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Phylogeography and evolutionary history of the burrowing scorpion genus Scorpio Linnaeus, 1758 (Scorpiones: Scorpionidae) in the Mediterranean Basin
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Khammassi, Manel, Nouira, Saïd, Badry, Ahmed, Sadine, Salah Eddine, Harris, D. James (2024): Phylogeography and evolutionary history of the burrowing scorpion genus Scorpio Linnaeus, 1758 (Scorpiones: Scorpionidae) in the Mediterranean Basin. Organisms Diversity & Evolution 24 (3): 335-352, DOI: 10.1007/s13127-024-00649-8, URL: https://doi.org/10.1007/s13127-024-00649-8
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References
- Abdel-Rahman, M. A., Omran, M. A., Abdel-Nabi, I. M., Ueda, H., & McVean, A. (2009). Intraspecific variation in the Egyptian scorpion Scorpio maurus palmatus venom collected from different biotopes. Toxicon, 53, 349-359. https://doi.org/10.1016/j. toxicon.2008.12.007
- Ajirlu, M. S., Moazzen, M., & Hajialioghli, R. (2016). Tectonic evolution of the Zagros orogen in the realm of the Neotethys between the Central Iran and Arabian Plates: An ophiolite perspective. Central European Geology, 59(1-4), 1-27. https:// doi.org/10.1556/24.59.2016.001
- Al-Saraireh, M., Yagmur, E. A., Abu Afifeh, B., & Amr, Z. (2023). A new species of Scorpio from Jordan (Scorpiones: Scorpionidae). Euscorpius, 2023(369), 1-17.
- Albrecht, F., Hering, J., Fuchs, E., Illera, J. C., Ihlow, F., Shannon, T. J., Collinson, J. M., Wink, M., Martens, J., & Packert, M. (2020). Phylogeny of the Eurasian Wren Nannus troglodytes (Aves: Passeriformes: Troglodytidae) reveals deep and complex diversification patterns of Ibero-Maghrebian and Cyrenaican populations. PLoS ONE, 15(3), e0230151. https://doi. org/10.1371/journal.pone.0230151
- Altschul, S. F., Gish, W., Miller, W., Myers, E. W., & Lipman, D. J. (1990). Basic local alignment search tool. Journal of Molecular Biology, 215(3), 403-410. https://doi.org/10.1016/S0022- 2836(05)80360-2
- Antonelli, A., Kissling, W. D., Flantua, S. G., Bermudez, M. A., Mulch, A., Muellner-Riehl, A. N., Kreft, H., Peter Linder, H., Badgley, C., Fjeldsa, J., Fritz, S. A., Rahbek, C., Herman, F., Hooghiemstra, H., & Hoorn, C. (2018). Geological and climatic influences on mountain biodiversity. Nature Geoscience, 11(10), 718-725. https://doi.org/10.1038/s41561-018-0236-z
- Beauchamp, W., Allmendinger, R. W., Barazangi, M., Demnati, A., El Alji, M., & Dahmani, M. (1999). Inversion tectonics and the evolution of the High Atlas Mountains, Morocco, based on a geological-geophysical transect. Tectonics, 18(2), 163-184. https://doi.org/10.1029/1998TC900015
- Birula, A. (1910). Ueber Scorpio maurus Linne und seine Unterarten. Horae Societatis Entomologicae Rossicae, 39, 115-192.
