• Título:  ZOOGEOGRAFÍA Y ECOLOGÍA EVOLUTIVA DEL JABALÍ (SUS SCROFA) Y LAS LIEBRES (LEPUS SPP.) EN EL NOROESTE DE ÁFRICA
• Autor:  Soria Boix, Maria del Carmen
• Universidad:  Universidad de Alicante
• Departamento:  Ciencias ambientales y recursos naturales
• Fecha de Lectura:  14/10/2020
• Programa de doctorado:  Programa de Doctorado en Ciencias Experimentales y Biosanitarias por la Universidad de Alicante
• Dirección: 
  • URIOS MOLINER, VICENTE (DIRECTOR)
  • DONAT TORRES, MARIA DEL PILAR (DIRECTOR)
• Tribunal: 
  • Belliure Ferrer, Josabel (PRESIDENTE)
  • LIMIÑANA MORCILLO, RUBÉN (SECRETARIO)
  • Pérez García, Juan Manuel (VOCAL)
• Descriptores: 
  • GENETICA MOLECULAR
  • GENETICA DE POBLACIONES
  • ECOLOGIA ANIMAL
• Fichero de tesis:
• http://hdl.handle.net/10045/116886
• Resumen:
  INTRODUCCIÓN O MOTIVACIÓN DE LA TESIS Debido a las especiales características del noroeste de África, esta zona constituye un lugar de estudio interesante para entender cómo los cambios que se producen en el entorno pueden afectar a la distribución de las especies y a su historia evolutiva. El estudio de las especies desde el punto de vista de la ecología evolutiva y la zoogeografía, junto a la realización de análisis moleculares, sirve tanto para entender por qué las especies han llegado a ser como son en la actualidad, como para entender los procesos evolutivos de cada especie. Con este trabajo buscamos aportar nuevos conocimientos sobre algunas especies que habitan este lugar; buscamos el profundizar en el estudio de las especies de interés, relacionando los resultados obtenidos mediante el análisis de su ADN con las características bioclimáticas y geográficas actuales y pasadas. Para esta tesis los animales seleccionados fueron el jabalí (Sus scrofa) y las liebres (Lepus spp.). Capítulo I. Jabalíes (Sus scrofa): Los objetivos iniciales para este caso incluyen el profundizar en las relaciones filogenéticas del jabalí del norte de África a partir del análisis de su ADN, y el estudio de los tiempos de divergencia obtenidos de los árboles filogenéticos; de esta manera podremos saber cuándo se produjeron los eventos que nos interesa conocer como, por ejemplo, el momento en el que se pudo producir el contacto entre jabalíes africanos y europeos. Evaluar si los eventos climáticos o la acción humana han tenido algo que ver en que estos contactos se produzcan, y analizar la teoría de que el estrecho de Gibraltar haya facilitado el movimiento de ejemplares entre África y Europa. Capítulo II. Liebres (Lepus spp.): En resumen, los objetivos iniciales son: adentrarnos en el entendimiento de la ecología evolutiva de las liebres del noroeste de África; entender la forma en la que han evolucionado las liebres en esta área geográfica y la posibilidad de que allí haya un grupo de especies crípticas; estudiar la presencia o ausencia de la liebre del Cabo (Lepus capensis) en el noroeste de África, así como la posible presencia de otras especies; en el caso de obtener datos que apunten a la presencia de una nueva especie, seguir las pautas pertinentes para describirla como tal. Si se tratan de especies ya descritas anteriormente, realizar una revisión bibliográfica para sugerir el nombre más acorde para ellas. Por lo tanto, en esta tesis se busca aportar nueva información sobre los jabalíes y liebres del noroeste de África a partir de aunar diferentes enfoques: el de la ecología evolutiva, la zoogeografía, la filogenómica y, en menor medida, la morfología. Se espera contribuir a saber más sobre el papel que juegan los cambios bioclimáticos y otras presiones evolutivas en la especiación de los animales en el noroeste de África. DESARROLLO TEÓRICO En el noroeste de África hay muchas especies cuya revisión es interesante para entender, a través de su análisis molecular, su ecología evolutiva y posición zoogeográfica. Esto es así porque en la actualidad aún hay pocos estudios al respecto. Tener un enfoque multidisciplinar a la hora de estudiar los procesos evolutivos que subyacen a la especiación y distribución actual de las especies es necesario. El manejo de distintas disciplinas permite importantes avances en la comprensión de los procesos ecológicos y evolutivos de los animales, sobre todo en las especies con procesos evolutivos más complejos. Las herramientas moleculares son muy útiles para aclarar ciertos aspectos que la taxonomía tradicional por sí sola no puede, pero pueden aparecer limitaciones a la hora de interpretar los resultados por sí solos. La ecología evolutiva tiene en cuenta la historia evolutiva de las especies. Analiza las adaptaciones evolutivas de los organismos al medio y las interacciones entre especies. La interpretación de la especiación y distribución de los organismos en términos de eventos de dispersión y aislamiento en respuesta a los cambios climáticos y de la vegetación es el objetivo principal de la biogeografía. En el caso de la zoogeografía, se centra en la diversidad de especies animales en el tiempo y espacio, ayudándonos a entender el porqué de la presencia o ausencia de una especie en un lugar y momento determinados. Hay numerosos trabajos que relacionan el estudio de las especies en el momento presente en relación con su zoogeografía, y otros que analizan lo que ocurrió en el pasado. En esta tesis nos centramos en el estudio del jabalí (Sus scrofa) y las liebres (Lepus spp.) del noroeste de África. En la actualidad, la distribución de los jabalíes en el norte de África se limita a parte del Magreb (Marruecos, Túnez y Argelia). La bibliografía actual agrupa a los jabalíes del norte de África con los jabalíes de Europa, cuando lo esperado sería que fueran más cercanos a los jabalíes del Cercano Oriente. Por lo tanto, vimos interesante el estudio del cómo y cuándo del contacto entre ejemplares europeos y africanos. Por otro lado, estudiamos las liebres, cuya posición filogenética en el noroeste de África aún no está clara. Por tanto, se decidió adentrarnos en el estudio de las liebres presentes en el noroeste de África, con un punto de vista multidisciplinar, para entender mejor las relaciones filogenéticas existentes. CONCLUSIONES Este amplio estudio nos ha llevado a las siguientes conclusiones: - Hemos encontrado haplotipos de jabalí europeo en jabalíes africanos, lo que indicaría un contacto entre ambas poblaciones. - Los tiempos de divergencia de los árboles filogenéticos indican que estos haplotipos compartidos por los jabalíes europeos y africanos aparecieron hace una media de 90.000 años. Para que aparezcan en jabalíes de ambos continentes, tienen que haberse producido flujo genético posterior a esta época. - Proponemos que el estrecho de Gibraltar hace 90.000 años en adelante (durante el Pleistoceno Superior) no constituiría una barrera en épocas glaciales, siendo permeable al paso de animales terrestres, y permitiendo que el jabalí europeo se dispersara hacia el sur a través del mismo. - La ecología evolutiva de las liebres muestra el porqué de la semejanza morfológica entre distintas especies, provocada por la similitud de las condiciones ambientales de los lugares donde habitan, y a su misma respuesta etológica y morfológica. - Los análisis moleculares realizados demuestran la existencia de Lepus mediterraneus, descrita con anterioridad, como especie diferenciada presente en el norte de Marruecos, Túnez, Libia, Egipto y en Cerdeña. - Demostramos la existencia de Lepus schlumbergeri, también descrita con anterioridad, como especie diferenciada cuya distribución abarca desde el sur del Rif hasta el Sáhara, solapándose en el norte con Lepus mediterraneus y en el sur con Lepus saharae sp. nov. - Describimos una nueva especie, Lepus saharae sp. nov., especialmente adaptada al desierto y que se distribuye en el oeste del desierto del Sáhara desde la costa Atlántica hasta Mali, según los datos existentes, aunque posiblemente tenga una distribución mayor. - Estas nuevas descripciones deberían sustituir a lo que hay aceptado actualmente que indica la presencia de una sola especie en todo el norte de África, Lepus mediterraneus o Lepus capensis. BIBLIOGRAFÍA CONSULTADA Abramson, N.I., Rodchenkova, E.N., Kostygov, A. 2009. Genetic variation and phylogeography of the bank vole (Clethrionomys glareolus, Arvicolinae, Rodentia) in Russia with special reference to the introgression of the mtDNA of a closely related species, red-backed vole (C. rutilus). Genetika. 