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layer 17 of the archaeological site of Hummal (El Kowm, Central Syria): Preliminary results
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Original article
Microvertebrates from unit G/layer 17 of the archaeological site of Hummal (El Kowm, Central Syria): Preliminary results Les microvertébrés de l’unité G/couche 17 du site archéologique de Hummal (El Kowm, Syrie centrale) : résultats préliminaires Lutz Christian Maul a,*, Krister T. Smith b, Georgy Shenbrot c, Angela A. Bruch d, Fabio Wegmüller e, Jean-Marie Le Tensorer e a Senckenberg Research Station of Quaternary Palaeontology, Weimar, Germany Senckenberg Research Institute and Natural History Museum, Frankfurt/Main, Germany c Ben-Gurion University of the Negev, Midreshet Ben-Gurion, Israel d ROCEEH, Senckenberg Research Institute and Natural History Museum, Frankfurt/Main, Germany e Institute for Prehistory and Archaeological Science, University of Basel, Basel, Germany b
Available online 14 November 2015
Abstract We present a first look at the microvertebrate fauna of the Middle Pleistocene site of Hummal in Central Syria. Some 2000 microvertebrate remains (1200 mammalian; 230 reptilian; 600 unidentified) were found in unit G/layer 17 by screen-washing sediments in an area of 4 m2. The following taxa have been identified: Reptilia: Agaminae indet., Gekkota indet., Lacertidae indet. (2–3 taxa), Eryx sp., Natricinae indet.; and Mammalia: Crocidurinae indet., Chiroptera indet., Lepus sp., Arvicolinae indet., Ellobius sp., Microtus sp., Murinae indet. (large form), Mus sp., Meriones sp., Gerbillus sp. The presence of Ellobius indicates a Middle Pleistocene age of the fauna. This genus does not occur in Syria today, but is recorded in Israel in Late Acheulean to Early Mousterian sites. In North Africa, the occurrence of this genus is restricted to the early part of the Middle Pleistocene. The ecological requirements of the nearest living relatives of the recorded taxa indicate mainly open habitats, but also the presence of vegetation and wet conditions, at least close to the site. Lepus, Ellobius, Meriones, Gerbillus, and Eryx live to various extents in steppes, semideserts, and deserts. Various extant species of large murids that come into consideration occupy a variety of wooded habitats, grassland and savannah and require the presence of water. Hummal offers the most
* Corresponding author. E-mail address: [email protected] (L.C. Maul). http://dx.doi.org/10.1016/j.anthro.2015.10.010 0003-5521/# 2015 Elsevier Masson SAS. All rights reserved.
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detailed Pleistocene portrait yet of hominin palaeoenvironment in an area that, at the end of the Pleistocene, became known as part of the Fertile Crescent. # 2015 Elsevier Masson SAS. All rights reserved. Keywords: Microvertebrate fauna; Palaeoenvironment; Levant; El Kowm; Hummal; Unit G
Résumé Dans cet article, nous présentons un premier aperçu de la faune des microvertébrés du site Pléistocène moyen de Hummal en Syrie centrale. Environ 2000 restes de microvertébrés (1200 mammifères ; 230 reptiles ; 600 non identifiés) ont été récoltés par tamisage à l’eau des sédiments d’une surface de 4 m2 dans l’unité G/couche 17. Les taxons suivants ont été identifiés : Reptilia : Agaminae indet., Gekkota indet., Lacertidae indet. (2–3 taxa), Eryx sp., Natricinae indet. ; et Mammalia : Crocidurinae indet., Chiroptera indet., Lepus sp., Arvicolinae indet., Ellobius sp., Microtus sp., Murinae indet. (grande espèce), Mus sp., Meriones sp., Gerbillus sp. La présence de Ellobius indique un âge Pléistocène moyen de la faune. Ce genre n’est pas connu aujourd’hui en Syrie, mais est représenté en Israël dans les sites de l’Acheuléen supérieur au début du Moustérien. En Afrique du Nord, ce genre est limité à la partie la plus ancienne du Pléistocène moyen. Les exigences écologiques espèces