- Bonnefille, R. (2010). Cenozoic vegetation, climate changes and hominid evolution in tropical Africa. Global and Planetary Change, 72(4), 390-411. https://doi.org/10.1016/j.gloplacha. 2010.01.015
- Bosworth, W. (2015). Geological evolution of the Red Sea: Historical background, review, and synthesis. The Red Sea, 45-78. https:// doi.org/10.1007/978-3-662-45201-1_3
- Bryja, J., Sumbera, R., Kerbis Peterhans, J. C., Aghova, T., Bryjova, A., Mikula, O., Nicolas, V., Denys, C., & Verheyen, E. (2017). Evolutionary history of the thicket rats (genus Grammomys) mirrors the evolution of African forests since late Miocene. Journal of Biogeography, 44(1), 182-194. https://doi.org/10.1111/jbi.12890
- Bryson, R. W., Prendini, L., Savary, W. E., & Pearman, P. B. (2014). Caves as microrefugia: Pleistocene phylogeography of the troglophilic North American scorpion Pseudouroctonus reddelli. BMC Evolutionary Biology, 14(1), 1-17. https://doi.org/10.1186/ 1471-2148-14-9
- Bryson, R. W., Jr., Riddle, B. R., Graham, M. R., Smith, B. T., & Prendini, L. (2013). As old as the hills: Montane scorpions in southwestern North America reveal ancient associations between biotic diversification and landscape history. PLoS ONE, 8(1), e52822. https://doi.org/10.1371/journal.pone.0052822
- Cain, S., Loria, S. F., Ben-Shlomo, R., Prendini, L., & Gefen, E. (2021). Dated phylogeny and ancestral range estimation of sand scorpions (Buthidae: Buthacus) reveal Early Miocene divergence across land bridges connecting Africa and Asia. Molecular Phylogenetics Evolution, 164, 107-212. https://doi.org/10.1016/j. ympev.2021.107212
- Charles-Dominique, T., Davies, T. J., Hempson, G. P., Bezeng, B. S., Daru, B. H., Kabongo, R. M., Maurinc, O., Muasyaa, A. M., van der Bank, M., & Bond, W. J. (2016). Spiny plants, mammal browsers, and the origin of African savannas. Proceedings of the National Academy of Sciences, 113(38), E5572-E5579. https:// doi.org/10.1073/pnas.1607493113
- Chernomor, O., von Haeseler, A., & Minh, B. Q. (2016). Terrace aware data structure for phylogenomic inference from supermatrices.Systematic Biology, 65, 997-1008. https://doi.org/10.1093/sysbio/syw037
- Couvreur, T. L., Chatrou, L. W., Sosef, M. S., & Richardson, J. E. (2008). Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees. BMC Biology, 6(1), 1-10. https://doi.org/10.1186/1741-7007-6-54
- Couvreur, T. L., Dauby, G., Blach-Overgaard, A., Deblauwe, V., Dessein, S., Droissart, V., Hardy, O. J., Harris, D. J., Janssens, S. B., Ley, A. C., Mackinder, B. A., Sonke, B., Sosef, M. S. M., Stevart, T., Svenning, J.-C., Wieringa, J. J., Faye, A., Missoup, A. D., Tolley, K. A., Nicolas, V., Ntie, S., Fluteau, F., Robin, C., Guillocheau, F., Barboni, D., & Sepulchre, P. (2021). Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna. Biological Reviews, 96(1), 16-51. https://doi.org/10.1111/brv.12644
- Drake, N., El-Hawat, A., Turner, P., Armitage, S., Salem, M., White, K., & McLaren, S. (2008). Palaeohydrology of the Fazzan Basin and surrounding regions: The last 7 million years. Palaeogeography, Palaeoclimatology, Palaeoecology, 263(3-4), 131-145. https://doi.org/10.1016/j.palaeo.2008.02.005
- Drummond, A. J., Suchard, M. A., Xie, D., & Rambaut, A. (2012). Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29(8), 1969-1973. https:// doi.org/10.1093/molbev/mss075
- Edgar, R. C. (2004). MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32(5), 1792-1797. https://doi.org/10.1093/nar/gkh340
- Fjeldsa, J., Johansson, U. S., Lokugalappatti, L., & Bowie, R. C. (2007). Diversification of African greenbuls in space and time: Linking ecological and historical processes. Journal of Ornithology, 148(2), 359-367. https://doi.org/10.1007/s10336-007-0179-4
- Folmer, O., Black, M., Hoeh, W., Lutz, R., & Vrijenhoek, R. (1994). DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 35, 294-299.