45(5):610-23. https://doi.org/10.1134/S1022795409050044 PMID: 19534420 Agadjanian, A.K., Shunkov, M.V. 2018. Paleolithic Man of Denisova Cave and Zoogeography of Pleistocene Mammals of Northwestern Altai. Paleontol. J. 52:66¿89. https://doi.org/10.1134/S0031030118010021 Agrawal, A.A. 2011. Current trends in the evolutionary ecology of plant defence. Func. Ecol. 25:420-432. https://doi.org/10.1111/j.1365-2435.2010.01796.x Agustí, J., Garcés, M., Krijgsman, W. 2006. Evidence for African-Iberian exchanges during the Messinian in the Spanish mammalian record. Palaeogeogr. Palaeoclimatol. Palaeoecol. 238:5¿14. https://doi.org/10.1016/j.palaeo.2006.03.013 Alves, P.C., Ferrand, N., Suchentrunk, F., Harris, D.J. 2003. Ancient introgression of Lepus timidus mtDNA into L. granatensis and L. europaeus in the Iberian Peninsula. Mol. Phylogent. Evol. 27:70¿80. https://doi.org/10.1016/S1055-7903(02)00417-7 PMID: 12679072 Alves, E., Ovilo, C., Rodriguez, M.C., Silio, L. 2003. Mitochondrial DNA sequence variation and phylogenetic relationships among Iberian pigs and other domestic and wild pig populations. Anim. Genet. 34(5):319¿24. https://doi.org/10.1046/j.1365-2052.2003.01010.x PMID: 14510666 Alves, P.C., Harris, D.J., Melo-Ferreira, J., Branco, M., Ferrand, N., Suchentrunk, F., MeloFerreira, J., Boursot, P. 2006. Hares on thin ice: introgression of mitochondrial DNA in hares and its implications for recent phylogenetic analyses. Mol. Phylogenet. Evol. 40:640¿641. https://doi.org/10.1016/j.ympev.2006.02.016 PMID: 16624594 Alves, P.C., Melo-Ferreira, J., Freitas, H., Boursot, P. 2008. The ubiquitous mountain hare mitochondria: multiple introgressive hybridization in hares, genus Lepus. Phil. Trans. R. Soc. B. 363:2831¿2839. https://doi.org/10.1098/rstb.2008.0053 PMC2606744 PMID: 18508749 Amills, M., Megens, H., Manunza, A., Ramos-Onsins, S., Groenen, M. 2017. A Genomic Perspective on Wild Boar Demography and Evolution. In: Ecology, Conservation and Management of Wild Pigs and Peccaries (Melletti, M., Meijaard, E., eds), pp. 376-387. Cambridge: Cambridge University Press. https://doi.org/10.1017/9781316941232.036 Amills, M., Ramírez, O., Galman-Omitogun, O. Clop, A. 2012. Domestic pigs in Africa. African Archaeological Review. 30:73¿82. https://doi.org/10.1007/s10437-012-9111-2 Anderung, C., Bouwman, A., Persson, P., Carretero, J.M., Ortega, AI., Elburg, R., et al. 2005. Prehistoric contacts over the Straits of Gibraltar indicated by genetic analysis of Iberian Bronze Age cattle. Proc. Natl. Acad. Sci. United. States. Am102(24):8431¿5. https://doi.org/10.1073/pnas. 0503396102 PMC1150856 PMID: 15941827 Ansell, W.F. 1971. Order Artiodactyla. In: The mammals of Africa: An identification manual (Meester, J., Setzer, H.W., eds), pp. 1-84 [issued 1974]. Smithsonian Institution Press, Washington, D.C. Arribas, P. 2013. Ecología evolutiva, biogeografía y conservación de los coleópteros acuáticos en ecosistemas salinos del mediterráneo. Thesis. Spain: University of Murcia. Available from: https://digitum.um.es/digitum/handle/10201/35938 Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., Ferrusquía-Villafranca, I. 2009. A perspective on mammal biodiversity and zoogeography in the Late Pleistocene of México. Quaternary Int. 212(2):187-197. https://doi.org/10.1016/j.quaint.2009.05.012 Asher, R.J., Meng, J., Wible, J.R., McKenna, M.C., Rougier, G.W., Dashzeveg, D., et al. 2005. Stem Lagomorpha and the antiquity of Glires. Science. 307:1091-1094. https//doi.org/10.1126/science.1107808 PMID: 15718468 Aulagnier, S., Cuzin, F., Thévenot, M. 2017. Mammifères Sauvages du Maroc: Peuplement, Répartition, Ecologie. 1st ed. Société Française pour l¿Étude et la Protection des mammifères. Austin, J.D., Lougheed, S.C., Moler, P.E., Boag, P.T. 2003. Phylogenetics, zoogeography, and the role of dispersal and vicariance in the evolution of the Rana catesbeiana (Anura: Ranidae) species group. Biol. J. Linn. Soc. 80:601¿624. https://doi.org/10.1111/j.1095-8312.2003.00259.x Awadi, A., Suchentrunk, F., Makni, M., Ben Slimen, H. 2016. Variation of partial transferrin sequences and phylogenetic relationships among hares (Lepus capensis, Lagomorpha) from Tunisia. Genetica. 144:497. https://doi.org/ 10.1007/s10709-016-9916-z PMID: 27485731 Bandelt, H.J., Forster, P., Röhl, A. 1999. Median-joining networks for inferring intraspecific phylogenies. Mol. Biol. Evol. 16(1):37¿48. https://doi.org/10.1093/oxfordjournals.molbev.a026036 PMID: 10331250 Barata, M., Perera, A., Martínez-Freiría, F., Harris, D.J. 2012. Cryptic diversity within the Moroccan endemic day geckos Quedenfeldtia (Squamata: Gekkonidae): a multidisciplinary approach using genetic, morphological and ecological data. Biol. J. Linn. Soc. Lond. 106:828¿850. https://doi.org/10.1111/j.1095-8312.2012.01903.x Barton, N., Hewitt, G. 1989. Adaptation, speciation and hybrid zones. Nature. 341:497¿503. https://doi.org/10.1038/341497a0 Barratt, E.M., Deaville, R., Burland, T.M., Bruford, M.W., Jones, G., Racey, P.A., et al. 1997. DNA answers the call of pipistrelle bat species. Nature. 387(6629):138¿139. https://doi.org/10.1038/387138b0 PMID: 9144281 Ben Slimen, H. 2008a. Phylogénie morphologique et moléculaire des lièvres d¿Afrique du Nord du genre Lepus. Thesis. Tunisia: Faculty of Sciences of Tunisia. Ben Slimen, H., Schaschl, H., Knauer, F., Suchentrunk, F. 2017. Selection on the mitochondrial ATP synthase 6 and the NADH dehydrogenase 2 genes in hares (Lepus capensis L., 1758) from a steep ecological gradient in North Africa. BMC. Evol. Biol. 17:46. https://doi.org/10.1186/s12862-017-0896-0 PMC5297179 PMID: 28173765 Ben Slimen, H., Suchentrunk, F., Benammar, A. 2008b. On shortcomings of using mtDNA sequence divergence for the systematics of hares (genus Lepus): an example from cape hares. Mamm. Biol. 73:25¿39. https://doi.org/10.1016/j.mambio.2007.02.003 Ben Slimen, H., Suchentrunk, F., Memmi, A., Benammar, A. 2005. Biochemical genetic relationships among Tunisian hares (Lepus sp.), South African cape hares (L. capensis), and European brown hares (L. europaeus). Biochem. Genet. 43:577¿ 596. https://doi.org/10.1007/s10528-005-9115-6 PMID: 16382363 Ben Slimen, H., Suchentrunk, F., Memmi, A., Sert, H., Kruyger, U., Alves, P.C., et al. 2006. Evolutionary relationships among hares from North Africa (Lepus sp. or Lepus spp.), cape hares (L. capensis) from South Africa, and brown hares (L. europaeus), as inferred from mtDNA PCR-RFLP and allozyme data. J. Zool. Syst. Evol. Res. 44:88¿99. https://doi.org/10.1111/j.1439-0469.2005.00345.x Ben Slimen, H., Suchentrunk, F., Shahin, A.B., Benammar, A. 2007. Phylogenetic analysis of mtCR-1 sequences of Tunisian and Egyptian hares (Lepus sp. or spp., Lagomorpha) with different coat colours. Mamm. Biol. 72:224¿239. https://doi.org/10.1016/j.mambio.2006.03.002 Ben Slimen, H., Suchentrunk, F., Stamatis, C., Mamuris, Z., Sert, H., Alves, P.C. et al. 2008c. Population genetics of cape and brown hares (Lepus capensis and L. europaeus): a test of Petter¿s hypothesis of conspeci¿city. Biochem. Syst. Ecol. 36:22¿39. https://doi.org/10.1016/j.bse.2007.06.014 Benabid, A. 2000. Flore et écosystemes du Maroc. Evaluation et préservation de la biodiversité. Ibis Press, Rabat. Benabid, A., Ruiz, A., Sanjuán, M.E., Sainz, H., Simón, J.C. 2015. UICN. Libro rojo de los ecosistemas forestales de Marruecos. Gland, Suiza y Málaga, España: UICN. Available from: http://www.iucn-redlistecosystems-morocco.org/data/pdf/lref_marruecos.pdf Benson, D.A., Karsch-Mizrachi, I., Clark, K., Lipman, D.J., Ostell, J., Sayers, E.W. 2012. GenBank. Nucleic. Acids. Res. Oxford University Press, pp. D48¿53. https://doi.org/10.1093/nar/gkr1202 PMC3245039 PMID: 22144687 Berthier, P., Excoffier, L., Ruedi, M. 2006. Recurrent replacement of mtDNA and cryptic hybridization between two sibling bat species Myotis myotis and Myotis blythii. Proc. Biol. Sci. 273(1605):3101-9. https://doi.org/10.1098/rspb.2006.3680 PMC1679893 PMID: 17018432 Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K.L., Meier, R., Winker, K., et al. 2007. Cryptic species as a window on diversity and conservation. Trends. Ecol. Biol. 22(3):148¿155. https://doi.org/10.1016/j.tree.2006.11.004 PMID: 17129636 Boratynski, Z., Brito, J.C., Mappes, T. 2012. The origin of two cryptic species of African desert jerboas (Dipodidae: Jaculus). Biol. J. Linn. Soc. 105:435¿445. https://doi.org/10.1111/j.1095-8312.2011.01791.x Blench, R.M. 1993. Ethnographic and linguistic evidence for the prehistory of African rumiinant livestock, horses and ponies. In: The archaelogy of Africa. Food, metals and towns (Shaw, T., Sinclair, P., Andah, B., Okpoko, A., eds), pp. 71-103. London: Routledge. Blench, R.M. 1995. A history of domestic animals in northeastern Nigeria. Cahiers de Siences Humaine. ORSTOM. 