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Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan

Published online by Cambridge University Press:  25 January 2019

James T. Pokines*
Affiliation:
Department of Anatomy and Neurobiology, Boston University School of Medicine, Boston, Massachusetts 02118, USA; Office of the Chief Medical Examiner, 720 Albany Street, Boston, Massachusetts02118, USA
Adrian M. Lister
Affiliation:
Vertebrates and Anthropology, Earth Sciences Department, Natural History Museum, Cromwell Road, LondonSW7 5BD, United Kingdom
Christopher J. H. Ames
Affiliation:
University of Wollongong, School of Earth and Environmental Sciences, Wollongong, New South Wales, Australia2522 Department of Anthropology, University of Victoria, Cornett Building B228, 3800 Finnerty Road, Victoria, British ColumbiaV8P 5C2, Canada
April Nowell
Affiliation:
Department of Anthropology, University of Victoria, Cornett Building B228, 3800 Finnerty Road, Victoria, British ColumbiaV8P 5C2, Canada
Carlos E. Cordova
Affiliation:
Department of Geography, Oklahoma State University, 337 Murray Hall, Stillwater, Oklahoma74078-4073, USA
*
*Corresponding author at: Department of Anatomy and Neurobiology, Boston University School of Medicine, Boston, Massachusetts 02118, USA. E-mail address: jamespokines@hotmail.com (J.T. Pokines).

Abstract

Excavations from 2013 to 2015 at the site of Shishan Marsh 1 (SM1) in the Azraq Basin of eastern Jordan have yielded substantial late middle Pleistocene lithic assemblages in association with faunal remains. Faunal preservation is poor, but multiple taxa have been identified, including cf. Panthera leo, Gazella sp., Bos cf. primigenius, Camelus sp., Equus spp., cf. Stephanorhinus hemitoechus, Palaeoloxodon cf. recki, and Elephas cf. hysudricus. The overall Azraq habitat may have been most similar to a savanna ecosystem, with a mixture of open grassland/shrub habitats and more closed vegetation along the wetlands margins. These taxa were drawn to the relatively lush oasis environment, where they were a dietary resource of the hominin groups exploiting the wetlands resources.

Type
Research Article
Copyright
Copyright © University of Washington. Published by Cambridge University Press, 2018 

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References

REFERENCES

Abu Baker, M., Qarqaz, M., Amr, Z., 2005. Small mammal remains recovered from pellets of the Barn Owl (Tyto alba) at Shawmari Wildlife Reserve, eastern Jordan. Casopis Národního Muzea, Rada prirodovedná 174, 127129.Google Scholar
Abuhelaleh, B., Al Nahar, M., Hohenstein, U.T., Berruti, G.L.F., Cancellieri, E., 2015. An experimental study of bone artifacts from the Neolithic site of Tell Abu Suwwan (PPNB-PN), Jordan. Adumatu 32, 720.Google Scholar
Abu-Zinada, A.H., Habibi, K., Seitre, R., 1988. The Arabian oryx programme in Saudi Arabia. In: Dixon, A., Jones, D. (Eds.), Conservation and Biology of Desert Antelopes, Including the Proceedings of the 25th Anniversary Celebration of ‘Operation Oryx’. Christopher Helm, London, pp. 41–46.Google Scholar
Albayrak, E., Lister, A.M., 2012. Dental remains of fossil elephants from Turkey. Quaternary International 276–277, 198211.Google Scholar
Al-Eisawi, D., 1996. Vegetation of Jordan. UNESCO, Cairo.Google Scholar
Al-Eisawi, D., 2005. Water scarcity in relation to food security and sustainable use of biodiversity in Jordan. In: Hamdy, A, Monti, R. (Eds.), Food Security under Water Scarcity in the Middle East: Problems and Solutions. Options Méditerranéenes: Série A. Séminaires Méditerranéenes no. 65. CIHEAM, Bari, Italy, pp. 239248.Google Scholar
Al-Kharabsheh, A., 2000. Ground-water modelling and long-term management of the Azraq basin as an example of arid area conditions (Jordan). Journal of Arid Environments 44, 143153.Google Scholar
Ames, C.J.H., 2014. Hominin Occupation and Landscape Evolution during the Middle and Late Pleistocene at the Druze Marsh Site in Northeast Jordan. PhD dissertation, McGill University, Montreal.Google Scholar
Ames, C.J.H., Cordova, C.E., 2014. Geoarchaeology and prehistoric transitions in the Azraq Druz Basin, Jordan: preliminary data and hypothesis for a 4-dimensional model of landscape change. In: Finlayson, W., Rollefson, G. (Eds.), Prehistoric Jordan: Recent and Future Research. Department of Antiquities, Amman, Jordan, pp. 6980.Google Scholar
Ames, C.J.H., Cordova, C.E., 2015. Middle and Late Pleistocene landscape evolution at the Druze Marsh Site in northeast Jordan: implications for population continuity and hominin dispersal. Geoarchaeology 30, 307329.Google Scholar
Ames, C.J.H., Nowell, A., Cordova, C.E., Pokines, J.T., Bisson, M.S., 2014a. Paleoenvironmental change and settlement dynamics in the Druze Marsh: results of recent excavation at an open-air Paleolithic site. Quaternary International 331, 6073.Google Scholar