- Frigola, A., Prange, M., & Schulz, M. (2018). Boundary conditions for the middle Miocene climate transition (MMCT v1. 0). Geoscientific Model Development, 11(4), 1607-1626. https://doi.org/10. 5194/gmd-11-1607-2018
- Froufe, E., Sousa, P., Alves, P. C., & Harris, D. J. (2008). Genetic diversity within Scorpio maurus (Scorpiones: Scorpionidae) from Morocco: Preliminary evidence based on CO1 mitochondrial DNA sequences. Biologia, 63(6), 1157-1160. https://doi. org/10.2478/s11756-008-0176-y
- Fuchs, J., Pons, J.-M., & Bowie, R. C. (2017). Biogeography and diversification dynamics of the African woodpeckers. Molecular Phylogenetics Evolution,108, 88-100. https://doi.org/10.1016/j. ympev.2017.01.007
- Gantenbein, B., Fet, V., Gantenbein-Ritter, I. A., & Balloux, F. (2005). Evidence for recombination in scorpion mitochondrial DNA (Scorpiones: Buthidae). Proceedings of the Royal Society b: Biological Sciences, 272(1564), 697-704. https://doi.org/10. 1098/rspb.2004.3017
- Georgalis, G. L., Gawad, M. K. A., Hassan, S. M., El-Barkooky, A. N., & Hamdan, M. A. (2020). Oldest co-occurrence of Varanus and Python from Africa-First record of squamates from the early Miocene of Moghra Formation, Western Desert, Egypt. Peerj, 8, e9092. https://doi.org/10.7717/peerj.9092
- Gladstone, R., Flecker, R., Valdes, P., Lunt, D., & Markwick, P. (2007). The Mediterranean hydrologic budget from a Late Miocene global climate simulation. Palaeogeography, Palaeoclimatology, Palaeoecology, 251(2), 254-267. https://doi.org/10.1016/j. palaeo.2007.03.050
- Graham, M. R., Jaeger, J. R., Prendini, L., & Riddle, B. R. (2013). Phylogeography of Beck's desert scorpion, Paruroctonus becki, reveals Pliocene diversification in the eastern California Shear Zone and postglacial expansion in the Great Basin Desert. Molecular Phylogenetics Evolution, 69(3), 502-513. https://doi.org/10.1016/j.ympev.2013.07.028
- Graham, M. R., Myers, E. A., Kaiser, R. C., & Fet, V. (2019). Cryptic species and co-diversification in sand scorpions from the Karakum and Kyzylkum deserts of Central Asia. Zoologica Scripta, 48(6), 801-812. https://doi.org/10.1111/zsc.12381
- Graham, M. R., Olah-Hemmings, V., & Fet, V. (2012). Phylogeography of co-distributed dune scorpions identifies the Amu Darya River as a long-standing component of Central Asian biogeography: (Scorpiones: Buthidae). Zoology in the Middle East, 55(1), 95-110. https://doi.org/10.1080/09397140.2012. 10648924
- Graham, M. R., Wood, D. A., Henault, J. A., Valois, Z. J., & Cushing, P. E. (2017). Ancient lakes, Pleistocene climates and river avulsions structure the phylogeography of a large but littleknown rock scorpion from the Mojave and Sonoran deserts. Biological Journal of the Linnean Society, 122(1), 133-146. https://doi.org/10.1093/biolinnean/blx058
- Greenbaum, E., Portillo, F., Jackson, K., & Kusamba, C. (2015). A phylogeny of Central African Boaedon (Serpentes: Lamprophiidae), with the description of a new cryptic species from the Albertine Rift. African Journal of Herpetology, 64(1), 18-38. https://doi. org/10.1080/21564574.2014.996189
- Griffin, D. L. (1999). The late Miocene climate of northeastern Africa: Unravelling the signals in the sedimentary succession. Journal of the Geological Society, 156(4), 817-826. https://doi.org/10. 1144/gsjgs.156.4.0817
- Griffin, D. L. (2002). Aridity and humidity: Two aspects of the late Miocene climate of North Africa and the Mediterranean. Palaeogeography, Palaeoclimatology, Palaeoecology, 182(1-2), 65-91. https://doi.org/10.1016/S0031-0182(01)00453-9
- Grossman, A., Calvo, R., Lopez-Antonanzas, R., Knoll, F., Hartman, G., & Rabinovich, R. (2019).First record of Sivameryx (Cetartiodactyla, Anthracotheriidae) from the lower Miocene of Israel highlights the importance of the Levantine Corridor as a dispersal route between Eurasia and Africa. Journal of Vertebrate Paleontology, 39(2), e1599901. https://doi.org/10.1080/02724634.2019.1599901
- Guiller, A., & Madec, L. (2010). Historical biogeography of the land snail Cornu aspersum: A new scenario inferred from haplotype distribution in the Western Mediterranean basin. BMC Evolutionary Biology, 10(1), 1-20. https://doi.org/10.1186/ 1471-2148-10-18
- Guiraud, R., Bosworth, W., Thierry, J., & Delplanque, A. (2005). Phanerozoic geological evolution of Northern and Central Africa: An overview. Journal of African Earth Sciences, 43(1-3), 83-143. https://doi.org/10.1016/j.jafrearsci.2005.07.017
- Habel, J. C., Husemann, M., Schmitt, T., Zachos, F. E., Honnen, A.-C., Petersen, B., Parmakelis, A., & Stathi, I. (2012). Microallopatry caused strong diversification in Buthus scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa). PLoS ONE, 7(2), e29403. https://doi.org/10.1371/journal.pone.0029403
- Haffer, J. (2008). Hypotheses to explain the origin of species in Amazonia. Brazilian Journal of Biology, 68, 917-947. https://doi.org/ 10.1590/S1519-69842008000500003
- Hall, T. A. (1999). BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Paper presented at the Nucleic acids symposium series.
