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FIGURE 1. Location of the Middle Pleistocene Qesem Cave.

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FIGURE 2. Squares of the excavation area of the Middle Pleistocene Qesem Cave with concentration (green areas) of microvertebrate fossils.

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FIGURE 3. Remains of (3.1-6) Rhinolophus mehelyi Matschie, 1901, (3.7-12) Rhinolophus euryale Blasius, 1853, and (3.13-24) Rhinolophus ferrumequinum (Schreber, 1774) from the Middle Pleistocene Qesem Cave: (3.1-3) TAU-QC/CHIR-12b, left mandibular fragment, occlusal, buccal, lingual view; (3.4-6) TAU-QC/CHIR-25, left mandibular fragment, occlusal, buccal, lingual view; (3.7-9) TAU-QC/CHIR-37, right upper canine, mesial, lingual, buccal view; (3.10-12) TAU-QC/CHIR-03, right mandibular fragment, occlusal, buccal, lingual view; (3.13-15) TAU-QC/CHIR-10, right upper canine, mesial, lingual, buccal view; (3.16-18), distal epiphysis of right humerus, TAU-QC/CHIR-05, internal, posterior, external view; (3.19-21) TAU-QC/CHIR-12a and (3.22-24) TAU-QC/CHIR-46, left mandibular fragments, occlusal, lingual, buccal view; .

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FIGURE 4. Remains of (4.1-9) Miniopterus cf. schreibersii (Kuhl, 1817), and (4.10-15) Myotis blythii (Tomes, 1857) from the Middle Pleistocene Qesem Cave: (4.1-3) TAU-QC/CHIR-04, right mandibular fragment, occlusal, buccal, lingual view; (4.4-6) TAU-QC/CHIR-07, left mandibular fragment, occlusal, buccal, lingual view; (4.7-9) TAU-QC/CHIR-48, left mandibular fragment, occlusal, lingual, buccal view; (4.10-12) TAU-QC/CHIR-40, left mandibular fragment, occlusal, buccal, lingual view; (4.13-15) TAU-QC/CHIR-02, left mandibular fragment, occlusal, buccal, lingual view.

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FIGURE 5. Remains of cf. Rousettus sp. from the Middle Pleistocene Qesem Cave: (5.1-3) TAU-QC/CHIR-26: right ?M1 fragment, occlusal, buccal, lingual view; (5.4-7) TAU-QC/CHIR-51: left C fragment, buccal, mesial, palatal, distal view.

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FIGURE 6. Extant Rousettus aegyptiacus (Qualamun, Dakhla, Egypt, Coll. Univ. Praha): (6.1 –7) M1 and (6.8–23) C. The views correspond to those of the fossil specimens figured in Figure 5. Upper dentitions of extant (6.24–25) Eidolon helvum (Malawi, Coll. Univ. Ceske Budejovice) and (6.26–27) Rousettus aegyptiacus (Rash130, Dakhla, Egypt, Coll. Univ. Praha), lateral and palatal views.

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Bat remains (Mammalia, Chiroptera) from the Middle Pleistocene site of Qesem Cave, Israel, with the first Pleistocene record of Fruit Bats in the Mediterranean region

Ivan Horáček, Lutz Christian Maul, K.T. Smith, Ran Barkai, and Avi Gopher

Plain Language Abstract

The Eastern Mediterranean (including the Levant and Near East) is an essential hot-spot of global biodiversity and a crossroad of faunal development of the Western Palearctic. The extant bat fauna in this region is composed of 64 species of European, African, and Asiatic origin; 60 of these reach margin of their distribution ranges here. To understand this biogeographic pattern requires data on history of bats in the region and on past dynamics of the range margins of particular species. Reliable information on these topics can be directly inferred only from the fossil record.

This paper briefly surveys the scarce fossil record of bats available from the Levantine zone of the E Mediterranean region and supplements it with new data from the recently excavated Middle Pleistocene site Qesem Cave, Israel, famous for records of human artifacts and numerous vertebrate remains. The five cave-dwelling microchiropteran species identified in the sample are common also in extant communities in the region, and exhibit the same phenotypic difference from northern populations as extant populations do. We also found two tooth fragments that are the first Pleistocene record of fruit bats outside of the tropics. The fragments, tentatively referred to as cf. Rousettus sp., suggest, in contrast to previous studies, that the Quaternary history of fruit bats in this region can be traced back further than the Holocene. In general, the Pleistocene record of bats in the Levant (including Qesem) is characterized by the absence of the African elements that dominate the extant Levantine fauna; instead, the vast majority of the fossil bat forms are elements of the Palaearctic clades.

Resumen en Español

Restos de murciélagos (Mammalia, Chiroptera) del Pleistoceno medio de la localidad de Qesem Cave (Israel), con el primer registro pleistoceno de murciélagos frugívoros de la región mediterránea

Son escasos los datos disponibles en la actualidad de mamíferos fósiles del Próximo Oriente, una región de gran importancia biogeográfica para este grupo en el Paleártico occidental. En este trabajo se estudia una colección de murciélagos fósiles del Pleistoceno medio de la localidad de Qesem Cave (Israel), famosa por los artefactos líticos, restos de homínidos y ricas faunas de vertebrados. Hemos identificado cinco especies de microquirópteros, concretamente Rhinolophus ferrumequinum, R. euryale, R. mehelyi, Miniopterus cf. schreibersii y Myotis blythii, todas ellas habitantes de cuevas y frecuentes todavía en las comunidades actuales de la región. Su presencia sugiere un clima suave con vegetación en mosaico durante el depósito. Junto a esas especies se han encontrado, además, dos fragmentos de dientes provisionalmente identificados como cf. Rousettus sp., lo que constituye el primer registro pleistoceno de murciélagos frugívoros fuera de los trópicos. Si se confirma esa determinación, la historia cuaternaria de los murciélagos frugívoros en esta región se remontaría hasta el Pleistoceno, antes de lo que se pensaba anteriormente, y la población actual extralimital de Rousettus aegyptiacus podría considerarse finalmente como un elemento paleócoro de la biota mediterránea. En general, el registro pleistoceno de murciélagos en el Levante mediterráneo (incluyendo Qesem) se caracteriza por la ausencia de los elementos etiópicos que dominan la fauna levantina actual, siendo la mayor parte de las formas fósiles de murciélagos miembros de clados paleárticos.