Ames, C.J.H., Nowell, A., Pokines, J.T., Cordova, C.E., 2014b. New evidence of Paleolithic occupation in the Shishan Marsh, Jordan: report on the 2013 field season of the Azraq Marshes Archaeological and Paleoecological Project. Paper presented at the 79th Annual Meeting of the Society for American Archaeology, Austin, TX, April 26, 2014.Google Scholar
Amr, Z.S., 2000. Mammals of Jordan. National Library for United Nations Environment Programme, Amman, Jordan.Google Scholar
Amr, Z.S., Al-Melhim, W.N., Yousef, M.A., 1997. Mammal remains from pellets of the Eagle Owl, Bubo bubo, from Azraq Nature Reserve, Jordan. Zoology in the Middle East 14, 510.Google Scholar
Andrews, I.J., 1995. The Birds of the Hashemite Kingdom of Jordan. Burns and Harris, Dundee, Scotland.Google Scholar
Asmodé, J.F., Khoja, A.R., 1990. Arabian oryx captive breeding and reintroduction in Saudi Arabia. In: Seal, U., Sausman, K., Mikolai, J. (Eds.), CBSG Aridland Antelope Workshop, San Antonio, TX, 13–15 September 1989. IUCN, Captive Breeding Specialist Group, Apple Valley, MN, pp. 109127.Google Scholar
Bakken, D., 2000. Hunting strategies of Late Pleistocene Zarzian populations from Palegawra Cave, Iraq and Warwasi Rock Shelter, Iran. In: Mashkour, M., Choyke, A.M., Bittenhuis, H., Poplin, F. (Eds.), Archaeology in the Near East IV A: Proceedings of the Fourth International Symposium on the Archaeology of Southwestern Asia and Adjacent Areas. ARC Publicaties 32. Centre for Archaeological Research and Consultancy, Groningen, the Netherlands, pp. 11–17.Google Scholar
Ballatore, M., Breda, M., 2013. Stephanorhinus hundsheimensis (Rhinocerontidae, Mammalia) teeth from the early Middle Pleistocene of Isernia La Pineta (Molise, Italy) and comparison with coeval British material. Quaternary International 302, 169183 Google Scholar
Bar-Oz, G., Dayan, T., Kaufman, D., 1998. Taphonomic analysis of the faunal remains from Neve David. In: Buitenhuis, H., Bartosiewicz, L., Choyke, A.M. (Eds.), Archaeozoology of the Near East III. ARC Publicaties 18. Centre for Archaeological Research and Consultancy, Groningen, the Netherlands, pp. 7279.Google Scholar
Bar-Yosef, O., Belfer-Cohen, A., 2013. Following Pleistocene road signs of human dispersals across Eurasia. Quaternary International 285, 3043.Google Scholar
Bar-Yosef, O., Belmaker, M., 2011. Early and Middle Pleistocene faunal and hominins dispersals through southwestern Asia. Quaternary Science Reviews 30, 13181337.Google Scholar
Bar-Yosef, O., Tchernov, E., 1972. On the Palaeo-Ecological History of the Site of Ubeidiya. Israel Academy of Science and Humanities, Jerusalem.Google Scholar
Beech, M., Mashkour, M., Huels, M., Zazzo, A., 2009. Prehistoric camels in south-eastern Arabia: the discovery of a new site in Abu Dhabi’s Western Region, United Arab Emirates. Proceedings of the Seminar for Arabian Studies 39, 1730.Google Scholar
Bennett, D., Hoffman, R.S., 1999. Equus caballus. Mammalian Species 628, 114.Google Scholar
Bilgin, S., Fidanbaş, Z.U.C., 2011. Nutritional properties of crab (Potamon potamios Olivier, 1804) in the Lake of Eğirdir (Turkey). Pakistan Veterinary Journal 31, 239243.Google Scholar
Bisson, M., Nowell, A., Cordova, C.E., Kalchgruber, R., 2007a. Human evolution at the crossroads: an archaeological survey in NW Jordan. Near Eastern Archaeology 69, 7385.Google Scholar
Bisson, M., Nowell, A., Cordova, C.E., Kalchgruber, R., 2007b. Neandertals at the crossroads. In: Levy, T.E., Daviau, P.M.M., Younker, R.W., Shaer, M. (Eds.), Crossing Jordan: North American Contributions to the Archaeology of Jordan. Equinox, London, pp. 179186.Google Scholar
Bisson, M., Nowell, A., Cordova, C.E., Poupart, M., Pokines, J.T., Ghaleb, B., 2014. WE-2: the Middle Paleolithic of the Wādī Al-‘Unqiyya revisited. In: Finlayson, W., Rollefson, G. (Eds.), Prehistoric Jordan: Recent and Future Research. Department of Antiquities, Amman, Jordan, pp. 3547.Google Scholar
Boëda, E., Noël-Soriano, S., Griggo, C., 2001. Différents modes d’occupation du site d’Umm el Tlel au cours du Paléolithique moyen (El Kowm, Syrie centrale). Paléorient 27, 1328.Google Scholar
Bouchud, J., 1974. Étude preliminaire de la faune provenant de la grotte du Djebel Qafzeh pres de Nazareth (Israel). Paléorient 2, 87102.Google Scholar
Brain, C.K., 1974. Some suggested procedures in the analysis of bone accumulations from Southern African Quaternary sites. Annals of the Transvaal Museum 29, 18.Google Scholar
Brühl, E., 2003. The small flint tool industry from Bilzingsleben-Steinrinne. In: Burdukiewicz, J.M., Ronen, A. (Eds.), Lower Palaeolithic Small Tools in Europe and the Levant. Archaeopress, Oxford, UK, pp. 4964.Google Scholar
Burke, A., Eisenmann, V., Ambler, G.K., 2003. The systematic position of Equus hydruntinus, an extinct species of Pleistocene equid. Quaternary Research 59, 459469.Google Scholar
Byrd, B.F., Garrard, A.N., 1990. The last glacial maximum in the Jordanian desert. In: Bar-Yosef, O., Gamble, C., (Eds.), The World at 18,000 BP. Vol. 2, Low Latitudes. Cambridge University Press, Cambridge, pp. 7896.Google Scholar