- Hamon, N., Sepulchre, P., Lefebvre, V., & Ramstein, G. (2013). The role of eastern Tethys seaway closure in the Middle Miocene Climatic Transition (ca. 14 Ma). Climate of the Past, 9(6), 2687- 2702. https://doi.org/10.5194/cp-9-2687-2013
- Harzhauser, M., Kroh, A., Mandic, O., Piller, W. E., Gohlich, U., Reuter, M., & Berning, B. (2007). Biogeographic responses to geodynamics: A key study all around the Oligo-Miocene Tethyan Seaway. Zoologischer Anzeiger-A Journal of Comparative Zoology, 246(4), 241-256. https://doi.org/10.1016/j. jcz.2007.05.001
- Hemp, C., Kehl, S., Schultz, O., Waegele, J. W., & Hemp, A. (2015). Climatic fluctuations and orogenesis as motors for speciation in East Africa: Case study on Parepistaurus Karsch, 1896 (Orthoptera). Systematic Entomology, 40(1), 17-34.https://doi.org/10.1111/syen.12092
- Howard, R. J., Edgecombe, G. D., Legg, D. A., Pisani, D., & Lozano- Fernandez, J. . (2019). Exploring the evolution and terrestrialization of scorpions (Arachnida: Scorpiones) with rocks and clocks. Organisms Diversity & Evolution, 19, 71-86. https://doi.org/10. 1007/s13127-019-00390-7
- Huntley, J. W., Harvey, J. A., Pavia, M., Boano, G., & Voelker, G. (2018). The systematics and biogeography of the Bearded Greenbuls (Aves: Criniger) reveals the impact of Plio-Pleistocene forest fragmentation on Afro-tropical avian diversity. Zoological Journal of the Linnean Society, 183(3), 672-686. https://doi.org/ 10.1093/zoolinnean/zlx086
- 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. Journal of Biogeography, 46(2), 381-391. https://doi.org/10.1111/jbi.13505
- Husemann, M., Schmitt, T., Stathi, I., & Habel, J. (2012). Evolution and radiation in the scorpion Buthus elmoutaouakili Lourenco and Qi 2006 (Scorpiones: Buthidae) at the foothills of the Atlas Mountains (North Africa). The Journal of Heredity, 103(2), 221-229. https://doi.org/10.1093/jhered/esr130
- Hussen, F. S., & Ahmed, S. T. (2020). New data of scorpion fauna, include two new records with identification key of scorpion species (Arachnida: Scorpiones) in Iraq. Plant Archives, 20(2), 6711-6725.
- Jacobs, B. F. (2004). Palaeobotanical studies from tropical Africa: Relevance to the evolution of forest, woodland and savannah biomes. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 359(1450), 1573-1583. https://doi.org/10.1098/rstb.2004.1533
- Jacobs, B. F., Pan, A. D., Scotese, C. R., Werdelin, L., & Sanders, W. (2010). A review of the Cenozoic vegetation history of Africa. In L. Werdelin (Eds.), Cenozoic Mammals of Africa. Oakland, CA, 2010. https://doi.org/10.1525/california/9780520257214. 003.0005
- Jacquet, F., Nicolas, V., Colyn, M., Kadjo, B., Hutterer, R., Decher, J., Akpatou, B., Cruaud, C., & Denys, C. (2014). Forest refugia and riverine barriers promote diversification in the West African pygmy shrew (Cocidura obscurior complex, Soricomorpha). Zoologica Scripta, 43(2), 131-148. https://doi.org/10.1111/zsc. 12039
- Keppel, G., Van Niel, K. P., Wardell-Johnson, G. W., Yates, C. J., Byrne, M., Mucina, L., Schut, A. G. T., Hopper, S. D., & Franklin, S. E. (2012). Refugia: Identifying and understanding safe havens for biodiversity under climate change. Global Ecology Biogeography, 21(4), 393-404. https://doi.org/10.1111/j.1466- 8238.2011.00686.x
- Khammassi, M., Harris, D. J., Sadine, S. E., Bouhissi, M. E., & Nouira, S. (2023). Description of a new species of Scorpio (Scorpiones: Scorpionidae) from Northwestern Algeria using morphological and molecular data. Biologia, 1-12. https://doi.org/10.1007/ s11756-022-01306-1
- Klesser, R., Husemann, M., Schmitt, T., Sousa, P., Moussi, A., & Habel, J. C. (2021). Molecular biogeography of the Mediterranean Buthus species complex (Scorpiones: Buthidae) at its southern Palaearctic margin. Biological Journal of the Linnean Society, 133(1), 166-178. https://doi.org/10.1093/biolinnean/ blab014
- Koc, H., Sipahi, H., & Yagmur, E. A. (2017). Genetic diversity within Scorpio maurus (Scorpiones: Scorpionidae) from Turkey. NorthWestern Journal of Zoology, 13(1), 27-33.