31(1):181-238. Blench, R.M. 1999. A history of pigs in Africa. In: Origins and development of African livestock: archaeology, genetics, linguistics and ethnography (Blench, R.M., Mac Donald, K., eds), pp. 355¿367. Florence, KY: Routledge Books. Briedermann, L. 1970. Zum Körper- und Organwachstum des Wildschweines in der DDR. Arch. Forstwes. 19(4):401-420. Briedermann, L. 1990. Schwarzwild. VEB Deutscher Landwirtschaftsverlag, Berlin: pp 540. Brito, J.C., Godinho, R., Martinez-Freiria, F., Pleguezuelos, J.M., Rebelo, H., Santos, X., et al. 2014. Unravelling biodiversity, evolution and threats to conservation in the Sahara-Sahel. Biol. Rev. 89:215¿231. https://doi.org/10.1111/brv.12049 PMID: 23848599 Brown, W.M., George, M., Wilson, A.C. 1979. Rapid evolution of animal mitochondrial DNA. Proc. Natl. Acad. Sci. 76(4):1967¿71. PMC383514 PMID: 109836 Brumm, A., Jensen, G.M., van den Bergh, G.D., Morwood, M.J., Kurniawan, I., Aziz, F., et al. 2010. Hominins on Flores, Indonesia, by one million years ago. Nature. 464(7289):748¿52. https://doi.org/10.1038/ nature08844 PMID: 20237472 Cabrera, A. 1932. Los mamíferos de Marruecos. Serie Zoológica, 57. Junta para la Ampliación de Estudios e Investigaciones Científicas. Madrid, España. Castella, V., Ruedi, M., Excoffier, L., Ibáñez, C., Arlettaz, R., Hausser, J. 2000. Is the Gibraltar Strait a barrier to gene flow for the bat Myotis myotis (Chiroptera: Vespertilionidae)? Mol Ecol. 9(11):1761¿72. https:// doi.org/10.1046/j.1365-294x.2000.01069.x PMID: 11091312 Chen, K., Baxter, T., Muir, W.M., Groenen, M.A., Schook, L.B. 2007. Genetic Resources, Genome Mapping and Evolutionary Genomics of the Pig (Sus scrofa). Int. J. Biol. Sci. 3(3):153-65. https://doi.org/10.7150/ijbs.3.153 PMC1802013 PMID: 17384734 Cliffe, K.M., Day, A.E., Bagga, M., Siggens, K., Quilter, C.R., Lowden, S., et al. 2010. Analysis of the non-recombining Y chromosome defines polymorphisms in domestic pig breeds: ancestral bases identified by comparative sequencing. Anim. Genet. 41(6):619¿29. https://doi.org/10.1111/j.1365-2052.2010. 02070.x PMID: 20477804 Clop, A., Amills, M., Noguera, J.L., Fernández, A., Capote, J., Ramón, M.M., et al. 2004. Estimating the frequency of Asian cytochrome B haplotypes in standard European and local Spanish pig breeds. Genet. Sel. Evol. 36(1):97¿104. https://doi.org/10.1186/1297-9686-36-1-97 PMC2697182 PMID: 14713412 Cloudsley-Thompson, J.L. 1991. Ecophysiology of desert arthropods and reptiles. Springer Verlag GmbH & Co. Coe, M.J., Skinner, J.D. 1993. Connections, disjunctions and endemism in the Eastern and southern African mammal faunas. Trans. Roy. Soc. S. Afr. 48:233-255. https://doi.org/10.1080/00359199309520273 Cooke, H.B.S., Wilkinson, A.F. 1978. Suidae and Tayassuidae. In: Evolution of African mammals (Maglio, V.J., Cooke, H.B.S., eds), pp. 435-482. Cambridge, MA: Harvard University Press. Corbet, G.B. 1986. Relationships and origins of European lagomorphs. Mammal Rev. 16:105¿110. https://doi.org/10.1111/j.1365-2907.1986.tb00029.x Cuzin, F. 2003. Les grands Mammifères du Maroc méridional (Haut Atlas, Anti Atlas et Sahara): Distribution, écologie et conservation. Montpellier, France: EPHE. Cuzin, F., Randi, E. 2013. Sus scrofa wild boar (Eurasian wild pig). In: Mammals of Africa. Volume VI. Pigs, hippopotamuses, chevrotain, giraffes, deer and bovids (Kingdon, J., Hoffmann, M., eds), pp. 28-31. Bloomsbury Publ., London. Darriba, D., Taboada, G.L., Doallo, R., Posada, D. 2012. jModelTest 2: more models, new heuristics and high-performance computing. Nat. Methods. 9(8):772. https://doi.org/10.1038/ nmeth.2109 PMC4594756 PMID: 22847109 Dawson, M.R. 1981. Evolution of modern leporids. In: Proceedings of the World Lagomorph Conference (Myers, K., MacInnes, C.D., eds), pp. 1-8. University of Guelph Press, Ontario. de Winton, W.E. 1898. XXV. On the hares of Western Europe and North Africa. Ann. Mag. Nat. Hist. 1:149¿158. https://doi.org/10.1080/00222939808677945 Demenocal, P.B. 1995. Plio-Pleistocene African climate. Science. 270:53-59. https://doi.org/10.1126/xcience.270.5233.53 Diong, C.H. 1973. Studies of the Malayan wild pig in Parak and Johore. Malayan Nature Journal 26: 120151. Dobson, M. 1998. Mammal distributions in the western Mediterranean: the role of human intervention. Mamm. Rev. 28(2):77¿88. https://doi.org/10.1046/j.1365-2907.1998.00027.x Dodson, J.J., Aubin-Horth, N. Thériault, V., Páez, D.J. 2013. The evolutionary ecology of alternative migratory tactics in salmonid fishes. Biol. Rev. Camb. Philos. Soc. 88(3):602-25. https://doi.org/10.1111/brv.12019. PMID: 23347290 Douady, C.J., Catzeflis, F., Raman, J., Springer, M.S., Stanhope, M.J. 2003. The Sahara as a vicariant agent, and the role of Miocene climatic events, in the diversification of the mammalian order Macroscelidea (elephant shrews). Proc. Natl. Acad. Sci. 100(14):8325¿8330. https://doi.org/10.1073/pnas.0832467100 PMC166228 PMID:12821774 Drake, N.A., Blench, R.M., Armitage, S.J., Bristow, C.S., White, K.H. 2011. Ancient watercourses and biogeography of the Sahara explain the peopling of the desert. Proc. Natl. Acad. Sci. USA. 108(2):458-462. https://doi.org/10.1073/pnas.1012231108 PMC3021035 PMID: 21187416 Drummond, A.J. HoS, Y., Phillips, M.J., Rambaut, A. 2006. Relaxed phylogenetics and dating with confidence. PLoS Biol. 4(5): e88. https://doi.org/10.1371/journal.pbio.0040088 PMC1395354 PMID: 16683862 Drummond, A.J., Suchard, M.A., Xie, D., Rambaut, A. 2012. Bayesian Phylogenetics with BEAUti and the BEAST 1.7. Mol. Biol. Evol. 29(8):1969¿73. https://doi.org/10.1093/molbev/ mss075 PMC3408070 PMID: 22367748 Elmastour, A. 1982. Contribution à l¿étude de la biologie et du mode de vie du sanglier (Sus scrofa barbarus, Sclater), Mém. 3ème Cycle, Inst. Agro. Vét. Hassan II, Rabat. pp 45. Elmastour, A., Perhutis, R., Popesco, C.P. 1983. Recherches préliminaires sur la bioloogie et l¿éco-éthologie du Sanglier marocain Sus scrofa barbarus. Symp. Intn. Gest. Conserv. Faune Sauvage Médit., Fès., pp 35. Emberger, L. 1955. Une classification biogéographique des climats. Recherches et Travaux des Laboratoires de Géologie, Botanique et Zoologie, Faculté des Sciences, Univ., Montpellier, France, pp. 1¿43. Epstein, J., Bichard, M. 1984. In: Evolution of Domesticated Animals (Mason, I.L., ed), pp. 145¿162. Longman, New York. Esmaeili, H.R., Coad, B.W., Mehraban, H.R., Masoudi, M., Khaefi, R., Abbasi, K., et al. 2014. An updated checklist of fishes of the Caspian Sea basin of Iran with a note on their zoogeography. Iran. J. Ichthyol. 1(3):152¿184. https://dx.doi.org/10.22034/iji.v1i3.18 Ewer, R.F., Cooke, H.B.S. 1964. The Pleistocene mammals of southern Africa. In: Ecological studies in southern Africa (Davis, D.H.S., DeMeillon, B., Harington, J.S., Kalk, M., eds), pp. 35-48. W. Junk Publishers, The Hague. Ezzine, H., Bouziane, A., Ouazar, D., Hasnaoui, M.D. 2017. Sensitivity of NDVI-Based Spatial Downscaling Technique of Coarse Precipitation to Some Mediterranean Bioclimatic Stages. IEEE Geosci. Remote Sens. Lett. 14(9):1518-1521. https://doi.org/10.1109/LGRS.2017.2720166 Fang, M., Andersson, L. 2006. Mitochondrial diversity in European and Chinese pigs is consistent with population expansions that occurred prior to domestication. Proc. R. Soc. B. Biol. Sci. 273(1595):1803¿10. https://doi.org/10.1098/rspb.2006.3514 PMC1634785 PMID: 16790414 Fedorov, V.B., Stenseth, N.C. 2001. Glacial survival of the Norwegian lemming (Lemmus lemmus) in Scandinavia: inference from mitochondrial DNA variation. Proc. Biol. Sci. 268(1469):809-814. https://doi.org/10.1098/rspb.2001.1584 PMC1088673 PMID: 11345325 Fernández, AI., Alves, E., Óvilo, C., Rodríguez, M.C., Silió, L. 2011. Divergence time estimates of East Asian and European pigs based on multiple near complete mitochondrial DNA sequences. Anim. Genet. 42(1):86¿8. https://doi.org/10.1111/j.1365-2052.2010.02068.x PMID: 20477794 Ferreira, E., Souto, L., Soares, A., Fonseca, C. 2009. Genetic structure of the wild boar population in Portugal: evidence of a recent bottleneck. Mammal. Biol. 74:274-285. https://doi.org/10.1016/j.mambio.2008.05.009 Fickel, J., Hauffe, H.C. Pecchioli, E., Soriguer, R., Vapa, L., Pitra, C. 2008. Cladogenesis of the European Brown hare (Lepus europaeus Pallas, 1778). Eur. J. Wildl. Res. 54:495. https://doi.org/10.1007/s10344-008-0175-x Flynn, L., Winkler, A., Erbajeva, M., Alexeeva, N., Anders, U., Angelone, C., et al. 2014. The Leporid Datum: A late Miocene biotic marker. Mamm. Rev. 2014:164¿76. https://doi.org/10.1111/mam.12016 Flux, J.E.C. 1983. Introduction to taxonomic problems in hares. Acta Zool. Fennica, 174:7¿ 10. Flux, J.E.C., Angemann, R. 1990. The hares and jackrabbits. In: Rabbits, Hares and Pikas: status conservation Action Plan. IUCN resources. (Chapmann, J.A., Flux, J.E.C., eds), pp. 61¿94. Gland, Swiitzerland. Frankham, R., Ballou, J.D., Briscoe, D.A. 2004. A primer for conservation genetics. Cambridge University Press, Cambridge. Futuyma, D.J. 2009. Evolution. Sinauer, Sunderland. Frantz, L., Madsen, O., Megens, H.