Çakirlar, C., Berthon, R., 2014. Caravans, camel wrestling and cowrie shells: towards a social zooarchaeology of camel hybridization in Anatolia and adjacent regions. Anthropozoologica 49, 237252.Google Scholar
Cerdeño, E., 1998. Diversity and evolutionary trends of the Family Rhinocerotidae (Perissodactyla). Palaeogeography, Palaeoclimatology, Palaeoecology 141, 1334.Google Scholar
Chazan, M., Avery, D.M., Bamford, M.K., Berna, F., Brink, J., Fernandez-Jalvo, Y., Goldberg, P., et al., 2012. The Oldowan horizon in Wonderwerk Cave (South Africa): archaeological, geological, paleontological and paleoclimatic evidence. Journal of Human Evolution 63, 859866.Google Scholar
Clark, G., Lindly, J., Donaldson, M., Garrard, A., Coinman, N., Schuldenrein, J., Fish, S., Olszewski, D., 1988. Excavations at Middle, Upper and Epipaleolithic sites in the Wadi Hasa, west-central Jordan. In: Garrard, A.N., Gebel, H.G. (Eds.), The Prehistory of Jordan: The State of Research in 1986. British Archaeological Reports (BAR) International Series 396(1). BAR, Oxford, UK, pp. 209285.Google Scholar
Clutton-Brock, J., 1970. The fossil fauna from an Upper Pleistocene site in Jordan. Journal of Zoology, London 162, 1929.Google Scholar
Clutton-Brock, J., 1989. A re-consideration of the fossil fauna from C-spring, Azraq. In: Copeland, L., Hours, F. (Eds.), The Hammer on the Rock: Studies in the Early Palaeolithic of Azraq, Jordan. British Archaeological Reports (BAR) International Series 540(ii). BAR, Oxford, UK, pp. 391397.Google Scholar
Compagnoni, B., Tosi, M., 1978. The camel: its distribution and state of domestication during the third millennium B.C. in light of finds from Sharh-i Sokhta. In: Meadow, R., Zeder, M.A., (Eds.), Approaches to Faunal Analysis in the Middle East. Peabody Museum of Archaeology and Ethnology Bulletin No. 2. Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge, MA, pp. 91103.Google Scholar
Copeland, L., 1988. Environment, chronology, and Lower-Middle Paleolithic occupations of the Azraq Basin, Jordan. Paléorient 14, 6675.Google Scholar
Copeland, L., 1991. The Late Acheulean knapping-floor at C-Spring, Azraq Oasis, Jordan. Levant 23, 16.Google Scholar
Cordova, C.E., Nowell, A., Ames, C.J.H., Pokines, J.T., DeWitt, R., Al-Souliman, A., 2015. Geoarchaeology of complex stratigraphic contexts in the Paleolithic of the Azraq-Shishan, Jordan: a preliminary reconstruction of paleolandscapes. Poster presented at the XIX INQUA Congress, Nagoya, Japan, July 31, 2015.Google Scholar
Cordova, C.E., Nowell, A., Bisson, M., Ames, C.J.H., Pokines, J., Chang, M., al-Nahar, M., 2013. Interglacial and glacial desert refugia and the Middle Paleolithic of the Azraq Oasis, Jordan. Quaternary International 300, 94110.Google Scholar
Cordova, C.E., Nowell, A., Bisson, M.S., Pokines, J., Ames, C.J.H., al-Nahar, M., 2009. The Druze Marsh Paleolithic Project, North Azraq, Jordan: stratigraphic sequences from the 2008 and 2009 seasons. Annual of the Department of Antiquities of Jordan 53, 311320.Google Scholar
Cordova, C.E., Rollefson, G.O., Kalchgruber, R., Wilke, P., Quintero, L.A., 2008. Natural and cultural stratigraphy of ‘Ayn as-Sawda, Azraq Wetland Reserve: 2007 excavation report and discussion of finds. Annual of the Department of Antiquities of Jordan 52, 417425.Google Scholar
Davies, P., Lister, A.M., 2007. Palaeoloxodon . In: Chazan, M., Horwitz, L.K. (Eds.), Holon: A Lower Paleolithic Site in Israel. American School of Prehistoric Research Bulletin No. 50. Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge, MA, pp. 123131.Google Scholar
Davis, S.J., 1974. Animal remains from the Kebaran site of Ein Gev I, Jordan Valley, Israel. Paléorient 2, 453462.Google Scholar
Davis, S.J., 1980. Late Pleistocene and Holocene equid remains from Israel. Zoological Journal of the Linnean Society 70, 289312.Google Scholar
Dirks, W., 1998. Preliminary account of Pleistocene mammals from ‘Ain Soda, Azraq Basin, Jordan. Journal of Vertebrate Paleontology 18, Supplement 3, 38A39A.Google Scholar
Disi, A.M., Modry, D., Necas, P., Rifai, L., 2001. Amphibians and Reptiles of the Hashemite Kingdom of Jordan: An Atlas and Field Guide. Edition Chimaira, Frankfurt, Germany.Google Scholar
Ducos, P., 1975. A new find of an equid metatarsal bone from Tell Mureibet in Syria and its relevance to the identification of equids from the Early Holocene of the Levant. Journal of Archaeological Science 2, 7173.Google Scholar
Edwards, P.C., 1991. Wadi Hammeh 27: an early Natufian site at Pella, Jordan. In: Bar-Yosef, O., Valla, F.R., (Eds.), The Natufian Culture in the Levant. University of Michigan Press, Ann Arbor, pp. 123148.Google Scholar
Edwards, P.C., Bourke, S.J., Colledge, S.M., Head, J., Macumber, P.G., 1988. Late Pleistocene prehistory in the Wadi al-Hammeh, Jordan Valley. In: Garrard, A.N., Gebel, H.G. (Eds.), The Prehistory of Jordan: The State of Research in 1986. British Archaeological Reports (BAR) International Series 396(2). BAR, Oxford, UK, pp. 525565.Google Scholar