- Kohler, C. M., Heslop, D., Krijgsman, W., & Dekkers, M. (2010). Late Miocene paleoenvironmental changes in North Africa and the Mediterranean recorded by geochemical proxies (Monte Gibliscemi section, Sicily). Palaeogeography, Palaeoclimatology, Palaeoecology, 285(1-2), 66-73. https://doi.org/10.1016/j. palaeo.2009.10.025
- Koufos, G. D., Kostopoulos, D. S., & Vlachou, T. D. (2005). Neogene/ Quaternary mammalian migrations in eastern Mediterranean. Belgian Journal of Zoology, 135(2), 181.
- Kovarik, F. (2009). Illustrated catalog of scorpions. Part I. Introductory remarks; keys to families and genera; subfamily Scorpioninae with keys to Heterometrus and Pandinus species. Prague: Clairon Production.
- Kumar, S., Stecher, G., Li, M., Knyaz, C., & Tamura, K. (2018). MEGA X: Molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution, 35(6), 1547-1549. https://doi.org/10.1093/molbev/msy096
- Lamoral, B. (1978). Soil hardness, an important and limiting factor in burrowing scorpions of the genus Opisthophthalmus CL Koch, 1837 (Scorpionidae, Scorpionida). Symposium of Zoology Society of London, 42, 171-181.
- Lamoral, B. (1979). The scorpions of Namibia (Arachnida-Scorpionida). Annals of the Natal Museum, 23, 497-784.
- Lanfear, R., Calcott, B., Ho, S. Y., & Guindon, S. (2012). PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29(6), 1695-1701. https://doi.org/10.1093/molbev/mss020
- Lanfear, R., Frandsen, P. B., Wright, A. M., Senfeld, T., & Calcott, B. (2016). PartitionFinder 2: New methods for selecting partitioned models of evolution formolecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 1-2, msw260. https://doi.org/10.1093/molbev/msw260
- Levy, G., & Amitai, P. (1980). Fauna Palaestina. Arachnida I: Scorpiones. Israel Academy of Sciences and Humanities.
- Loader, S. P., Pisani, D., Cotton, J. A., Gower, D. J., Day, J. J., & Wilkinson, M. (2007). Relative time scales reveal multiple origins of parallel disjunct distributions of African caecilian amphibians. Biology Letters, 3(5), 505-508. https://doi.org/10. 1098/rsbl.2007.0266
- Lopez-Antonanzas, R., Renaud, S., Pelaez-Campomanes, P., Azar, D., Kachacha, G., & Knoll, F. (2019). First levantine fossil murines shed new light on the earliest intercontinental dispersal of mice. Scientific Reports, 9(1), 1-16. https://doi.org/10.1038/ s41598-019-47894-y
- Lourenco, W. R. (2009). Reanalysis of the genus Scorpio Linnaeus 1758 in sub-Saharan Africa and description of one new species from Cameroon (Scorpiones, Scorpionidae). Entomologische Mitteilungen Aus Dem Zoologischen Museum Hamburg, 15(181), 99-113.
- Lourenco, W. R., & Cloudsley-Thompson, J. L. (2009). A new species of the genus Scorpio Linnaeus 1758 from Sudan (Scorpiones, Scorpionidae). Boletin De La SEA, 45, 123-126.
- Lourenco, W., & Cloudsley-thompson, J. (2012). About the enigmatic presence of the genus Scorpio Linnaeus, 1758 in Congo with the description of a new species from Niger (Scorpiones, Scorpionidae). Serket, 13(1-2), 1-7.
- Lourenco, W. R., Duhem, B., & Cloudsley-Thompson, J. L. (2012). Scorpions from Ennedi, Kapka and Tibesti, the mountains of Chad, with descriptions of nine new species (Scorpiones: Buthidae, Scorpionidae). Arthropoda Selecta, 21(4), 307-338.
- Machado, L., Harris, D. J., & Salvi, D. (2021). Biogeographic and demographic history of the Mediterranean snakes Malpolon monspessulanus and Hemorrhois hippocrepis across the Strait of Gibraltar. BMC Ecology and Evolution, 21(1), 1-14. https:// doi.org/10.1186/s12862-021-01941-3
- Mizerovska, D., Nicolas, V., Demos, T. C., Akaibe, D., Colyn, M., Denys, C., Kaleme, P. K., Katuala, P., Kennis, J., Kerbis Peterhans, J. C., Laudisoit, A., Missoup, A. D., Sumbera, R., Verheyen, E., & Bryja, J. (2019). Genetic variation of the most abundant forest-dwelling rodents in Central Africa (Praomys jacksoni complex): Evidence for Pleistocene refugia in both montane and lowland forests. Journal of Biogeography, 46(7), 1466-1478. https://doi.org/10.1111/jbi.13604
- Morley, R. J. (2000). Origin and evolution of tropical rain forests. John Wiley & Sons.