Palabras clave: Pleistoceno medio; Israel; Chiroptera; taxonomía; tafonomía; biogeografía; Rousettus

Traducción: Miguel Company

Résumé en Français

Des restes de chauve-souris (Mammalia, Chiroptera) du site pléistocène moyen de la grotte de Qesem, Israël, et le premier enregistrement pléistocène de chauves-souris frugivores dans la région méditerranéenne.

Jusqu'à maintenant, seules de rares données sont disponibles sur les chauves-souris fossiles de Proche-Orient, une région de grande importance biogéographique pour ce groupe dans le paléarctique de l'Ouest. Nous présentons ici une collection de chauves-souris fossiles du site pléistocène moyen de la grotte de Qesem, Israël, connue pour ses artefacts lithiques, ses restes d'homininés et sa riche faune de vertébrés. Nous identifions cinq espèces de microchiroptères, viz. Rhinolophus ferrumequinum, R. euryale, R. mehelyi, Miniopterus cf. schreibersii, et Myotis blythii, tous des habitants de grottes commun aussi dans les communautés actuelles de la région. Leur présence suggère des conditions douces avec une végétation variée au moment du dépôt. De plus, deux fragments de dents, provisoirement identifiés comme cf. Rousettus sp., fournissent le premier enregistrement pléistocène de chauves-souris frugivores au delà des tropiques. Si ces restent sont confirmés, l'histoire quaternaire des chauves-souris frugivores dans cette région pourrait remonter au Pléistocène, plus tôt que précédemment supposé, et donc la population actuelle extra-limite de Rousettus aegyptiacus pourrait éventuellement être vue comme un élément ancestral du biota méditerranéen. En général, le registre pléistocène des chauves-souris dans la Levant (incluant Qesem) est caractérisé par l'absence d'éléments éthiopiens qui dominent la faune Levantine actuelle, alors que la vaste majorité des formes de chauves-souris fossiles sont des éléments de clades paléarctiques.

Mots clés : Pléistocène moyen ; Israël ; Chiroptera; taxonomie; taphonomie; biogéographie; Rousettus

Translator: Olivier Maridet

Deutsche Zusammenfassung

In progress

Translator: Eva Gebauer

Arabic

380 arab

Translator: Ashraf M.T. Elewa

 

 

horacekIvan Horáček
Department of Zoology
Charles University
Vinicna 7
CZ 128 44 Praha|
Czech Republic
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Ivan Horacek is a professor and past chair in vertebrate zoology at Faculty of Science, Charles University in Prague, Czech Republic. Born 1952, graduated in vertebrate zoology in 1976 (Charles Univ. Prague 1976), PhD in paleontology and Quaternary geology in 1984 (Geological Institute of Czech Academy of Sciences).

Published about 200 papers on biology and taxonomy of bats (including fossils), Late Cenozoic small mammals of Europe and comparative morphology of vertebrates

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maulLutz Christian Maul
Research Station of Quaternary Palaeontology
Senckenberg Research Institute and Natural History Museum
Am Jakobskirchhof 4
99423 Weimar
Germany
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Dr. Lutz Christian Maul, graduated 1990 in Zoology at the Humboldt University in Berlin, employed at the Institute of Quaternary Palaeontology, at the University of Jena, since 2000 at Senckenberg Research Institute (Dpt. Quaternary Palaeontology in Weimar). Interests: Neogene and Quaternary Micromammals.

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smithK.T. Smith
Department of Palaeoanthropology and Messel Research
Senckenberg Research Institute and Natural History Museum
Senckenberganlage 25
60325 Frankfurt
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Krister Smith completed his Bachelor’s degree in geology at the University of California at Berkeley in 2000. He then moved to Yale University, where he received his PhD in 2006. He had post-docs at the University of Texas at Austin and the Senckenberg Research Institute in Frankfurt, Germany, where he is now head of the division palaeoherpetology. His active research projects are in Mesozoic and Cenozoic reptiles, taphonomy, and quantitative paleoecology.

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barkaiRan Barkai
Department of Archaeology
Tel Aviv University
Ramat Aviv
Tel Aviv 69978
Israel
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Ran Barkai is a professor of prehistoric archaeology at the Department of Archaeology and Near Eastern Cultures, Tel-Aviv University, Israel. He has been excavating the Middle Pleistocene site of Qesem Cave (in collaboration with Prof. Avi Gopher) for the last decade and has been involved in Palaeolithic and Neolithic research in the Levant.

 

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gopherAvi Gopher
Department of Archaeology
Tel Aviv University
Ramat Aviv
Tel Aviv 69978
Israel
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Prof. Avi Gopher, Dept. of Archaeology and Ancient Near Eastern Cultures of Tel Aviv University has excavated mainly Neolithic sites and studied their finds, mainly lithics. In recent years he is part of a research group concentrated on plant domestication in the Levant. In the last decade he co-directs the Qesem Cave project - a Middle Pleistocene, Lower Palaeolithic site in central Israel, not far from Tel Aviv.