Eisenmann, V., 1986. Comparative osteology of modern and fossil horses, half-asses, and asses. In: Meadow, R.H., Uerpmann, H.-P. (Eds.), Equids in the Ancient World. Beihefte zum Tübinger Atlas des Vorderen Orients, Reihe A, Wiesbaden, Germany, pp. 67116.Google Scholar
Eisenmann, V., Beckouche, S., 1986. Identification and discrimination of metapodials of modern and Pleistocene Equus, wild and domestic. In: Meadow, R.H., Uerpmann, H.-P. (Eds.), Equids in the Ancient World. Beihefte zum Tübinger Atlas des Vorderen Orients, Reihe A, Wiesbaden, Germany, pp. 117163.Google Scholar
El-Naqa, A., 2010. Study of Salt Water Intrusion in the Upper Aquifer in Azraq Basin, Jordan. International Union for the Conservation of Nature (IUCN) and the University of Jordan, Amman.Google Scholar
Fleischer, R.C., Perry, E.A., Muralidharan, K., Stevens, E.E., Wemmer, C.M., 2001. Phylogeography of the Asian elephant (Elephas maximus) based on mitochondrial DNA. Evolution 55, 18821892.Google Scholar
Forsten, A., 1986. A review of the Süssenborn horses and the origin of Equus hydruntinus Regalia. Quartärpalaeontologie 6, 4352.Google Scholar
Fortelius, M., Mazza, P., Sala, B., 1993. Stephanorhinus (Mammalia: Rhinocerotidae) of the Western European Pleistocene, with a revision of S. etruscus (Falconer, 1868). Palaeontographia Italica 80, 63155.Google Scholar
France, R., 2010. Rehabilitation of a historic wetland: Jordan’s Azraq oasis. In: France, R. (Ed.), Restorative Redevelopment of Devastated Ecocultural Landscapes. CRC Press, Boca Raton, FL, pp. 85115.Google Scholar
Garrard, A.N., 1991. Natufian settlement in the Azraq Basin, eastern Jordan. In: Bar-Yosef, O., Valla, F.R. (Eds.), The Natufian Culture in the Levant. University of Michigan Press, Ann Arbor, pp. 235244.Google Scholar
Garrard, A.N., Betts, A., Byrd, B., Hunt, C., Colledge, S., Copeland, L., Montague, R., West, B., 1988a. Summary of palaeoenvironmental and prehistoric investigations in the Azraq Basin. In: Garrard, A.N., Gebel, H.G. (Eds.), The Prehistory of Jordan: The State of Research in 1986. British Archaeological Reports (BAR) International Series 396(1). BAR, Oxford, UK, pp. 311337.Google Scholar
Garrard, A., Colledge, S., Montague, R., Hunt, C., 1988b. Environment and subsistence during the Late Pleistocene and Early Holocene in the Azraq Basin. Paléorient 14, 4049.Google Scholar
Gautier, A., 1966. Camelus thomasi from the northern Sudan and its bearing on the relationship C. thomasi-C. bactrianus . Journal of Paleontology 40, 13681372.Google Scholar
Gauthier-Pilters, H., Dagg, A.I., 1981. The Camel: Its Evolution, Ecology, Behavior, and Relationship to Man. University of Chicago Press, Chicago.Google Scholar
Gillet, H., 1989. The diet of the Arabian oryx (Oryx leucoryx) at the National Wildlife Research Center, Taif. In: Abuzinada, A.H., Goriup, P.D., Nader, I.A. (Eds.), Wildlife Conservation and Development in Saudi Arabia. National Commission for Wildlife Conservation and Development (NCWCD) Publication No.3. NCWCD, Riyadh, Saudi Arabia, pp. 327334.Google Scholar
Google Earth, 2018. Google Earth Pro, version 7.3.1.4507 (accessed March 24, 2018). https://www.google.com/earth/desktop/.Google Scholar
Gordon, C.G., Buikstra, J.E., 1981. Soil pH, bone preservation, and sampling bias at mortuary sites. American Antiquity 46, 566571.Google Scholar
Goren-Inbar, N., Lister, A., Werker, E., Chech, M., 1994. A butchered elephant skull and associated artifacts from the Acheulian site of Gesher Benot Ya’aqov, Israel. Paléorient 20, 99112.Google Scholar
Grantham, B., 1995. Dinner in Buqata: the symbolic nature of food animals and meal sharing in a Druze village. In: Ryan, K., Crabtree, P.J. (Eds.), The Symbolic Role of Animals in Archaeology. MASCA Research Papers in Science and Archaeology 12. MASCA, University Museum of Archaeology and Anthropology, University of Pennsylvania, Philadelphia, pp. 73–78.Google Scholar
Grigson, C., 1983. A very large camel from the Upper Pleistocene of the Negev Desert. Journal of Archaeological Science 10, 311316.Google Scholar
Grigson, C., 2002. Camels, copper and donkeys in the Early Iron Age of the southern Levant: Timna revisited. Levant 44, 82100.Google Scholar
Grigson, C., Gowlett, J.A.J., Zarins, J., 1989. The camel in Arabia—a direct radiocarbon date, calibrated to about 7000 BC. Journal of Archaeological Science 16, 355362.Google Scholar
Groves, C.P., 1983. Phylogeny of the living species of rhinoceros. Zeitschrift fur Zoologische Systematik und Evolutionsforschung 21, 293313.Google Scholar
Groves, C.P., 1986. The taxonomy, distribution, and adaptations of recent equids. In: Meadow R.H., Uerpmann, H.-P. (Eds.), Equids in the Ancient World. Beihefte zum Tübinger Atlas des Vorderen Orients, Reihe A, Wiesbaden, Germany, pp. 1165.Google Scholar
Haas, S.K., Hayssen, V., Krausman, P.R., 2005. Panthera leo. Mammalian Species 762, 111.Google Scholar
Hall, S.J.G., 2008. A comparative analysis of the habitat of the extinct aurochs and other prehistoric mammals in Britain. Ecography 31, 187190.Google Scholar