- Morley, R. (2011). Cretaceous and Tertiary climate change and the past distribution of megathermal rainforests. In M. Bush, J. Flenley, & W. Gosling (Eds.), Tropical rainforest responses to climatic change (pp. 1-34). Springer, Berlin, Heidelberg: Springer Praxis Books.
- Newlands, G. (1972). Notes on psammophilous scorpions and a description of a new species (Arachnida: Scorpionides). Annals of the Transvaal Museum, 27(12), 241-254.
- Nguyen, L. T., Schmidt, H. A., von Haeseler, A., & Minh, B. Q. (2015). IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268-274. https://doi.org/10.1093/molbev/msu300
- Nouira, S., & Blanc, C. (2003). Distribution spatiale des Lacertides (Sauria, Reptilia) en Tunisie; caracteristiques des biotopes et role des facteurs ecologiques. Ecologia Mediterranea, 29(1), 71-86.
- Novello, A., Barboni, D., Sylvestre, F., Lebatard, A.-E., Pailles, C., Bourles, D. L., Andossa Likius, A., Mackaye, H. T., Vignaud, P., & Brunet, M. (2017). Phytoliths indicate significant arboreal cover at Sahelanthropus type locality TM266 in northern Chad and a decrease in later sites. Journal of Human Evolution, 106, 66-83. https://doi.org/10.1016/j.jhevol.2017.01.009
- Ojanguren-Affilastro, A. A., Mattoni, C. I., Ochoa, J. A., Ramirez, M. J., Ceccarelli, F. S., & Prendini, L. (2016). Phylogeny, species delimitation and convergence in the South American bothriurid scorpion genus Brachistosternus Pocock 1893: Integrating morphology, nuclear and mitochondrial DNA. Molecular Phylogenetics Evolution, 94, 159-170. https://doi.org/10.1016/j.ympev. 2015.08.007
- Pandolfi, L., Calvo, R., Grossman, A., & Rabinovich, R. (2021). Rhinocerotidae from the early Miocene of the Negev (Israel) and implications for the dispersal of early Neogene rhinoceroses. Journal of Paleontology, 95(6), 1340-1351. https://doi.org/10. 1017/jpa.2021.64
- Peter, B. D. (2004). African climate change and faunal evolution during the Pliocene-Pleistocene. Earth Planetary Science Letters, 220(1-2), 3-24. https://doi.org/10.1016/S0012-821X(04) 00003-2
- Pokorny, L., Riina, R., Mairal, M., Meseguer, A. S., Culshaw, V., Cendoya, J., Miguel Serrano, M., Carbajal, R., Ortiz, S., Heuertz, M., & Sanmartin, I. (2015). Living on the edge: Timing of Rand Flora disjunctions congruent with ongoing aridification in Africa. Frontiers in Genetics, 6, 154. https://doi.org/10.3389/ fgene.2015.00154
- Popov, S. V., Shcherba, I. G., Ilyina, L. B., Nevesskaya, L. A., Paramonova, N. P., Khondkarian, S. O., & Magyar, I. (2006). Late Miocene to Pliocene palaeogeography of the Paratethys and its relation to the Mediterranean. Palaeogeography, Palaeoclimatology, Palaeoecology, 238(1-4), 91-106. https:// doi.org/10.1016/j.palaeo.2006.03.020
- Portillo, F., Branch, W. R., Conradie, W., Rodel, M.-O., Penner, J., Barej, M. F., Kusamba, C., Muninga, W. M., Aristotei, M. M., Bauer, A. M., Jean-Francois Trape, J.-F., Nagy, Z. T., Carlinom, P., Pauwels, O. S. G., Menegon, M., Burger, M., Mazuch, T., Jackson, K., Hughesa, D. F., Behangana, M., & Greenbaum, E. (2018). Phylogeny and biogeography of the African burrowing snake subfamily Aparallactinae (Squamata: Lamprophiidae). Molecular Phylogenetics Evolution, 127, 288-303. https://doi.org/10.1016/j.ympev.2018.03.019
- Pound, M. J., Haywood, A. M., Salzmann, U., & Riding, J. B. (2012). Global vegetation dynamics and latitudinal temperature gradients during the Mid to Late Miocene (15.97-5.33 Ma). Earth- Science Reviews, 112(1-2), 1-22. https://doi.org/10.1016/j. earscirev.2012.02.005
- Pound, M. J., Haywood, A. M., Salzmann, U., Riding, J. B., Lunt, D. J., & Hunter, S. J. (2011). A Tortonian (late Miocene, 11.61- 7.25 Ma) global vegetation reconstruction. Palaeogeography, Palaeoclimatology, Palaeoecology, 300(1-4), 29-45. https:// doi.org/10.1016/j.palaeo.2010.11.029
- Prendini, L. (2001). Substratum specialization and speciation in southern African scorpions: The effect hypothesis revisited. In V. Fet & P. A. Selden (Eds.), Scorpions 2001 (pp. 113-138). British Arachnological Society.
