diversity in Ethiopian Grass Frogs (Genus Ptychadena).” Royal Society Open Science 4, no. 8: 170021.

[125] Soares, C., and J. C. Brito. 2007. “Environmental Correlates for Species Richness Among Amphibians and Reptiles in a Climate Transition Area.” Biodiversity and Conservation 16, no. 4: 1087–1102.

[126] *Spawls, S., K. Howell, H. Hinkel, and M. Menegon. 2018. Field Guide to East African Reptiles. Bloomsbury. 2nd ed.

[127] *Speybroeck, J., W. Beukema, C. Dufresnes, et al. 2020. “Species List of the European Herpetofauna – 2020 Update by the Taxonomic Committee of the Societas Europaea Herpetologica.” Amphibia-Reptilia 41, no. 2: 139–189.

[128] Stenger, L., A. Große Hovest, T. Q. Nguyen, et al. 2023. “Assessment of the Threat Status of Reptile Species From Vietnam - Implementation of the One Plan Approach to Conservation.” Nature Conservation 53: 183–221.

[129] Stuart, S. N., J. S. Chanson, N. A. Cox, et al. 2004. “Status and Trends of Amphibian Declines and Extinctions Worldwide.” Science 306, no. 5702: 1783–1786.

[130] *Tamar, K., L. Chirio, M. Shobrak, S. Busais, and S. Carranza. 2019. “Using Multilocus Approach to Uncover Cryptic Diversity Within Pseudotrapelus Lizards From Saudi Arabia.” Saudi Journal of Biological Sciences 26, no. 7: 1442–1449.

[131] Tamar, K., and J. Moravec. 2022. “First Exact Record and Phylogenetic Position of the Gecko Ptyodactylus hasselquistii (Squamata: Phyllodactylidae) From Lebanon.” Zootaxa 5200, no. 5: 449–469.

[132] Tamar, K., S. Scholz, P. A. Crochet, et al. 2016. “Evolution Around the Red Sea: Systematics and Biogeography of the Agamid Genus Pseudotrapelus (Squamata: Agamidae) From North Africa and Arabia.” Molecular Phylogenetics and Evolution 97: 55–68.

[133] Tan, W. C., A. Herrel, and D. Rödder. 2023. “A Global Analysis of Habitat Fragmentation Research in Reptiles and Amphibians: What Have We Done so Far?” Biodiversity and Conservation 32: 439–468.

[134] Temesgen, F., and B. Warkineh. 2018. “Biodiversity Status & Conservation Challenges of Protected Areas of Ethiopia: Awash & Nechsar National Parks in Focus.” Journal of Natural Sciences Research 8, no. 5: 46–61.

[135] Therneau, T. 2023. A Package for Survival Analysis in R. R package Version 3.5-5. https://CRAN.R-project.org/package=survival.

[136] Tilman, D., M. Clark, D. R. Williams, K. Kimmel, S. Polasky, and C. Packer. 2017. “Future Threats to Biodiversity and Pathways to Their Prevention.” Nature 546, no. 7656: 73–81.

[137] *Tiutenko, A., C. Koch, M. Pabijan, and O. Zinenko. 2022. “Generic Affinities of African House Snakes Revised: A New Genus for Boodon erlangeri (Serpentes: Elapoidea: Lamprophiidae: Lamprophiinae).” Salamandra 58, no. 4: 235–262.

[138] Tiutenko, A., and O. Zinenko. 2021. “A New Species of Leptopelis (Anura, Arthroleptidae) From the South-Eastern Slope of the Ethiopian Highlands, With Notes on the Leptopelis gramineus Species Complex and the Revalidation of a Previously Synonymised Species.” ZooKeys 1023: 119–150.

[139] Tolley, K. A., G. J. Alexander, W. R. Branch, P. Bowles, and B. Maritz. 2016. “Conservation Status and Threats for African Reptiles.” Biological Conservation 204: 63–71.

[140] *Trape, J. F.2021. “Three New Snakes of the Genus Lycophidion Fitzinger, 1843 (Squamata: Lamprophiidae) in Central Africa.” Bulletin de la Société Herpétologique de France 178: 24–37.

[141] *Trape, J. F., P. A. Crochet, D. G. Broadley, et al. 2019. “On the Psammophis sibilans Group (Serpentes, Lamprophiidae, Psammophiinae) North of 12° S, With the Description of a New Species From West Africa.” Bonn Zoological Bulletin 68, no. 1: 61–91.