Harrison, D.L., Bates, P.J.J., 1991. The Mammals of Arabia. 2nd ed. Harrison Zoological Museum, Sevenoaks, UK.Google Scholar
Hedges, R.E.M., Housley, R.A., Law, I.A., Perry, C., Gowlett, J.A.J., 1987. Radiocarbon dates from the Oxford AMS system: archaeometry datelist 6. Archaeometry 29, 289306.Google Scholar
Heide, M., 2011. The domestication of the camel: biological, archaeological and inscriptional evidence from Mesopotamia, Egypt, Israel and Arabia, and literary evidence from the Hebrew Bible. Ugarit-Forschungen 42, 331382.Google Scholar
Henry, D.O., Garrard, A.N., 1988. Tor Hamar: an Epipaleolithic rockshelter in southern Jordan. Palestine Exploration Quarterly 120, 125.Google Scholar
Jagher, R., Elsuede, H., Le Tensorer, J.-M., 2015. El Kowm Oasis, human settlement in the Syrian Desert during the Pleistocene. L’Anthropologie 119, 542580.Google Scholar
Jones, J.R., 2012. Using gazelle dental cementum studies to explore seasonality and mobility patterns of the Early-Middle Epipalaeolithic Azraq Basin, Jordan. Quaternary International 252, 195201.Google Scholar
Jones, M.D., Richter, T., 2011. Paleoclimatic and archeological implications of Pleistocene and Holocene environments in Azraq, Jordan. Quaternary Research 76, 363372.Google Scholar
Jungius, H., 1985. The Arabian oryx: its distribution and former habitat in Oman and its reintroduction. Journal of Oman Studies 8, 4964, plates 1–8.Google Scholar
Kalbe, J., Ames, C., Nowell, A., Cordova, C., Pokines, J.T., 2014. Pleistocene wetlands in the Jordan desert—environmental reconstructions by ostracod and gastropod analysis. Paper presented at the Middle Palaeolithic in the Desert II Conference in Bordeaux, France, December 11, 2014.Google Scholar
Khoury, F., Amr, Z., Hamidan, N., Al Hassani, I., Mir, S., Eid, E., Bolad, N., 2012. Some introduced vertebrate species to the Hashemite Kingdom of Jordan. Vertebrate Zoology 62, 435451.Google Scholar
Klingel, H., 1998. Observations on social organization and behaviour of African and Asiatic Wild Asses Equus africanus and Equus hemionus . Applied Animal Behaviour Science 60, 103113.Google Scholar
Kӧhler, I., 1984. The dromedary in modern pastoral societies and implications for its process of domestication. In: Clutton-Brock, J., Grigson, C. (Eds.), Animals and Archaeology 3: Early Herders and Their Flocks. British Archaeological Reports (BAR) International Series 202. BAR, Oxford, UK, pp. 201206.Google Scholar
Köhler-Rollefson, I., 1991. Camelus dromedarius. Mammalian Species 375, 18.Google Scholar
Köhler-Rollefson, I., 1993. Camels and camel pastoralism in Arabia. Biblical Archaeologist 56, 180188.Google Scholar
Kruuk, H., 1972. The Spotted Hyaena. University of Chicago Press, Chicago.Google Scholar
Kruuk, H., 2002. Hunter and Hunted: Relationships between Carnivores and People. Cambridge University Press, Cambridge.Google Scholar
Lacombat, F., 2006. Morphological and biometrical differentiation of the teeth from Pleistocene species of Stephanorhinus (Mammalia, Perissodactyla, Rhinocerotidae) in Mediterranean Europe and the Massif Central, France. Palaeontographica Abteilung A: Paläozoologie – Stratigraphie 274, 71111.Google Scholar
Lister, A.M., 2013. The role of behaviour in adaptive morphological evolution of African proboscideans. Nature 500, 331334.Google Scholar
Lister, A.M., 2016. Dating the arrival of straight-tusked elephant (Palaeoloxodon spp.) in Eurasia. Bulletin du Musée d’Anthropologie préhistorique de Monaco, année 2015, Supplément n° 6, 123128.Google Scholar
Lister, A.M., Dirks, W., Assaf, A., Chazan, M., Goldberg, P., Applbaum, Y.H., Greenbaum, N., Horwitz, L.K., 2013. New fossil remains of Elephas from the southern Levant: implications for the evolutionary history of the Asian elephant. Palaeogeography, Palaeoclimatology, Palaeoecology 386, 119130.Google Scholar
Lister, A.M., Sher, A.V., 2015. Evolution and dispersal of mammoths across the Northern Hemisphere. Science 350, 805809.Google Scholar
Lister, A.M., Stuart, A.J., 2010. The West Runton mammoth (Mammuthus trogontherii) and its evolutionary significance. Quaternary International 228, 180209.Google Scholar
Lyman, R.L., 1994. Vertebrate Taphonomy. Cambridge University Press, Cambridge.Google Scholar
Maglio, V.J., 1973. Origin and evolution of the Elephantidae. Transactions of the American Philosophical Society 63, 1149.Google Scholar
Marom, N., Bar-Oz, G., 2008. “Measure for measure”: a taphonomic reconsideration of the Kebaran site of Ein Gev I, Israel. Journal of Archaeological Science 35, 214227.Google Scholar
Martin, L., Edwards, Y., Garrard, A., 2010. Hunting practices at an eastern Jordanian Epipalaeolithic aggregation site: the case of Kharaneh IV. Levant 42, 107135.Google Scholar
Martin, L., Edwards, Y., Garrard, A., 2013. Broad spectrum or specialised activity? Birds and tortoises at the Epipaleolithic site of Wadi Jilat 22 in the eastern Jordan steppe. Antiquity 87, 649665.Google Scholar