-J., Schraiber, J., Paudel, Y., Bosse, M., et al. 2015- Evolution of Tibetan Wild Boars. Nat. Genet. 47(3):188¿189. https://doi.org/10.1038/ng.3197 PMC4455000 PMID: 25711859 Frantz, L., Meijaard, E., Gongora, J., Haile, J., Groenen, M., Larson, G. 2016. The Evolution of Suidae. Annu. Rev. Anim. Biosci. 4:61-85. https://doi.org/10.1146/annurev-animal-021815-111155 PMID: 26526544 Frantz, L., Schraiber, J., Madsen, O., Megens, H.-J., Bosse, M., Paudel, Y., et al. 2013. ¿ Genome sequencing reveals fi ne scale diversifi cation and reticulation history during speciation in Sus. Genom. Biol. 14 (9):1-12. http://dx.doi.org/10.1186%2Fgb2013-14-9-r107 PMC4053821 PMDID: 24070215 Frantz, L., Schraiber, J., Madsen, O., Megens, H.-J., Cagan, A., Bosse, M., et al. 2015. Evidence of long-term gene flow and selection during domestication from analyses of Eurasian wild and domestic pig genomes. Nat. Genet. 47(10):1141-8. https://doi.org/10.1038/ng.3394 PMID: 26323058 Gantenbein, B., Largiadèr, C.R. 2003. The phylogeographic importance of the Strait of Gibraltar as a gene flow barrier in terrestrial arthropods: a case study with the scorpion Buthus occitanus as model organism. Mol. Phyl. Evol. 28(1):119¿30. https://doi.org/10.1016/S1055-7903(03)00031-9 PMID: 12801475 García-Mudarra, J., Ibáñez, C., Juste, J. 2009. The Straits of Gibraltar: barrier or bridge to Ibero-Moroccan bat diversity? Biol. J. Linn. Soc. 96(2):434¿50. https://doi.org/10.1111/j.1095-8312.2008.01128.x Garzón, P. 1991. Biología y ecología del jabalí (Sus scrofa L., 1758) en el Parque Natural de Monfragüe. Thesis. Spain: Universidad Autónoma de Madrid. Available from: https://repositorio.uam.es/handle/10486/666215 Gharaibeh, B.M. 1997. Systematics, distribution, and zoogeography of mammals of Tunisia. Thesis. Graduate Faculty of Texas Tech University. Available from: https://ttu-ir.tdl.org/ttu-ir/handle/ 2346/9796 Gibert, J., Gibert, L., Iglesias, A., Maestro, E. 1998. Two ¿Oldowan¿ assemblages in the Plio-Pleistocene deposits of the Orce region, southeast Spain. Antiquity. 72(275):17¿25. https://doi.org/10.1017/ S0003598X00086233 Gibert, J., Gibert, L., Iglesias, A. 2003. The Gibraltar Strait: A Pleistocene door of Europe? Hum. Evol. 18(3):147¿60. https://doi.org/10.1007/BF02436283 Gibert, L., Scott, G.R., Scholz, D., Budsky, A., Ferràndez, C., Ribot, F., et al. 2016. Chronology for the Cueva Victoria fossil site (SE Spain): Evidence for Early Pleistocene Afro-Iberian dispersals. J. Hum. Evol. 90:183¿97. https://doi.org/10.1016/j.jhevol.2015.08.002 PMID: 26581114 Gilman, A. 1975. The later prehistory cultures of East Africa: an historical inquirí. Thesis. Anthropology, Michigan: Ann Arbor. Giuffra, E., Kijas, J.M., Amarger, V., Carlborg, O., Jeon, J.T., Andersson, L. 2000. The origin of the domestic pig: independent domestication and subsequent introgression. Genetics. 154(4):1785¿91. PMC1461048 PMID: 10747069 Goodman, S. M., Ramasindrazana, B. 2015. Description of a new species of the Miniopterus aelleni group (Chiroptera: Miniopteridae) from upland areas of central and northern Madagascar. Zootaxa. 3936(4):538¿558. http://dx.doi.org/10.11646/zootaxa.3936.4.4 PMID: 25947452 Gonçalves, D.V., Brito, J.C., Crochet, P.-A., Geniez, P., Padial, J.M., Harris, D.J. 2012. Phylogeny of North African Agama lizards (Reptilia: Agamidae) and the role of the Sahara desert in vertebrate speciation. Mol. Phylogenet. Evol. 64(3):582¿591. https://doi.org/10.1016/j.ympev.2012.05.007 PMID: 22634241 Gonçalves, D.V., Martínez-Freiría, F., Crochet, P.-A., Geniez, P., Carranza, S., Brito, J.C. 2018. The role of climatic cycles and trans-Saharan migration corridors in species diversification: Biogeography of Psammophis schokari group in North Africa. Mol. Phylogenet. Evol. 118:64¿74. Graciá, E., Giménez, A., Anadón, J.D., Harris, D.J., Fritz, U., Botella, F. 2013. The uncertainty of Late Pleistocene range expansions in the western Mediterranean: a case study of the colonization of south-eastern Spain by the spur-thighed tortoise, Testudo graeca. J. Biogeogr. 40(2):323¿34. https://doi.org/10. 1111/jbi.12012 Grant, W.S., Leslie, R.W. 1993. Effect of metapopulatiom structure and organellar DNA variability in semi-arid environments of southern Africa. S. Afr. J. Sci. 89:287-293. Groves, C.P. 1981. Ancestors for the Pigs: Taxonomy and Phylogeny of the genus Sus. Tech. Bull. N3, Australian National University Press, Canberra: pp 96. Groves, C.P., Grubb, P. 1993. The Eurasian suids (Sus and Babyrousa). Taxonomy and description. In: Pigs, Peccaries, and Hippos: Status Survey and Conservation Action Plan (Oliver, W.L.R., ed), pp. 107-111. International Union for the Conservation of Nature, Gland, Switzerland. Grubb, P. 1978. Patterns of speciation in African mammals. In: Ecology and taxonomy of African small mammals (Schlitter, D.A., ed), pp. 1-26. Bulletin of Carnegie Museum of Natural History. Number 6, Pittsburgh. Guinea, E. 1945. Aspecto forestal del desierto. La vegetación leñosa y los pastos del Sahara español. Madrid, Instituto Forestal de Investigaciones y Experiencias. Available from: https://bibdigital.rjb.csic.es/viewer/10160/?offset=#page=1&viewer=picture&o=bookmark&n=0&q= Guinea, E. 1948. Catálogo razonado de las plantas del Sahara español. An. Jard. Bot. Madrid, 8:357-442. Guzmán, B., Vargas, P. 2005. Systematics, character evolution, and biogeography of Cistus L. (Cistaceae) based on ITS, trnl-trnF, and matK sequences. Mol. Phylogenet. Evol. 37:644¿660. https://doi.org/10.1016/j.ympev.2005.04.026 PMID: 16055353 Habel, J.C., Dieker, P., Schmitt, T. 2009. Biogeographical connections between the Maghreb and the Mediterranean peninsulas of southern Europe. Biol. J. Linn. Soc. 98:693¿703. Habel, J.C., Rödder, D., Stefano, S., Meyer, M., Schmitt, T. 2010. Strong genetic cohesiveness between Italy and North Africa in four butterfly species. Biol. J. Linn. 99(4):818¿30. https://doi.org/10.1111/j. 1095-8312.2010.01394.x Haberle, S., Stevenson, J., Prebble, M. 2010. Altered Ecologies: Fire, Climate and Human Influence on Terrestrial Landscapes. ANU E Press. Australia. Hajji, G.E.M., Zachos, F.E. 2011. Mitochondrial and nuclear DNA analyses reveal pronounced genetic structuring in Tunisian wild boar Sus scrofa. Eur. J. Wildl. Res. 57(3):449¿56. https://doi.org/10.1007/s10344010-0452-3 Halanych, K.M., Robinson, T.J. 1999. Multiple substitutions affect the phylogenetic utility of Cytochrome b and 12S rDNA data: examining a rapid radiation in leporid (Lagomorpha) evolution. J. Mol. Evol. 48:369¿379. https://doi.org/10.1007/PL00006481 PMID: 10093227 Hall, T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl. Acids. Symp. Ser. 41:95¿98. Hernández, C.L., Soares, P., Dugoujon, J.M., Novelletto, A., Rodríguez, J.N., Rito, T., et al. 2015. Early Holocenic and Historic mtDNA African Signatures in the Iberian Peninsula: The Andalusian Region as a Paradigm. PLoS One. 10(10):1¿24. https://doi.org/10.1371/journal.pone.0139784 PMC4624789 PMID: 26509580 Hernández-Pacheco, E., Hernández-Pacheco, F., Alía Medina, M., Vidal Box, C., Guinea, E. 1949. El Sahara español. Estudio geológico, geográfico y botánico. Madrid, Instituto de Estudios Africanos. Hewitt, G.M. 1988. Hybrid zones-natural laboratorios for evolutionary sutides. Trends. Ecol. Evol. 3(7):158-167. https://doi.org/10.1016/0169-5347(88)90033-X. PMID: 21227192 Hibbard, C.W. 1963. The origin of the P3 pattern of Sylvilagus, Caprolagus, Oryctolagus and Lepus. J. Mamm. 44(1):1-15. https://doi.org/10.2307/1377162 Hinojosa, J. C., Koubínová, D., Szenteczki, M. A., Pitteloud, C., Dinc¿, V., Alvarez, N., et al. 2019. A mirage of cryptic species: Genomics uncover striking mitonuclear discordance in the butterfly Thymelicus sylvestris. Mol. Ecol. 28(17):3857¿3868. https://doi.org/10.1111/mec.15153 PMID: 31233646 Hoffman, J.I., Grant, S.M. Forcada, J., Phillips, C.D. 2011. Bayesian inference of a historical bottleneck in a heavily exploited marine mammal. Mol. Ecol. (19):3989-4008. https://doi.org/10.1111/j.1365-294X.2011.05248.x PMID: 21895820 Hoffmann, R. S., Smith, A. T. 2005. Lagomorphs. In: Mammal species of the World. 