apparentées les plus proches des taxons enregistrés indiquent surtout des habitats ouverts, mais aussi la présence de végétation et de conditions humides, au moins à proximité du site. Lepus, Ellobius, Meriones, Gerbillus et Eryx vivent à des degrés divers dans les steppes, les semi-déserts et les déserts. Les diverses espèces existantes de grands muridés occupent une grande variété d’habitats boisés, prairies et savanes et ont besoin de la présence d’eau. Hummal offre le cadre le plus détaillé du paléoenvironnement humain au Pléistocène dans une région qui, à la fin du Pléistocène, est considérée comme une partie du Croissant fertile. # 2015 Elsevier Masson SAS. Tous droits réservés. Mots clés : Faune de microvertébrés ; Paléoenvironnement ; El Kowm ; Hummal ; Unité G
1. Introduction Microvertebrates are rare in Syrian Pleistocene sites. That is unfortunate, since these kinds of fossils are particularly suitable for palaeoecological reconstruction and biostratigraphical inference. The small scale of typical terrestrial microvertebrate habitats allows inferences about palaeoenvironment very close to the site of deposition. The suitability of small mammals, in particular, for biostratigraphic age estimation results from their rather high evolutionary rates; for this reason, the identified evolutionary stages allow detailed age determination. In other parts of the Near East, particularly in Israel, the Pleistocene record of microvertebrates is much more dense (for a compilation see Tchernov, 1996 and references therein). On the other hand, there is only one sufficiently large sample is known from Syria: Late Pleistocene Douara Cave near Palmyra (Payne, 1983). We report on a concentration of microfaunal remains unearthed in a small excavation area (unit G, layer 17) of the archaeological site of Hummal. These remains occur together with numerous crushed and fragmented large bones (Wegmüller, 2011; Wegmüller et al., 2012). The site of Hummal is located in the El Kowm oasis in the desert steppe of Central Syria, a key area for the Palaeolithic in the Levant (Jagher and Le Tensorer, 2011; Le Tensorer et al., 2011; Jagher this volume). The sequence begins in the Lower Palaeolithic and concludes in the Upper
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Palaeolithic (Le Tensorer et al., 2011). In this paper, we present preliminary results of our investigations of the microvertebrates from Hummal and their palaeoecological and biostratigraphical implications. 2. Material and methods All microvertebrate remains mentioned here originate from an area of 4 m2 in layer 17 of unit G at Hummal (for details see Wegmüller, 2011; Wegmüller et al., 2012). This layer is situated c. 12.5 m below the surface. It is c. 10–15 cm thick and consists of dark grey to black compact clay. The precompactional thickness of the sediment layer was presumably greater. The layer is also rich in bone fragments of large mammals but poor in artefacts. It conformably overlies layer 18, a level very rich in artefacts. All layers superjacent to layer 17 are of Yabrudian, Hummalian or Mousterian age. Generally speaking, the lower part of the profile (layers 15–23) has yielded archaic artefacts, some of which are comparing to those from Ubeidiya in Israel (Wegmüller, this volume). More than 2000 microvertebrate remains was extracted by screen-washing of the sediments. Sixty percent of the remains clearly belong to small mammals, and ten percent to small reptiles, while 600 bones are of small vertebrates but cannot be identified more precisely. Many small mammal bone fragments are part of post-cranial elements. In general, these remains are unsuitable for identification to a fine taxonomic level. The incisors of rodents were merely identified as belonging to the order Rodentia. In the following descriptions, lower case denotes lower teeth and upper case denotes upper teeth. Dental morphological nomenclature of murines is after Vandebroek (1961–1962), that of arvicolines after van der Meulen (1973). 