Martin, L., Edwards, Y., Roe, J., Garrard, A., 2016. Faunal turnover in the Azraq Basin, eastern Jordan 28,000 to 9,000 cal BP, signalling climate change and human impact. Quaternary Research 86, 200219.Google Scholar
Martini, P., Costeur, L., Le Tensorer, J.-M., Schmid, P., 2015. Pleistocene camelids from the Syrian Desert: the diversity in El Kowm. L’Anthropologie 119, 687693.Google Scholar
Martini, P., Geraads, D., 2018. Camelus thomasi Pomel, 1893 from the Pleistocene type-locality Tighennif (Algeria). Comparisons with modern Camelus. Geodiversitas 40, 115134.Google Scholar
Mashkour, M., Monchot, H., Trinkaus, E., Reyss, J.-L., Biglari, F., Bailon, S., Heydari, S., Abdi, K., 2009. Carnivores and their prey in the Wezmeh Cave (Kermanshah, Iran): a Late Pleistocene refuge in the Zagros. International Journal of Osteoarchaeology 19, 678694.Google Scholar
Morel, P., 1997. La faune de la chasse. In: Le Tensorer, J.-M., Muhesen, S. (Eds.), Les Premiers Hommes du Désert Syrien—Fouilles Syro-Suisses à Nadaouiyeh Aïn Askar. Catalogue de l’Exposition, Musée de l’Homme. Editions du Muséum d’Histoire Naturelle, Paris, pp. 4551.Google Scholar
Munro, N.D., Kennerty, M., Meier, J.S., Samei, S., al-Nahar, M., Olszewski, D.I., 2016. Human hunting and site occupation intensity in the Early Epipaleolithic of the Jordanian western highlands. Quaternary International 396, 3139.Google Scholar
Nanda, A.C., 2002. Upper Siwalik mammalian faunas of India and associated events. Journal of Asian Earth Sciences 21, 4758.Google Scholar
Nanda, A.C., 2008. Comments on the Pinjor mammalian fauna of the Siwalik Group in relation to the post-Siwalik faunas of peninsular India and Indo-Gangetic Plain. Quaternary International 192, 613.Google Scholar
Nanda, A.C., 2013. Upper Siwalik mammalian faunas of the Himalayan foothills. Journal of the Palaeontological Society of India 58, 7586.Google Scholar
Nelson, J.B., 1973. Azraq: Desert Oasis. Ohio University Press, Columbus.Google Scholar
Nelson, J.B., 1985. Azraq—a case study. In: Hadidi, A. (Ed.), Studies in the History and Archaeology of Jordan, II. Department of Antiquities, Amman, Jordan, pp. 3944.Google Scholar
Nowell, A., 2014. Reversals of fortune: Neandertals and modern humans in the Levantine Middle Paleolithic, a view from the Druze Marsh, North Azraq (Jordan). In: Finlayson, W., Rollefson, G. (Eds.), Prehistoric Jordan: Recent and Future Research. Department of Antiquities, Amman, Jordan, pp. 2334.Google Scholar
Nowell, A., Walker, C., Cordova, C.E., Ames, C.J.H., Pokines, J.T., Stueber, D., DeWitt, R., al-Souliman, A.S.A., 2016. Middle Pleistocene subsistence in the Azraq Oasis, Jordan: protein residue and other proxies. Journal of Archaeological Science 72, 3644.Google Scholar
Olsen, S.J., 1988. The camel in ancient China and an osteology of the camel. Proceedings of the Academy of Natural Sciences of Philadelphia 140, 1858.Google Scholar
Orlando, L., Mashkour, M., Burke, A., Douady, C.J., Eisenmann, V., Hänni, C., 2006. Geographic distribution of an extinct equid (Equus hydruntinus: Mammalia, Equidae) revealed by morphological and genetical analyses of fossils. Molecular Ecology 15, 20832093.Google Scholar
Osborn, D.J., Helmy, I., 1980. The Contemporary Land Mammals of Egypt (Including Sinai). Fieldiana: Zoology, New Series, No. 5. Field Museum of Natural History, Chicago.Google Scholar
Osborn, H.F., 1942. The Proboscidea: A Monograph of the Discovery, Evolution, Migration and Extinction of the Mastodonts and Elephants of the World. Vol. 2, Stegodontoidea, Elephantoidea. American Museum of Natural History, New York.Google Scholar
Pandolfi, L., Tagliacozzo, A., 2015. Stephanorhinus hemitoechus (Mammalia, Rhinocerotidae) from the Late Pleistocene of Valle Radice (Sora, Central Italy) and re-evaluation of the morphometric variability of the species in Europe. Geobios 48, 169191.Google Scholar
Payne, S., 1983. The animal bones from the 1974 excavations at Douara Cave. Bulletin of the University Museum, University of Tokyo 21, 1108.Google Scholar
Payne, S., Garrard, D., 1983. Camelus from the Upper Pleistocene of Mount Carmel, Israel. Journal of Archaeological Science 10, 243247.Google Scholar
Peters, J., 1998. Camelus thomasi Pomel, 1893, a possible ancestor of the one-humped camel? Zeitschrift für Säugetierkunde 63, 372376.Google Scholar
Peters, J., von den Driesch, A., 1997. The two-humped camel (Camelus bactrianus): new light on its distribution, management and medical treatment in the past. Journal of Zoology, London 242, 651679.Google Scholar
Pokines, J.T., Ames, C.J.H., 2015a. Test excavations at Wadi Zarqa Ma’in 1, a natural sinkhole faunal trap site, Hashemite Kingdom of Jordan. Bulletin of the American Schools of Oriental Research 373, 121137.Google Scholar
Pokines, J.T., Ames, C.J.H., 2015b. Weathering and dispersal of a cattle (Bos taurus) carcass in the desert of eastern Jordan over a six-year interval. Journal of Taphonomy 13, 1731.Google Scholar