3rd Ed, (Wilson, D.E., Reeder, D.M., eds). Johns Hopkins University Press. Höglund, J. 2009. Evolutionary conservation genetics. Oxford University Press, Oxford. Hooghiemstra, H., Stalling, H., Agwu, C.O.C., Dupont, L.M. 1992. Vegetational and climatic changes at the northern fringe of the sahara 250,000¿5000 years BP: evidence from 4 marine pollen records located between Portugal and the Canary Islands. Rev. Palaeobot. Palynol. 74(1):1¿53. https://doi.org/10.1016/0034-6667(92)90137-6 Horreo, J.L., Alonso, J.C., Palacin, C., Milá, B. 2014. Genetic structure in Iberian and Moroccan populations of the globally threatened great bustard Otis tarda: a microsatellite perspective. J. Avian. Biol. 45(5):507¿13. https://doi.org/10.1111/jav.00401 Hotaling, S., Foley, M. E., Lawrence, N. M., Bocanegra, J., Blanco, M. B., Rasoloarison, R., et al. 2016. Species discovery and validation in a cryptic radiation of endangered primates: coalescent-based species delimitation in Madagascar¿s mouse lemurs. Mol. Ecol. 25(9):2029¿2045. https://doi.org/10.1111/mec.13604 PMID: 26946180 Hulme-Beaman, A., Cucchi, T., Evin, A., Searle, J. B., Dobney, K. 2018. Exploring Rattus praetor (Rodentia, Muridae) as a possible species complex using geometric morphometrics on dental morphology. Mammal. Biol. 92:62¿67. https://doi.org/10.1016/j.mambio.2018.04.002 Hundertmark, K., Daele, L. 2010. Founder effect and bottleneck signatures in an introduced, insular population of elk. Conserv. Genet. 11:139-147. https://doi.org/10.1007/s10592-009-0013-z Huntley, J.W., Keith, K.D., Castellanos, A.A., Musher, L.J., Voelker, G. 2019. Underestimated and cryptic diversification patterns across Afro-tropical lowland forests. J. Biogeogr. 46(2):381¿391. https://doi.org/10.1111/jbi.13505 Husemann, M., Schmitt, T., Zachos, F.E., Ulrich, W., Habel, J.C. 2014. Palaearctic biogeography revisited: evidence for the existence of a North African refugium for Western Palaearctic biota. J. Biogeogr. 41(1):81¿94. https://doi.org/10.1111/jbi.12180 Iwase, M., Satta, Y., Hirai, Y., Hirai, H., Imai, H., Takahata, N. 2003. The amelogenin loci span an ancient pseudoautosomal boundary in diverse mammalian species. Proc. Natl. Acad. Sci. U. S. A. 100(9):5258¿63. https://doi.org/10.1073/pnas.0635848100 PMC154332 PMID: 12672962 Jansson, G., Thulin, C.G. and Pehrson, A. 2007. Factors related to the occurrence of hybrids between brown hares Lepus europaeus and mountains hares L. timidus in Sweden. Ecography. 30:709-715. https://doi.org/10.1111/j.2007.0906-7590.05162.x Jing, Y., Flad, R.K. 2002. Pig domestication in Ancient China. Antiquity. 76(293):724-732. Juste, J., Bilgin, R., Munoz, J., Ibañez, C. 2009. Mitochondrial DNA signatures at different spatial scales: from the effects of the Straits of Gibraltar to population structure in the meridional serotine bat (Eptesicus isabellinus). Heredity (Edinb). 103(2):178¿87. https://doi.org/10.1038/hdy.2009.47 PMID: 19401715 Juste, J., Ruedi, M., Puechmaille, S. J., Salicini, I., Ibáñez, C. 2019. Two New Cryptic Bat Species within the Myotis nattereri Species Complex (Vespertilionidae, Chiroptera) from the Western Palaearctic. Acta Chiropterologica. 20(2):285¿300. https://doi.org/10.3161/15081109ACC2018.20.2.001 Kasapidis, P., Suchentrunk, F., Magoulas, A., Kotoulas, G. 2005. The shaping of mitochondrial DNA phylogeographic patterns of the brown hare (Lepus europaeus) under the combined in¿uence of Late Pleistocene climatic ¿uctuactions and antrhopogenic translocations. Mol. Phylogenet. Evol. 34:55¿66. https://doi.org/10.1016/j.ympev.2004.09.007 PMID: 15579381 Kawamoto, Y., Shotake T., Nozawa, K., Kawamoto, S., Tomari, K, Kawai, S. 2007. Postglacial population expansion of Japanese macaques (Macaca fuscata) inferred from mitochondrial DNA phylogeography. Primates. 48:27-40. https://doi.org/10.1007/s10329-006-0013-2 Keuling, O., Leus, K. 2019. Sus scrofa. The IUCN Red List of Threatened Species 2019: e.T41775A44141833. Available from: http://dx.doi.org/10.2305/IUCN.UK.2019-3.RLTS.T41775A44141833.en Kijas, J.M., Andersson, L. 2001. A phylogenetic study of the origin of the domestic pig estimated from the near-complete mtDNA genome. J. Mol. Evol. 52(3):302-308. https://doi.org/10.1007/s002390010158 PMID: 11428467 Kim, K.-I., Lee, J.-H., Li, K., Zhang, Y.-P., Lee, S.-S., Gongora, J., et al. 2002. Phylogenetic relationships of Asian and European pig breeds determined by mitochondrial DNA D-loop sequence polymorphism. Anim. Genet. 19¿25. https://doi.org/10.1046/j.1365-2052.2002.00784.x PMID: 11849133 Kim, K.-I., Yang, Y.-H., Lee, S.-S., Park, C., Ma, R., Bouzat, J.L., et al. 1999. Phylogenetic relationships of Cheju horses to other horse breeds as determined by mtDNA D-loop sequence polymorphism. Anim. Genet. 30(2):102¿8. https://doi.org/10.1046/j.1365-2052.1999.00419.x PMID: 10376300 Kingdon, J., Happold, D., Butynski, T.M., Hoffmann, M., Happold, M., Kalina, J. 2013. Mammals of Africa Volume I. A & C Black. Kishida, O., trussell, G.C., Mougi, A., Nishimura, K. 2009. Popul. Ecol. 52:37-46. https://doi.org/10.1007/s10144-009-0182-0 Klicka, J., Zink, R.M. 1997. The importance of recent ice ages in speciation: a failed paradigm. Science. 277(5332):1666-1669. https://doi.org/10.1126/science.277.5332.1666 Koepfli, K.P., Pollinger, J., Godinho, R., Robinson, J., Lea, A., Hendricks, S. et al. 2015. Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species. Curr. Boil. 25(16):2158-65. https://doi.org/10.1016/j.cub.2015.06.060 PMID: 26234211 Kong, L., Wang, W., Cong, H., Son Nguyen, T., Yang, Q., Wu, Y., et al. 2014. Molecular evidence revealed Lepus hainanus and L. peguensis have a conspecific relationship. Mitochondrial DNA. 27(1):265-9. https://doi.org/10.3109/19401736.2014.888550 PMID: 24548010 Koju, N.P., He, K., Chalise, M.K., Ray, C., Chen, Z., Zhang, B., et al. 2017. Multilocus approaches reveal underestimated species diversity and inter-specific gene flow in pikas (Ochotona) from southwestern China. Mol. Phylo. Evol. 107:239¿245. https://doi.org/10.1016/j.ympev.2016.11.005 PMID: 27838310 Kottek, M., Grieser, J., Beck, C., Rudolf, B., Rubel, F. 2006. World Map of the Köppen-Geiger Climate Classification Updated. Meteorological Zeitschrift. 15(3):259-263. https://doi.org/10.1127/0941-2948/ 2006/0130 Kryger, U. 2005. Genetic variation among South African hares (Lepus spec.) as inferred from mitocondrial DNA and microsatellites. Thesis. University of Petroria. Available from: https://repository.up.ac.za/handle/2263/25948 Kumar, S., Stecher, G., Tamura, K. 2016. MEGA7: Molecular Evolutionary Genetics Analysis Version 7.0 for Bigger Datasets. Mol. Biol. Evol. 33(7):1870¿4. Available from: https://doi.org/10.1093/molbev/msw054 Kuro-o, M., Yonekawa, H., Saito, S., Eda, M., Higuchi, H., Koike, et al. 2010. Unexpectedly high genetic diversity of mtDNA control región through severe bottleneck in culnerable albatros Phoebastria albatrus. Conserv. Genet. 11:127-137. https://doi.org/10.1007/s10592-009-0011-1 Ladle, R.J., Whittaker, M.S, eds. 2011. Conservation biogeography. Wiley-Blackwell, Chichester. Lado, S. 2015. Population history and taxonomy of African hares (genus Lepus) inferred from genetic variation. Thesis. Universidade du Porto. Lado, S., Alves, P.C., Islam, M.Z., Brito, J.C., Melo-Ferreira, J. 2019. The evolutionary history of the Cape hare (Lepus capensis sensu lato): insights for systematics and biogeography. Heredity (Edinb). 123(5):634¿46. Larson, G., Dobney, K., Albarella, U., Fang, M., Matisoo-Smith, E., Robins, J., et al. 2005. Worldwide Phylogeography of Wild Boar Reveals Multiple Centers of Pig Domestication. Science. 307(5715):1618¿1621. https://doi.org/10.1126/science.1106927 PMID: 15761152 Lataste, F. 1887. Catalogue critique des mammiferes appelagiques sauvages de la Tunisie. Imprimerie Natinonle, Paris, pp. 42. Lau, J.A., Lennon, J.T. 2011. Evolutionary ecology of plant-microbe interactions: soil microbial structure alters selection on plant traits. New Phytol. 192(1):215-24. https://doi.org/10.1111/j.1469-8137.2011.03790.x PMID: 21658184 Lavergne, S., Hampe, A., Arroyo, J. 2013. In and out of Africa: how did the Strait of Gibraltar affect plant species migration and local diversification? J. Biogeogr. 40:24¿36. https://doi.org/10.1111/j.1365-2699.2012.02769.x Lekagul, B., McNeely, J.A. 1977. Mammals of Thailand. Kurusapha Ladprao Press, Bangkok: pp 661-664. Li, W.H. 1997. Molecular evolution. 1st ed. Massachusetts: Sinauer Associates. Librado, P., Rozas, J. 2009. DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinforma. 