- Prendini, L., Crowe, T. M., & Wheeler, W. C. (2003). Systematics and biogeography of the family Scorpionidae (Chelicerata: Scorpiones), with a discussion on phylogenetic methods. Invertebrate Systematics, 17(2), 185-259. https://doi.org/10. 1071/IS02016
- Prendini, L., & Wheeler, W. C. (2005). Scorpion higher phylogeny and classification, taxonomic anarchy, and standards for peer review in online publishing. Cladistics, 21(5), 446-494. https://doi.org/10.1111/j.1096-0031.2005.00073.x
- Rambaut, A. (2012). Figtree 1.4. 0. Distributed by the author via: http://tree.bio.ed.ac.uk/software/figtree
- Rambaut, A., & Drummond, A. J. (2003). Tracer: MCMC trace analysis tool. University of Oxford.
- Rangel Lopez, J. A., Husemann, M., Schmitt, T., Kramp, K., & Habel, J. C. (2018). Mountain barriers and trans-Saharan connections shape the genetic structure of Pimelia darkling beetles (Coleoptera: Tenebrionidae). Biological Journal of the Linnean Society, 124(3), 547-556. https://doi.org/10.1093/ biolinnean/bly053
- Renaud, S., Michaux, J., Schmidt, D. N., Aguilar, J.-P., Mein, P., & Auffray, J.-C. (2005). Morphological evolution, ecological diversification and climate change in rodents. Proceedings of the Royal Society b: Biological Sciences, 272(1563), 609-617. https://doi.org/10.1098/rspb.2004.2992
- Rogl, F. (1999). Mediterranean and Paratethys. Facts and hypotheses of an Oligocene to Miocene Paleogeography (short overview). Geologica Carpathica, 50, 339-349.
- Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D. L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M. A., & Huelsenbeck, J. P. (2012). MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic biology, 61(3), 539-542. https://doi.org/10.1093/sysbio/sys029
- Ronquist, F., van der Mark, P., & Huelsenbeck, J. P. (2009). Bayesian phylogenetic analysis using MRBAYES. In P. Lemey, M. Salemi, & A.-M. Vandamme (Eds.), The phylogenetic handbook: A practical approach to phylogenetic analysis and hypothesis testing (pp. 210-266). Published by Cambridge University Press.
- Rosado, D., Rato, C., Salvi, D., & Harris, D. J. (2017). Evolutionary history of the Morocco lizard-fingered geckos of the Saurodactylus brosseti complex. Evolutionary Biology, 44, 386-400. https:// doi.org/10.1007/s11692-017-9417-8
- Salicini, I., Ibanez, C., & Juste, J. (2013). Deep differentiation between and within Mediterranean glacial refugia in a flying mammal, the Myotis nattereri bat complex. Journal of Biogeography, 40(6), 1182-1193. https://doi.org/10.1111/jbi.12062
- Santibanez-Lopez, C. E., Ojanguren-Affilastro, A. A., & Sharma, P. P. (2020). Another one bites the dust: Taxonomic sampling of a key genus in phylogenomic datasets reveals more non-monophyletic groups in traditional scorpion classification. Invertebrate Systematics, 34(2), 133-143. https://doi.org/10.1071/IS19033
- Schenkel, E. (1949). Mitteilungen uber Spinnentiere. Skorpioneaus Marokko. Verhandlungen der Naturforschenden Gesellschaftin Basel, 60, 186-201.