Pokines, J.T., Hoskins, J., Ames, C.J.H., Kravitz, G.A., Beck, J., 2014. Taphonomic and paleoecological investigations in the Paleolithic of Northern Jordan. In: Finlayson, W., Rollefson, G. (Eds.), Prehistoric Jordan: Recent and Future Research. Department of Antiquities, Amman, Jordan, pp. 4967.Google Scholar
Rabinovich, R., Hovers, E., 2004. Faunal analysis from Amud Cave: preliminary results and interpretations. International Journal of Osteoarchaeology 14, 287306.Google Scholar
Ramsey, M.N., Jones, M., Richter, T., Rosen, A., 2015. Modifying the marsh: evaluating Early Epipaleolithic hunter-gatherer impacts in the Azraq wetland, Jordan. Holocene 25, 15531564.Google Scholar
Reynaud Savioz, N., Morel, P., 2005. La faune de Nadaouiyeh Aïn Askar (Syrie centrale, Pléistocène moyen): Aperçu et perspectives. Revue de Paléobiologie, Genève 10, 3135.Google Scholar
Richter, T., Maher, L., Stock, J., Allcock, S., Jones, M., Martin, L., Thorne, B., 2009. New light on final Pleistocene settlement diversity in the Azraq Basin (Jordan): recent excavations at ‘Ayn Qasiyya. Paléorient 35, 4968.Google Scholar
Rifai, L.B., Al-Melhim, W.N., Gharaibeh, B.M., Amr, Z.S., 2000. The diet of the Desert Eagle Owl, Bubo bubo ascalaphus, in the Eastern Desert of Jordan. Journal of Arid Environments 44, 369472.Google Scholar
Rivals, F., Lister, A.M., 2016. Dietary flexibility and niche partitioning of large herbivores through the Pleistocene of Britain. Quaternary Science Reviews 146, 116133.Google Scholar
Rollefson, G.O., Quintero, L., Low, R., Schnurrenberger, D., Watson, R., 1996. Paleolithic hunters in the Azraq Oasis. Biblical Archaeologist 59, 239240.Google Scholar
Rollefson, G.O., Quintero, L., Wilke, P., Schnurrenburger, D., Low, R., Watson, R., 1997a. Excavations at ‘Ain Soda in Azraq. Occident and Orient 2, 1617.Google Scholar
Rollefson, G.O., Schnurrenburger, D., Quintero, L.A., Watson, R.P., Low, R., 1997b. ‘Ain Soda and ‘Ain Qasiya: new Late Pleistocene and Early Holocene sites in the Azraq Shishan area, eastern Jordan. In: Gebel, H.G.K., Kafafi, Z., Rollefson, G.O. (Eds.), The Prehistory of Jordan, II. Perspectives from 1997. Studies in Early Near Eastern Production, Subsistence, and Environment 4. Ex Oriente, Berlin, pp. 4558.Google Scholar
Saarinen, J., Karme, A., Cerling, T., Uno, K., Säilä, L., Kasiki, S., Ngene, S., et al., 2015. A new tooth wear-based dietary analysis method for Proboscidea (Mammalia). Journal of Vertebrate Paleontology 35, e918546.Google Scholar
Saegusa, H., Gilbert, W.H., 2008. Elephantidae. In: Gilbert, W.H., Asfaw, B. (Eds.), Homo erectus: Pleistocene Evidence from the Middle Awash, Ethiopia. The Middle Awash Series Vol. 1. University of California Press, Berkeley, pp. 193226.Google Scholar
Sanders, W.J., Gheerbrant, E., Harris, J.M., Saegusa, H., Delmer, C., 2010. Proboscidea. In: Werdelin, L., Sanders, W.J., (Eds.), Cenozoic Mammals of Africa. University of California Press, Berkeley, pp. 161251.Google Scholar
Sapir-Hen, L., Ben-Yosef, E., 2013. The introduction of domestic camels to the southern Levant: evidence from the Aravah Valley. Tel Aviv 40, 277285.Google Scholar
Scates, M.D., 1968. Notes on the hydrobiology of Azraq Oasis, Jordan. Hydrobiologia 31, 7380.Google Scholar
Schaller, G.B., 1972. The Serengeti Lion: A Study of Predator-Prey Relations. University of Chicago Press, Chicago.Google Scholar
Shatnawi, R.S., AlAyyash, S.M., Abdellhadi, A., 2014. Rainy season assessment of Azraq Basin in eastern Jordan. Research Journal of Applied Sciences, Engineering and Technology 8, 120123.Google Scholar
Shehab, A.H., Ciach, M., 2008. Diet composition of the Pharaoh Eagle Owl, Bubo ascalaphus, in Azraq Nature Reserve, Jordan. Turkish Journal of Zoology 32, 6569.Google Scholar
Shoshani, J., Eisenberg, J.F., 1982. Elephas maximus. Mammalian Species 182, 18.Google Scholar
Simmons, A.H., Rollefson, G.O., Kafafi, Z., Mandel, R.D., al-Nahar, M., Cooper, J., Köhler-Rollefson, I., Durand, K.R., 2001. Wadi Shuʿeib, a large Neolithic community in central Jordan: final report of test investigations. Bulletin of the American Schools of Oriental Research 321, 139.Google Scholar
Smuts, M.M.S., Bezuidenhout, A.J., 1987. Anatomy of the Dromedary. Clarendon, Oxford, UK.Google Scholar
Spassov, N., Stoytchev, T., 2004. The dromedary domestication problem: 3000 BC rock art evidence for the existence of wild one-humped camel in Central Arabia. Historia Naturalis Bulgarica 16, 151158.Google Scholar
Stanley, H.F., Kadwell, M., Wheeler, J.C., 1994. Molecular evolution of the family Camelidae: a mitochondrial DNA study. Proceedings of the Royal Society of London B: Biological Sciences 256, 16.Google Scholar
Steiger, C., 1990. Vergleichend Morphologische Untersuchungen an Einzelknochen des Postcranialen Skeletts der Atlweltkamele. PhD dissertation, Universität München, Munich, Germany.Google Scholar
Stewart, M., Louys, J., Price, G.J., Drake, N.A., Groucutt, H.S., Petraglia, M.D., 2017. Middle and Late Pleistocene mammal fossils of Arabia and surrounding regions: implications for biogeography and hominin dispersals. Quaternary International (in press). https://doi.org/10.1016/j.quaint.2017.11.052.Google Scholar