25(11):1451¿2. https://doi.org/10.1093/bioinformatics/btp187 PMID: 19346325 Liu, J., Yu, L., Arnold, M.L., Wu, C.H., Wu, S.F., Lu, X. et al. 2011. Reticulate evolution: frequent introgressive hybridization among Chinese hares (genus Lepus) revealed by analysis of multiple mitochondrial and nuclear DNA loci. BMC. Evol. Biol. 11:223¿236. https://doi.org/10.1186/1471-2148-11-223 PMC3155923 PMID: 21794180 López-Wilchis, R., Flores-Romero, M., Guevara-Chumacero, L. M., Serrato-Díaz, A., Díaz-Larrea, J., et al. 2016. Evolutionary Scenarios Associated with the Pteronotus parnellii Cryptic Species-Complex (Chiroptera: Mormoopidae). Acta Chiropterologica. 18(1):91¿116. https://doi.org/10.3161/15081109ACC2016.18.1.004 Lucchini, V., Meijaard, E., Diong, C.H., Groves, C.P., Randi, E. 2006. New phylogenetic perspectives among species of South¿east Asian wild pig (Sus sp.) based on mtDNA sequences and morphometric data. J. Zool. 266:25-35. https://doi.org/10.1017/S0952836905006588 Mace, G.M. 2004. The role of taxonomy in species conservation. Philos. Trans. R. Soc. Lond. B. Biol. Sci. 359(1444):711-719. https://doi.org/10.1098/rstb.2003.1454 Maghnouj, M. 1999. Quelques espèces de mammifères de la cédraie au Maroc, cas du singe Magot. T. XX, n3. pp. 101-105. Mamuris, Z., Motou, K., Stamatis, C., Sarafidou, T., Suchentrunk, F. 2010. Y DNA and mitochondrial lineages in European and Asian populations of the brown hare (Lepus europaeus). Mammal. Biol. 75:233¿242. https://doi.org/10.1016/j.mambio.2009.01.004 Manlius, N., Gautier, A. 1999. The wild boar of Egypt. C. R. Acad. Sci. III. 322(7):573¿7. PMID: 10488431 Marincovich, J.L., Gladenkov, A.Y., 1999. Evidence for an early opening of the Bering Strait. Nature 397:149¿151. https://doi.org/10.1038/16446 Marraha, M., Sehhar, E. 1997. Régime alimentaire du lièvre (Lepus capensis L.) dans la région du Souss. Ann. Rech. For. Maroc. 30:99-11. Maryanto, I., and Higashi, S. 2011. Comparison of zoogeography among rats, fruit bats and insectivorous bats on Indonesian islands. Treubia. 38:33-52. https://doi.org/10.14203/treubia.v38i0.545 Matthiesen, S., Haines, K. 2003. A hydraulic box model study of the Mediterranean response to postglacial sealevel rise. Paleoceanography. 18(4):1¿12. https://doi.org/10.1029/2003PA000880 Matthee, C.A., van Vuuren, B.J., Bell, D., Robinson, T.J. 2004. A molecular supermatrix of the rabbits and hares (Leporidae) allows for the identification of five intercontinental exchanges during the Miocene. Syst. Biol. 53:433¿447. https://doi.org/10.1080/10635150490445715 PMID: 15503672 Mayer, V.E., Frederickson, M.E., McKey, D., Blatrix, R. 2014. New Phytol. 202(3):749-64. https:// doi.org/10.1111/nph.12690 PMID: 24444030 McKenna, M.C. 1982. Lagomorph interrelationships. Geobios Mem. Spec. 6:213-223. https://doi.org/10.1016/S0016-6995(82)80115-0 Mecklenburg, C.W., Moller, P.R., Steinke, D. 2011. Biodiversity of arctic marine fishes: taxonomy and zoogeography. Mar. Biodiv. 41:109¿140. https://doi.org/10.1007/s12526-010-0070-z Meiri, M., Huchon, D., Bar-Oz, G., Boaretto, E., Horwitz, L.K., Maeir, A.M., et al. 2013. Ancient DNA and Population Turnover in Southern Levantine Pigs- Signature of the Sea Peoples Migration? Sci. Rep. 3:3035. https://doi.org/10.1038/srep03035 PMC3816294 PMID: 24186332 Melo-Ferreira, J., Alves, P.C., Freitas, H., Ferrand, N., Boursot, P. 2009. The genomic legacy from the extinct Lepus timidus to the three hare species of Iberia: contrast between mtDNA, sex chromosomes and autosomes. Mol. Ecol. 18:2643¿2658. https://doi.org/10.1111/j.1365-294X.2009.04221.x PMID: 19457181 Melo-Ferreira, J., Alves, P.C., Rocha, J., Ferrand, N., Boursot, P. 2011. Interspeci¿c XChromosome and Mitochondrial DNA Introgression in the Iberian hare: selection or Allele Sur¿ng? Evolution. 65: 1956¿1968. https://doi.org/10.1111/j.1558-5646.2011.01261.x PMID: 21729051 Melo-Ferreira, J., Boursot, P., Carneiro, M., Esteves, P.J., Farelo, L., Alves, P.C. 2012. Recurrent introgression of mitochondrial DNA among hares (Lepus spp.) revealed by species-tree inference and coalescent simulations. Syst. Biol. 61: 367¿381. https://doi.org/10.1093/sysbio/syr114 PMID: 22201159 Melo-Ferreira, J., Boursot, P., Randi, E., Kryukov, A., Suchentrunk, F., Ferrand, N. et al. 2007. The rise and fall of mountain hare (Lepus timidus) during Pleistocene glaciations: expansion and retreat with hybridization in the Iberian Peninsula. Mol. Ecol. 16(3):605-618. https://doi.org/10.1111/j.1365-294X.2006.03166.x PMID: 17257116 Melo-Ferreira, J., Boursot, P., Suchentrunk, F., Ferrand, N., Alves, P.C. 2005. Invasion from the cold past: extensive introgression of mountain hare (Lepus timidus) mitochondrial DNA into three other hare species in northern Iberia. Mol. Ecol. 14:2459¿2464. https://doi.org/10.1111/j.1365-294X.2005.02599.x PMID: 15969727 Melo-Ferreira, J., Farelo, L., Freitas, H., Suchentrunk, F., Boursot, P., Alves, P.C. 2014a. Home-loving boreal hare mitochondria survived several invasions in Iberia: the relative roles of recurrent hybridisation and allele sur¿ng. Heredity. 112: 265¿ 273. https://doi.org/10.1038/hdy.2013.102 PMC3931168 PMID: 24149657 Melo-Ferreira, J., Seixas, F.A., Cheng, E., Mills, S.L., Alves, P.C. 2014b. The hidden history of the snowshoe hare, Lepus americanus: extensive mitochondrial DNA introgression inferred from multilocus genetic variation. Mol. Ecol. 23:4617¿630. https://doi.org/doi:10.1111/mec.12886 PMID: 25113393 Melo-Ferreira, J., Vilela, J., Fonseca, M.M., da Fonseca, R.R., Boursot, F., Alves, P.C. 2014c. The elusive nature of adaptive mitochondrial DNA evolution of an arctic lineage prone to frequent introgression. Genom. Biol. Evol. 6:886¿896. https://doi.org/10.1093/gbe/evu059 PMC4007550 PMID: 24696399 Menni, R.C., Jaureguizar, A.J., Sthemann, M.F.W., Lucifora, L.O. 2009. Marine biodiversity at the community level: zoogeography of sharks, skates, rays and chimaeras in the southwestern Atlantic. Bio. Conserv. 19:775¿796. https://doi.org/10.1007/s10531-009-9734-z Mikolajewicz, U. 2011. Modeling Mediterranean Ocean climate of the Last Glacial Maximum. Clim Past. 7:161¿180. https://doi.org/10.5194/cp-7-161-2011 Molina-Venegas, R., Aparicio, A., Lavergne, S., Arroyo, J. 2015. The building of a biodiversity hotspot across a land-bridge in the Mediterranean. Proc. R. Soc. 282:20151116. http://dx.doi.org/10.1098/rspb.2015.1116 PMC4632623 PMID: 26246551 Mouliéras, A. 1905. Une tribu zènète anti-musulmane au Maroc. Paris. Moura, A.E., Janse van Rensburg, C., Pilot, M., Tehrani, A., Best, P.B., Thorton, M., et al. 2014. Killer whale nuclear genome and mtDNA reveal widespread population bottleneck during the last glacial máximum. Mol. Biol. Evol. 31(5):1121-31. https://doi.org/10.1093/molbev/msu058. PMC3995335 PMID: 24497033 Müller, P. 1974. Aspects of Zoogeography. Springer. Natoli, A., Phillips, K.P., Richardson, D.S., Jabado, R.W. 2017. Low genetic diversity after a bottleneck in a population of a critically endangered migratory marine turtle species. J. Exp. Mar. Biol. Ecol. 491:9-18. https://doi.org/10.1016/j.jembe.2017.01.009 Noce, A., Amills, M., Manunza, A., Muwanika, V., Muhangi, D., Aliro, T., et al. 2015. East African pigs have a complex Indian, Far Eastern and Western ancestry. Anim. Genet. 46(4):433¿6. https://doi.org/10. 1111/age.12305 PMID: 26011180 Ogilvie, H.A., Bouckaert, R.R., Drummond, A.J. 2017. StarBEAST2 Brings Faster Species Tree Inference and Accurate Estimates of Substitution Rates. Mol. Biol. Evol. 34:2101¿2114. https://doi.org/10.1093/molbev/msx126 PMC5850801 PMID: 28431121 Ojeda, A., Rozas, J., Folch, J.M., Pérez-Enciso, M. 2006. Unexpected High Polymorphism at the FABP4 Gene Unveils a Complex History for Pig Populations. Genetics. 174(4):2119¿27. https://doi.org/10.1534/genetics.106.063057 PMC1698616 PMID: 17057239 Olalde, I., Capote, J., Del-Arco, M.C., Atoche, P., Delgado, T., González-Anton, R., et al. 2015. Ancient DNA sheds light on the ancestry of pre-hispanic Canarian pigs. Genet. Sel. Evol. 47(1):1¿5. https://doi.org/10.1186/s12711-015-0115-7 PMC4421913 PMID: 25944642 Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, V.N., Underwood, E.C., et al. 2001. Terrestrial ecoregions of the world: a new map of life on Earth. BioScience. 