- Schuster, M., Duringer, P., Ghienne, J.-F., Vignaud, P., Mackaye, H. T., Likius, A., & Brunet, M. (2006). The age of the Sahara Desert. Science, 311(5762), 821-821. https://doi.org/10.1126/science.1120161
- Sharma, P. P., Fernandez, R., Esposito, L. A., Gonzalez-Santillan, E., & Monod, L. (2015). Phylogenomic resolution of scorpions reveals multilevel discordance with morphological phylogenetic signal. Proceedings of the Royal Society b: Biological Sciences, 282(1804), 20142953. https://doi.org/10.1098/rspb.2014.2953
- Solovyeva, E. N., Lebedev, V. S., Dunayev, E. A., Nazarov, R. A., Bannikova, A. A., Che, J., Robert, W., Murphy, R. W., & Poyarkov, N. A. (2018). Cenozoic aridization in Central Eurasia shaped diversification of toad-headed agamas (Phrynocephalus; Agamidae, Reptilia). PeerJ, 6, e4543. https://doi.org/10.7717/peerj.4543
- Soltan-Alinejad, P., Rafinejad, J., Dabiri, F., Onorati, P., Terenius, O., & Chavshin, A. R. (2021). Molecular analysis of the mitochondrial markers COI, 12S rDNA and 16S rDNA for six species of Iranian scorpions. BMC Research Notes, 14(1), 40. https://doi.org/10.1186/s13104-021-05449-3
- Sousa, P., Froufe, E., Harris, D. J., Alves, P. C., & Van der Meijden, A. (2011). Genetic diversity of Maghrebian Hottentotta (Scorpiones: Buthidae) scorpions based on CO1: New insights on the genus phylogeny and distribution. African Invertebrates, 52(1), 135-143. https://doi.org/10.5733/afin.052.0106
- Sousa, P., Harris, D., Froufe, E., & Meijden, A. (2012). Phylogeographic patterns of Buthus scorpions (Scorpiones: Buthidae) in the Maghreb and South Western Europe based on CO1 mtDNA sequences. Journal of Zoology, 288, 66-75. https://doi.org/10. 1111/j.1469-7998.2012.00925.x
- Suchard, M. A., Lemey, P., Baele, G., Ayres, D. L., Drummond, A. J., & Rambaut, A. (2018). Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10. Virus evolution, 4(1), vey016. https://doi.org/10.1093/ve/vey016
- Talal, S., Tesler, I., Sivan, J., Ben-Shlomo, R., Tahir, H. M., Prendini, L., Snir, S., & Gefen, E. (2015). Scorpion speciation in the Holy Land: Multilocus phylogeography corroborates diagnostic differences in morphology and burrowing behavior among Scorpio subspecies and justifies recognition as phylogenetic, ecological and biological species. Molecular Phylogenetics and Evolution, 91, 226-237. https://doi.org/10.1016/j.ympev.2015.04.028
- Tamura, K., Nei, M., & Kumar, S. (2004). Prospects for inferring very large phylogenies by using the neighbor-joining method. Proceedings of the National Academy of Sciences, 101(30), 11030- 11035. https://doi.org/10.1073/pnas.0404206101
- Tejero-Cicuendez, H., Patton, A. H., Caetano, D. S., Smid, J., Harmon, L. J., & Carranza, S. (2022). Reconstructing squamate biogeography in Afro-Arabia reveals the influence of a complex and dynamic geologic past. Systematic Biology, 71(2), 261-272. https://doi.org/10.1093/sysbio/syab025
- Tlili, W., Nefla, A., Delaugerre, M., Ouni, R., & Nouira, S. (2014). Factors determining Gekkotan (Reptilia, Sauria) distribution in Tunisia (North Africa). Acta Herpetologica, 9(2), 203-217. https://doi.org/10.13128/Acta_Herpetol-12875
- Torfstein, A., & Steinberg, J. (2020). The Oligo-Miocene closure of the Tethys Ocean and evolution of the proto-Mediterranean Sea. Scientific Reports, 10(1), 1-10. https://doi.org/10.1038/ s41598-020-70652-4
- Vachon, M. (1952). Etudes sur les scorpions. Description des Scorpions du Nord de l'Afrique. Archive de l'Institut Pasteur d'Algerie, Alger.
- van der Meijden, A., Kleinteich, T., & Coelho, P. (2012). Packing a pinch: Functional implications of chela shapes in scorpions using finite element analysis. Journal of Anatomy, 220(5), 423-434. https://doi.org/10.1111/j.1469-7580.2012.01485.x
- Voje, K. L., Hemp, C., Flagstad, O., Saetre, G. P., & Stenseth, N. C. (2009). Climatic change as an engine for speciation in flightless Orthoptera species inhabiting African mountains. Molecular Ecology, 18(1), 93-108. https://doi.org/10.1111/j.1365-294X. 2008.04002.x
- Ythier, E., & Francois, A. (2023). The scorpion fauna of the Oriental region in Morocco (Scorpiones: Buthidae, Scorpionidae) with description of three new species of the genus Scorpio Linnaeus, 1758. Faunitaxys, 11(3), 1-15. https://doi.org/10. 57800/faunitaxys-11(3)