Stimpson, C.M., Lister, A., Parton, A., Clark-Balzan, L., Breeze, P.S., Drake, N.A., Groucutt, H.S., et al., 2016. Middle Pleistocene vertebrate fossils from the Nefud Desert, Saudi Arabia: implications for biogeography and palaeoecology. Quaternary Science Reviews 143, 1336.Google Scholar
Stiner, M.C., 2005. The Faunas of Hayonim Cave, Israel: A 200,000-Year Record of Paleolithic Diet, Demography, and Society. American School of Prehistoric Research Bulletin No. 48. Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge, MA. Google Scholar
Stiner, M.C., Barkaib, R., Gopher, A., 2009. Cooperative hunting and meat sharing 400–200 kya at Qesem Cave, Israel. Proceedings of the National Academy of Sciences of the United States of America 106, 1320713212.Google Scholar
Tchernov, E., 1993. Exploitation of birds during the Natufian and Early Neolithic of the southern Levant. Archaeofauna 2, 121143.Google Scholar
Todd, N.E., 2005. Reanalysis of African Elephas recki: implications for time, space and taxonomy. Quaternary International 126–128, 6572.Google Scholar
Turnbull, P.F., Reed, C.A., 1974. The fauna from the Terminal Pleistocene of Palegawra Cave, a Zarzian occupation site in Northeastern Iraq. Fieldiana Anthropology 63, 81146.Google Scholar
Turnbull, P.M., 1989. Camel, canid, and other animal remains from Ain El-Assad (Lion’s Spring), Jordan. In: Copeland, L., Hours, F. (Eds.), The Hammer on the Rock: Studies in the Early Palaeolithic of Azraq, Jordan. British Archaeological Reports (BAR) International Series 540(ii). BAR, Oxford, UK, pp. 277299.Google Scholar
Tyrberg, T., 1998. Pleistocene Birds of the Palearctic: A Catalogue. Nuttall Ornithological Club, Cambridge, MA.Google Scholar
Uerpmann, H.-P., 1987. The Ancient Distribution of Ungulate Mammals in the Near East: Fauna and Archaeological Sites in Southwest Asia and Northeast Africa. Reichert, Wiesbaden, Germany.Google Scholar
Uerpmann, H.-P., Uerpmann, M., 2002. The appearance of the domestic camel in south-east Arabia. Journal of Oman Studies 12, 235260.Google Scholar
van Asperen, E.N., Kahlke, R.-D., 2015. Dietary variation and overlap in central and northwest European Stephanorhinus kirchbergensis and S. hemitoechus (Rhinocerotidae, Mammalia) influenced by habitat diversity. Quaternary Science Reviews 107, 4761.Google Scholar
van Vuure, T., 2003. History, morphology and ecology of the aurochs (Bos primigenius). Lutra 45, 16.Google Scholar
Vidya, T.N.C., Sukumar, R., Melnick, D.J., 2009. Range-wide mtDNA phylogeography yields insights into the origins of Asian elephants. Proceedings of the Royal Society B: Biological Sciences 276, 893902.Google Scholar
Villwock, W., Scholl, A., Krupp, F., 1983. Zur Taxonomie, Verbreitung und Speziation des Formenkreises Aphanius dispar (Rüppell, 1828) und Beschreibung von Aphanius sirhani n. sp. (Pisces: Cyprinodontidae). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 80, 251277.Google Scholar
von den Driesch, A., 1976. A Guide to the Measurement of Animal Bones from Archaeological Sites. Peabody Museum of Archaeology and Ethnology Bulletin No. 1. Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge, MA.Google Scholar
von den Driesch, A., Bruckner, H., Obermaier, H., Zander, A., 2008. The hunt for wild dromedaries during the 3rd and 2nd millennia BC on the United Emirates coast. Camel bone finds from the excavations at Al Sufouh 2, Dubai, UAE. In: Vila, E., Gourichon, L., Choyke, A.M., Buitenhuis, H. (Eds.), Archaeozoology of the Near East VIII. Maison de l’Orient et de la Méditerrannée, Lyon, France, pp. 487497.Google Scholar
von den Driesch, A., Obermaier, H., 2007. The hunt for wild dromedaries during the 3rd and 2nd millennia BC on the United Emirates coast: camel bone finds from the excavations at al-Sufouh 2, Dubai, UAE. Documenta Archaeobiologiae 5, 133167.Google Scholar
Walker, C., 2017. Dining out in the desert: results from protein residue analysis at the Azraq Oasis, Jordan. Paper presented at the 82nd Annual Meeting of the Society for American Archaeology, Vancouver, BC, Canada, March 30, 2017.Google Scholar
Wapnish, P., 1984. The dromedary and bactrian camel in Levantine historical settings: the evidence from Tell Jemmeh. In: Clutton-Brock, J., Grigson, C. (Eds.), Animals and Archaeology 3: Early Herders and Their Flocks. British Archaeological Reports (BAR) International Series 202. BAR, Oxford, UK, pp. 171200.Google Scholar
Watson, R.P., Burnett, G.W., 2001. On the origins of Azraq’s “Roman Wall.” Near Eastern Archaeology 64, 7279.Google Scholar
Woolfenden, W.B., Ababneh, L., 2011. Late Holocene vegetation in the Azraq Wetland Reserve, Jordan. Quaternary Research 76, 345351.Google Scholar
Zaidner, Y., 2013. Adaptive flexibility of Oldowan hominins: secondary use of flakes at Bizat Ruhama, Israel. PLoS ONE 8,e66851.Google Scholar
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