51(11):933-938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2 Osborne, A.H., Vance, D., Rohling, E.J., Barton, N., Rogerson, M., Fello, N. 2008. A humid corridor across the Sahara for the migration of early modern humans out of Africa 120,000 years ago. Proc. Natl. Acad. Sci. 105(43):16444¿16447. https://doi.org/10.1073/pnas.0804472105 PMC2575439 PMID: 18936490 Padial. J.M., Miralles, A., de la Riva, I., Vences, M. 2010. The integrative future of taxonomy. Front. Zool. 7:16. https://doi.org/10.1186/1742-9994-7-16 PMC2890416 PMID: 20500846 Pala, M., Olivieri, A., Achilli, A., Accetturo, M., Metspalu, E., Reidla, M., et al. 2012. Mitochondrial DNA signals of late glacial recolonization of Europe from near eastern refugia. Am. J. Hum. Genet. 90(5):915-24. https://doi.org/10.1016/j.ajhg.2012.04.003 PMC3376494 PMID: 22560092 Palacios, F. 1989. Biometric and morphologic features of the species of the genus Lepus in Spain. Mammalia, 73:227-264. Palacios, F. 1996. Systematics of the indigenous hares of Italy traditionally identified as Lepus europaeus Pallas, 1778 (Mammalia: Leporidae). Bonn. Zool.Beitr. 46:59¿91. Pauwels, W. 1980. Study of Sus scrofa vittatus, its ecology and behaviour in Ujung Kulon Nature Reserve, Java, Indonesia. Thesis. Switzerland: University of Basel. Peery, M.Z., Kirby, R., Reid, B.N. Stoelting, R., Doucet-Bëer, E., Robinson, S., et al. 2012. Reliability of genetic bottleneck tests for detecting recent population declines. Mol. Ecol. 21(14):3403-18 https://doi.org/10.1111/j.1365-294X.2012.05635.x PMID: 22646281 Pérez-Suárez, G., Palacios, F., Boursot, P. 1994. Speciation and paraphyly in western Mediterranean hares (Lepus castroviejoi, L. europaeus, L. granatensis, and L. capensis) revealed by mitochondrial DNA phylogeny. Biochem. Genet. 32(11-12):423-36. https://doi.org/10.1007/bf00566063 PMID: 7748159 Pesole, G., Gissi, C., De Chirico, A., Saccone, C. 1999. Nucleotide substitution rate of mammalian mitochondrial genomes. J. Mol. Evol. 48(4):427¿434. https://doi.org/10.1007/Pl00006487 PMID: 10079281 Petter, F. 1959. Eléments d¿une révision des lièvres africains du sous-genre Lepus. Mammalia. 23: 41¿ 67. Petter, F. 1961. Eléments d¿une révision des lièvres européens et asiatique du sous-genre Lepus. A. Säugetierkd. 26:30-40. Phoca-Cosmetatou, N., Rabett, R.J. 2014. Pleistocene Island Occupation in the Mediterranean: Insights from a Tied-Biome Approach to Glacial Refugia. In: Living in the Landscape: Essays in Honour of Graeme Barker (Boyle, K., Rabett, R.J., Hunt, C.O., eds), pp. 83¿108. Cambridge: McDonald Institute for Archaeological Research, McDonald Institute Monograph Series. Pierpaoli, M., Trocchi, V., Randi, E. 1999. Species distinction and evolutionary relationships of the Italian hare (Lepus corsicanus) as described by mitochondrial DNA sequencing. Mol. Ecol. 8:1805¿1817. https://doi.org/10.1046/j.1365-294x.1999.00766.x PMID: 10620225 Prater, S.H. 1971. The Book of Indian Animals. Bombay Nat. Hist. Sot., Bombay: 299-300. Rankou, H., Culham, A., Jury, S.L., Christenhusz, M.J. 2013. The endemic flora of Morocco. Phytotaxa. 78(1):1-69. http://dx.doi.org/10.11646/phytotaxa.78.1.1 Rambaut, A., Drummond, A.J., Xie, D., Baele, G., Suchard, M.A. 2018. Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Syst Biol. 67(5):901¿ 904. https://doi.org/10.1093/sysbio/syy032 Rambaut, A., Suchard, M.A., Xie, D., Drummond, A.J. 2014. Tracer v1.6. Available from http://beast.bio.ed. ac.uk/Tracer Ramírez, O., Ojeda, A., Tomàs, A., Gallardo, D., Huang, L.S., Folch, J.M., et al. 2009. Integrating Y-chromosome, mitochondrial, and autosomal data to analyze the origin of pig breeds. Mol. Biol. Evol. 26(9):2061¿72. https://doi.org/10.1093/molbev/msp118 PMID: 19535739 Ramos-Muñoz, J., Cantillo-Duarte, J.J., Bernal-Casasola, D., Barrena-Tocino, A., Domínguez-Bella, S., Vijande-Vila, E., et al. 2016. Early use of marine resources by Middle/Upper Pleistocene human societies: The case of Benzú rockshelter (northern Africa). Quat. Int. 407, Part B:6¿15. https://doi.org/10.1016/j. quaint.2015.12.092 Ravaoarimanana, L.B., Tiedemann, R., Montagnon, D., Rumpler, Y. 2004. Molecular and cytogenetic evidence for cryptic speciation within a rare endemic Malagasy lemur, the Northern Sportive Lemur (Lepilemur septentrionalis). Mol. Phylogenet. Evol. 31(2):440-448. https://doi.org/10.1016/j.ympev.2003.08.020 PMID: 15062786 Rhouda, T., Martínez-Redondo, D., Gómez-Durán, A., Elmtili, N., Idaomar, M., Díez-Sánchez, C., et al. 2009. Moroccan mitochondrial genetic background suggests prehistoric human migrations across the Gibraltar Strait. Mitochondrion. 9(6):402¿7. https://doi.org/10.1016/j.mito.2009.07.003 PMID: 19631765 Riddle, B. R., Jezkova, T., Eckstut, M. E., Oláh-Hemmings, V., Carraway, L. N. 2014. Cryptic divergence and revised species taxonomy within the Great Basin pocket mouse, Perognathus parvus (Peale, 1848), species group. J. Mammal. 95(1):9¿25. https://doi.org/10.1644/12-MAMM-A-252 Rose, K.D., DeLeon, V.B., Missiaen, P., Rana, R.S., Sahni, A., Singh, L. et al. 2008. Early Eocene lagomorph (Mammalian) from Western India and the early diversification of Lagomorpha. Proc. R. Soc. B. 275:1203-1208. https://doi.org/10.1098/rspb.2007.1661 PMC2602686 PMID: 18285282 Rosell, C., Fernández-Llario, P., Herrero, J. 2001. El jabalí (Sus scrofa Linnaeus, 1758). Galemys. 13(2). Rotter, M.C., Couture, J.J., Rothwell, E.M., Garcia, J., Holeski, L.M. 2018. Evolutionary ecology of plant resistance traits across the herbivore diet spectrum: a test in the model plant Mimulus guttatus. Evol. Ecol. Res. 19:423-440. Ruvinsky, A., Rothschild, M.F. 1998. Systematics and evolution of the pig. Iowa State University. Available from: https://lib.dr.iastate.edu/cgi/viewcontent.cgi?article=1415&context=ans_pubs Saint-Loup, R. 1984. Sur una espece marocaine du genre Lepus (Lepus schlumbergeri, Nova Species). pp. 168 ¿ 172. Bulletin de la Société zoologique de France. Scandura, M., Iacolina, L., Ben Slimen, H., Suchentrunk, F., Apollonio, M. 1999. Mitochondrial CR-1 Variation in Sardinian Hares and Its Relationships with Other Old World Hares (Genus Lepus). Biochem. Genet. 45:305¿323. https://doi.org/10.1007/s10528-007-9076-z Scandura, M., Iacolina, L., Crestanello, B., Pecchioli, E., Di Benedetto, M.F., Russo,V., et al. 2008. Ancient vs. recent processes as factors shaping the genetic variation of the European wild boar: Are the effects of the last glaciation still detectable? Mol. Ecol. 17(7):1745¿62. https://doi.org/10.1111/j.1365-294X.2008. 03703.x PMID: 18371016 Schmitt, T. 2007. Molecular biogeography of Europe: Pleistocene cycles and postglacial trends. Front Zool. 4(1):11. https://doi.org/10.1186/1742-9994-4-11 PMC1868914 PMID: 17439649 Sheets, E. A., Warner, P. A., Palumbi, S. R. 2018. Accurate population genetic measurements require cryptic species identification in corals. Coral Reefs. 37(2):549¿563. https://doi.org/10.1007/s00338-018-1679-9 Sher, A., 1999. Traffic lights at the Beringian crossroads. Nature 397:103¿104. https://doi.org/10.1038/16341 Singer, M.S. 2008. Evolutionary ecology of polyphagy: specialization, speciation, and radiation. In: The Evolutionary Biology of Herbivorous Insects (Tilmon, K.J., ed), pp. 29¿42. Berkeley, CA: University of California Press. Singer, M.S., Bernays, E.A. 2009. Specialized generalists: behavioral and evolutionary ecology of polyphagous woolly bear caterpillars. In: Tiger Moths and Woolly Bears: Behavior, Ecology, and Evolution of the Arctiidae (Conner, W.E., ed), pp. 83¿101. Oxford: Oxford University Press. Soisook, P., Karapan, S., Srikrachang, M., Dejtaradol, A., Nualcharoen, K., Bumrungsri, S., et al. 2016. Hill Forest Dweller: A New Cryptic Species of Rhinolophus in the (pusillus Group) (Chiroptera: Rhinolophidae) from Thailand and Lao PDR. Acta Chiropterologica. 18(1):117¿139. https://doi.org/10.3161/15081109ACC2016.18.1.005 Soria-Boix, C., Donat-Torres, M.P., Urios, V. 2017. Contacts in the last 90,000 years over the Strait of Gibraltar evidenced by genetic analysis of wild boar (Sus scrofa). PLOS ONE. 12(7): e0181929. https://doi.org/10.1371/journal.pone.0181929 PMC5526546 PMID: 28742834 Spitz, F. 1986. Current state of knowledge of wild boar biology. Pig News and Information. 7(2):171-175. Srinivasulu, C., Srinivasulu, A., Srinivasulu, B., Jones, G. 2019. Integrated approaches to identifying cryptic bat species in areas of high endemism: The case of Rhinolophus andamanensis in the Andaman Islands. PLOS ONE. 14(10):e0213562. https://doi.org/10.1371/journal.pone.0213562 PMC6786537 PMID: 31600196 Straus, L.G. 2001. Africa and Iberia in the Pleistocene. Quat Int. 75(1):91¿102. https://doi.org/10. 