[142] *Trape, J. F., Y. Mané, and Y. Ineich. 2006. “Atractaspis microlepidota, A. micropholis et A. watsoni en Afrique occidentale et centrale.” Bulletin de la Société Herpétologique de France 119: 5–16.

[143] *Trape, J. F., and O. Mediannikov. 2016. “Cinq serpents nouveaux du genre Boaedon Duméril, Bibron & Duméril, 1854 (Serpentes: Lamprophiidae) en Afrique centrale.” Société Herpétologique de France 159: 61–111.

[144] TTWG (Turtle Taxonomy Working Group), A. G. J. Rhodin, J. B. Iverson, et al. 2021. “ Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status.” In Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group, edited by A. G. J. Rhodin, J. B. Iverson and P. P. Dijk, et al., Chelonian Research Monographs, 1–472. 9th ed. Chelonian Research Foundation in association with the Turtle Conservancy.

[145] UN List of Least Developed Countries. UN Trade & Development. 2024. https://unctad.org/topic/least-developed-countries/list.

[146] UNEP-WCMC & IUCN (United Nations Environment Programme - World Conservation Monitoring Centre & International Union for Conservation of Nature and Natural Resources). 2023. Protected Area Profile for Ethiopia From the World Database on Protected Areas. www.protectedplanet.net.

[147] *Wagner, P., W. Freund, D. Modry, A. Schmitz, and W. Böhme. 2011. “Studies on African Agama IX. New Insights Into Agama finchi Böhme et al., 2005 (Sauria: Agamidae), With the Description of a New Subspecies.” Bonn Zoological Bulletin 60, no. 1: 25–34.

[148] *Wagner, P., T. Mazuch, and A. M. Bauer. 2013. “An Extraordinary Tail - Integrative Review of the Agamid Genus Xenagama.” Journal of Zoological Systematics and Evolutionary Research 51, no. 2: 144–164.

[149] Wahle, A., D. Rödder, D. G. Chapple, S. Meiri, A. Rauhaus, and T. Ziegler. 2021. “Skinks in Zoos: A Global Approach on Distribution Patterns of Threatened Scincidae in Zoological Institutions.” Global Ecology and Conservation 30: e01800.

[150] Wake, D. B., and V. T. Vredenburg. 2008. “Are We in the Midst of the Sixth Mass Extinction? A View From the World of Amphibians.” Proceedings of the National Academy of Sciences 105: 11466–11473.

[151] *Wallach, V., K. Williams, and J. Boundy. 2014. Snakes of the World. A Catalogue of Living and Extinct Species. CRC Press, Taylor & Francis Group.

[152] Wickham, H. 2016. ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag.

[153] Witzenberger, K. A., and A. Hochkirch. 2011. “Ex Situ Conservation Genetics: A Review of Molecular Studies on the Genetic Consequences of Captive Breeding Programmes for Endangered Animal Species.” Biodiversity and Conservation 20, no. 9: 1843–1861.

[154] Zancolli, G., I. Steffan-Dewenter, and M.-O. Rödel. 2014. “Amphibian Diversity on the Roof of Africa: Unveiling the Effects of Habitat Degradation, Altitude and Biogeography.” Diversity and Distributions 20, no. 3: 297–308.

[155] Ziegler, T., J. Kamphausen, F. Glaw, et al. 2022. “Threatened Malagasy Amphibians and Reptiles in Zoos—A Call for Enhanced Implementation of the IUCN's One Plan Approach.” Zoologischer Garten 90: 21–69.

[156] Ziegler, T., A. Rauhaus, C. Niggemann, J. Nicolaudius, M. Bernardes, and T. Q. Nguyen. 2020. “Developing a Conservation Breeding Network for Threatened Vietnamese Crocodile Newts.” AArk Newsletter 52: 9–12.

[157] *Zimkus, B. M.2008. “Geographic Distribution: Cacosternum plimptoni.” Herpetological Review 39, no. 2: 233.

[158] Zimkus, B. M., L. P. Lawson, M. F. Barej, et al. 2017. “Leapfrogging Into New Territory: How Mascarene Ridged Frogs Diversified Across Africa and Madagascar to Maintain Their Ecological Niche.” Molecular Phylogenetics and Evolution 106: 254–269.

[159] ZIMS (Zoological Information Management System). 2023. https://zims.species360.org.

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