1016/S1040-6182(00)00081-1 Suchentrunk, F., Ben Slimen, H., Sert, H. 2008. Phylogenetic Aspects of Nuclear and Mitochondrial Gene-Pool Characteristics of South and North African Cape Hares (Lepus capensis) and European Hares (Lepus europaeus). In: Lagomorph Biology: Evolution, Ecology and Conservation (Alves, P.C., Ferrand, N., Hacklander, K., eds), pp. 65¿88. Springer, Berlin, Heidelberg. Tamura, K., Stecher, G., Peterson, D., Filipski, A., Kumar, S. 2013. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Mol. Biol. Evol. 30(12):2725¿2729. https://doi.org/10.1093/ molbev/mst197 PMC3840312 PMID: 24132122 Tavaré, S. 1986. Some Probabilistic and Statistical Problems in the Analysis of DNA Sequences In American Mathematical Society: Lectures on Mathematics in the Life Sciences. AMS. 17:57¿86 Tegelström, H. 1987. Transfer of mitochondrial DNA from the northern red-backed vole (Clethrionomys rutilus) to the bank vole (C. glareolus). J. Mol. Evol. 24(3):218-27. https://doi.org/10.1007/bf02111235 PMID: 3106637 Teimori, A., Abd Jalil Jawad, L., Al-Kharusi, L.H., Al-Mamry, J.M., Reichenbacher, B. 2012. Late Pleistocene to Holocene diversification and historical zoogeography of the Arabian killifish (Aphanius dispar) inferred from otolith morphology. Sci. Mar. 76(4). https://doi.org/10.3989/scimar.03635.26C Thulin, C.G., Fang, M., Averianov, A.O. 2006a. Introgression from Lepus europaeus to L. timidus in Russia revealed by mitochondrial single nucleotide polymorphisms and nuclear microsatellites. Hereditas. 143:68¿76. https://doi.org/10.1111/j.2006.0018-0661.01952.x PMID: 17362337 Thulin, C.G., Jaarola, M., Tegelström, H. 1997. The occurrence of mountain hare mitochondrial DNA in wild brown hares. Mol. Ecol. 6:463¿467. https://doi.org/10.1046/j.1365-294X.1997.t01-1-00199.x PMID: 9161014 Thulin, C.G., Stone, J., Tegelström, H., Walker, C.W. 2006b. Species assignment and hybrid identi¿cation among Scandinavian hares Lepus europaeus and L. timidus. Wildlife Biol. 12:29¿38. https://doi.org/10.2981/0909-6396(2006)12[29:SAAHIA]2.0.CO;2 Thulin, C.G., Tegelström, H. 2002. Biased geographical distribution of mitochondrial DNA that passed the species barrier from mountain hares to brown hares (genus Lepus): an effect of genetic incompatibility and mating behavior? J. Zool. Lond. 258 (3):299-306. https://doi.org/10.1017/S0952836902001425 Tolesa, Z. 2014. Evolutionary Relationships among Hares (Lepus spp.) from Ethiopia: Multivariate Morphometry, Molecular Phylogenetics and Population Genetics. Thesis. Ethiopia: Addis Ababa University. Available from: http://etd.aau.edu.et/bitstream/handle/123456789/9087/Zelalem%20Gebremariam.pdf?sequence=1&isAllowed=y Tolesa, Z., Bekele, E., Tesfaye, K., Ben Slimen, H., Valqui, J., Getahun, A. et al. 2017. Mitochondrial and nuclear DNA reveals reticulate evolution in hares (Lepus spp., Lagomorpha, Mammalia) from Ethiopia. PLoS One. 12(8):e0180137. https://doi.org/10.1371/journal.pone.0180137 PMC5540492 PMID: 28767659 Urios, V., Donat-Torres, M.P., Ramírez, C., Monroy-Vilchis, O., Rgribi-Idrissi, H. 2015. El análisis del genoma mitocondrial del cánido estudiado en Marruecos manifiesta que no es ni lobo (Canis lupus) ni chacal euroasiático (Canis aureus). Altotero, Monografías. 3:1-24. https://doi.org/10.6084/m9.figshare.1524971 Urios, V., Soria, C. 2019. Situación Actual de los Grandes Depredadores Terrestres. In: Situación Actual de los Grandes Depredadores Terrestres (Monroy-Vilchis, O., Urios, V., Zarco-González, M., eds). Centro de Investigación en Ciencias Biológicas Aplicadas, Editorial Colofón, Universidad Autónoma de México, México. Valdés, B. 1991. Andalusia and the Rif: Floristic links and common flora. Bot. Chronika. 10: 117¿124. Valdés, B., Rejdali, M., Achhal El Kadmiri, A., Jury, S.L., Montserrat, J.M. eds. 2002. Catalogue des plantes vasculaires du nord du Maroc, incluant des clés d'identification = Checklist of vascular plants of N Morocco with identification keys; Volume II. Madrid: Consejo Superior de Investigaciones Científicas. van Asch, B., Pereira, F., Santos, L.S., Carneiro, J., Santos, N., Amorim, A. 2012. Mitochondrial lineages reveal intense gene flow between Iberian wild boars and South Iberian pig breeds. Anim. Genet. 43(1):35¿41. https://doi.org/10.1111/j.1365-2052.2011.02222.x PMID: 22221023 Verner, D., Tréguer, D., Redwood, J., Christensen, J., McDonnell, R., Elbert, C., et al. 2018. Climate Variability, Drought, and Drought Management in Morocco¿s Agricultural Sector. https://doi.org/10.1596/30603 Vigilant, L., Stoneking, M., Harpending, H., Hawkes, K., Wilson, A.C. 1991. African populations and the evolution of human mitochondrial DNA. Science. 253(5027):1503¿1507. http://doi.org/10.1126/science.1840702 PMID: 1840702 Vilaça, S.T., Biosa, D., Zachos, F., Iacolina, L., Kirschning, J., Alves, P.C., et al. 2014. Mitochondrial phylogeography of the European wild boar: the effect of climate on genetic diversity and spatial lineage sorting across Europe. J. Biogeogr. 41(5):987¿998. https://doi.org/10.1111/jbi.12268 Voorhies, M.R., Timperley, C.L. 1997. A new Pronotolagus (Lagomorph: Leporidae) and other leporids from Valentine railway quarries (Barstovian, Nebraska), and the archaeolagine-leporine transition. J. Vertebrate Paleontology. 17:725-737. Wagner, P., Melville, J., Wilms, T.M., Schmitz, A. 2011. Opening a box of cryptic taxa¿the first review of the North African desert lizards in the Trapelus mutabilis Merrem, 1820 complex (Squamata: Agamidae) with descriptions of new taxa. Zool. J. Linnean Soc. 163(3):884¿912. https://doi.org/10.1111/j.1096-3642.2011.00726.x Waltari, E., Cook, J.A. 2005. Hares on ice: phylogeography and historical demographics of Lepus arcticus, L. othus and L. timidus (Mammalia: Lagomorpha). Mol. Ecol. 14:3005¿3016. https://doi.org/10.1111/j.1365-294X.2005.02625.x PMID: 16101770 Waltari, E., Demboski, J.R., Klein, D.R., Cook, J.A. 2004. A molecular perspective on the historical biogeography of the northern high latitudes. J. Mammal. 85:591¿600. https://doi.org/10.1644/BER-101 Wares, J.P. 2001. Patterns of speciation inferred from mitochondrial DNA in North American Chthamalus (Cirripedia: Balanomorpha: Chthamaloidea). Mol. Phylogenet. Evol. 18(1):104-16. https//doi.org/10.1006/mpev.2000.0857 PMID: 11161747 Wilson, D.E., Leacher, T.E. Jr., Mittermeirer, R.A. 2016. Handbook of the mammals of the World. Volume 6. 1st ed. Lynx Edicions in association with Conservation International and IUCN. White, J.A. 1991. North American Leporidae (Mammalia, Lagomorpha) from late Miocene (Clarendonian) to late Pliocene (Blancan). J. Vertebrate Paleontology. 1:67-89. https://doi.org/10.1080/02724634.1991.10011376 Wu, C.H., Li, H.P, Wang, Y.X., Zhang, Y.P. 2000. Low genetic variation of the Yunnan Hare (Lepus comus Allen 1927) as revealed by mitochondrial Cytb gene sequences. Biochem. Genet. 38:149¿155. PMID: 1091905 Wu, C.H., Wu, J., Bunch, T.D., Li, Q., Wang, Y., Zang, Y. 2005. Molecular phylogenetics and biogeography of Lepus in Eastern Asia based on mitochondrial DNA sequences. Mol. Phylogenet. Evol. 37:45¿61. https://doi.org/10.1016/j.ympev.2005.05.006 PMID: 15990340 Wu, G.S., Yao, Y.G., Qu, K.X., Ding, Z.L., Li, H., Palanichamy, M.G., et al. 2007. Population phylogenomic analysis of mitochondrial DNA in wild boars and domestic pigs revealed multiple domestication events in East Asia. Genome Biol. 8:R245. https://doi.org/10.1186/gb-2007-8-11-r245 PMC2258183 PMID: 18021448 Wu, Y.H., Xia, L., Zhang, Q., Yang, Q. 2010. Habitat fragmentation affects genetic diversity and differentiation of the Yarkand hare. Conserv. Genet. 11:183¿194. https://doi.org/10.1007/s10592-009-0022-y Wu, Y.H., Xia, L., Zhang, Q., Yang, Q.S., Meng, X.X. 2011. Bidirectional introgressive hybridization between Lepus capensis and Lepus yarkandensis. Mol. Phyl. Evol. 59(3):545¿55. https://doi.org/10.1016/j.ympev.2011.03.027 PMID: 2146369 Yalden, D.W., Largen, M.J., Dock, D., Hilman, J.C. 1996. Catalogue of the Mammals of Ethiopia and Eritrea. 7. Revised check list, Zoogeography and Conservation. Trop. Zool. 9: 73-164. https://doi.org/10.1080/03946975.1996.10539304 Yamada, F. Takaki, M., Suzuki, H. 2002. Molecular phylogeny of Japanese Leporidae, the Amami rabbit Pentalagus furnessi, the Japanese hare Lepus brachyurus, and the mountain hare Lepus timidus, inferred from mitochondrial DNA sequences. Genes Genet. Syst. 77:107¿116. https://doi.org/10.1266/ggs.77.107 PMID: 12087193 Yoshikawa, S., Mimura, M., Watanabe, S., Lin, L.K., Ota, H., Mizoguchi, Y. 2016. Historical Relationships among Wild Boar Populations of the Ryukyu Archipelago and Other Eurasian regions, as Inferred from Mitochondrial Cytochrome b Gene Sequences. Zoolog. Sci. 33(5):520¿526. https://doi.org/10.2108/ zs160025