3. Taxonomy Class REPTILIA Laurenti, 1768 Order SQUAMATA Oppel, 1811 Suborder IGUANIA Cope, 1864 Family AGAMIDAE Fitzinger, 1826 Subfamily AGAMINAE Fitzinger, 1826 sensu Macey et al., 2000 AGAMINAE indet. Twenty-three cranial specimens, including maxillae (Fig. 1), ectopterygoids, quadrates, dentaries and coronoids, as well as a number of presacral and caudal vertebrae, are attributed to an agamine lizard. Caudal vertebrae lack autotomy planes, as in all members of Acrodonta (Etheridge, 1967), and the cheek tooth series includes two pleurodont tooth loci in the maxilla followed by a series of acrodont teeth, which is frequently interpreted as an apomorphy of Agamidae. The remains furthermore show characters of the African/West Asian clade Agaminae (sensu Macey et al., 2000), which have been interpreted as apomorphies (Maul et al., 2011): facial process (as indicated by its broken base) sharply folded medially along an oblique, posterodorsally trending axis to create a distinct anterodorsally directed surface behind the external nares; two pleurodont tooth loci in both maxilla and dentary; and acrodont teeth simple, triangular and unicuspid (also known in Hydrosaurus and Amphibolurinae). Presently available evidence on the identity of the Hummal agamid is therefore meagre but suggestive of Agaminae. Most members of that clade – especially those taxa found in the area
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today, including species of Trapelus (Wagner et al., 2011) – are considered ‘‘semiclimbers’’ and typical inhabitants of warm, temperate environments (Moody, 1980). Suborder GEKKOTA Cuvier, 1817 GEKKOTA indet. The most abundant squamate taxon at Hummal is a single species of gecko. Prefrontals and frontals are particularly common, but quadrates, compound bones, and dentigerous elements (Fig. 1) are also present. Derived gekkotan characters found in this species (see Estes et al., 1988; Gauthier et al., 2012) include: azygous frontal, subolfactory processes of frontal meet and fuse below olfactory tracts, and fused Meckelian groove. Comparative specimens of Near Eastern geckos are lamentably rare, precluding a more precise identification at this time, despite the propitiousness of the material. Suborder LACERTIFORMES Estes et al., 1988 Family LACERTIDAE Gray, 1825 LACERTIDAE indet. (at least two species) Forty-nine cranial bones, especially maxilla and dentary fragments (Fig. 1) but also frontals, prefrontals, an ectopterygoid (?), a jugal, postorbitofrontals, a quadrate, and a coronoid, are referred to Lacertidae. The frontals are ayzgous; the orbital margins show a narrow band where the supraocular osteoderms articulated. The postorbitofrontal is medially expanded, showing partial closure of the supratemporal fenestra. The exterior surfaces of the cranial bones (generally fused osteoderms) are sculptured. Jaws are the most abundant elements, and the posterior teeth are bicuspid. Tooth morphology and jaw size suggest that at least two species are represented. Suborder SERPENTES Linnaeus, 1758 Family BOIDAE Gray, 1825 Genus ERYX Daudin, 1802 Eryx sp. Three trunk vertebrae, a cloacal vertebra, and ten caudal vertebrae are attributed to a single species of the Old World sand boa clade Eryx. The trunk vertebrae are anteroposteriorly short, as in Boidae, and have low, restricted neural spines and somewhat depressed neural arches, as in Erycinae (Hoffstetter and Rage, 1972). The most diagnostic, and curiously also the most abundant, specimens are caudal vertebrae (Fig. 1) that evince the bifurcated neural spine and complicated accessory processes typical of erycine boids (Rage, 1974; Szyndlar and Rage, 2003; Smith, 2013). The more distal caudals are anteroposteriorly very short and tall, and the zygosphene-zygantral articulations are absent, as in Eryx and Bransapteryx (Szyndlar, 1994). As in most Eryx, a direct connection between the pre-zygapophyses and post-zygapophyses has been lost (Szyndlar, 1994; Szyndlar and Schleich, 1994). Further detailed studies of Eryx, incorportating the most recent results on phylogenetic structure within the taxon (Reynolds et al., 2014), will be necessary for an apomorphy-based identification of the Hummal species, but its attribution to Eryx appears secure. Only Eryx jaculus is known from Syria today. This species has a patchy distribution in North Africa, the Levant, and southeastern Europe, but does not presently occur in the hyperarid Syrian plains in which Hummal is situated. Szyndlar and Schleich (1994) previously described uppermost Miocene material from the south of Spain as Eryx cf. jaculus, postulating a formerly circum-Mediterranean distribution. Family COLUBRIDAE Oppel, 1811 sensu Lawson et al., 2005
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Subfamily NATRICINAE Bonaparte, 1838 NATRICINAE indet. Thirty elongate, delicate vertebrae appear to represent a single species of Natricinae. They all possess at least small hypapophyses, which on more posterior trunk vertebrae are short, pointed, and posteriorly directed. Class MAMMALIA Linnaeus, 1758 Some 700 post-cranial bone fragments are typical in size and shape of small mammals, but they have not been identified more precisely. Order SORICOMORPHA Gregory, 1910 Family SORICIDAE Fischer von Waldheim, 1814 Subfamily CROCIDURINAE Milne-Edwards, 1872 CROCIDURINAE indet. Despite its incompleteness, including loss of the processus articularis, a mandible with m1 and m2 and three isolated teeth clearly belong to a white-toothed shrew (subfamily Crocidurinae). Possible candidates are Suncus etruscus (Savi, 1822), Crocidura leucodon (Hermann, 1780), C. suaveolens (Pallas, 1811), C. katinka Bate, 1937, and C. caspica Thomas, 1907. However, for the moment, we cannot assign the fragments to a particular species or genus. Order CHIROPTERA Blumenbach, 1779 CHIROPTERA indet. Three molar fragments display characters diagnostic of bats: high cusps, crowns oblique in a longitudinal direction, and a strong cingulum. However, because of the fragmentary nature of the specimens, they cannot currently be referred to a particular taxon. Order LAGOMORPHA Brandt, 1855 Family LEPORIDAE Fischer von Waldheim, 1817 Genus LEPUS Linnaeus, 1758 Lepus sp. Several cheek teeth (two P2, two p3, two M) are characteristic of leporids. The diagnostic p3 (Fig. 2A) and P2 indicate the presence of the genus Lepus. The occlusal pattern with small buccal and lingual folds is similar to that of L. capensis Linnaeus, 1758, and the size of the teeth also
Fig. 1. Selected small reptile remains from Hummal. A. Partial left maxilla of Agaminae indet. (HERP3-c2). B. Partial left dentary of Gekkota indet. (HERP2-f4). C. Partial right dentary of Lacertidae indet. (HERP2-a10). D. Distal caudal vertebra of Eryx sp. (HERP1-a11). Sélection de restes de petits reptiles de Hummal. A. Portion de maxillaire gauche d’Agaminae indet. (HERP3-c2). B. Portion dentaire gauche de Gekkota indet. (HERP2-f4). C. Portion dentaire droite de Lacertidae indet. (HERP2-a10). D. Vertèbre caudale distale d’Eryx sp. (HERP1-a11).
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matches this species, which is smaller than L. europaeus Pallas, 1778. However, there is another small species, L. tolai Pallas, 1778, which is distributed in central Asia, westwards up to the Caspian Sea, and in north-eastern Asia. Due to the small number of specimens in Hummal and overlap in size, we tentatively refer the material to Lepus sp. Order RODENTIA Bowdich, 1821 RODENTIA indet. There are some 200 incisors of rodents, which have not been referred to the family or genus level. Family CRICETIDAE Fischer von Waldheim, 1817 Subfamily ARVICOLINAE Gray, 1821 ARVICOLINAE indet. Five molar fragments, consisting only of one or two prisms, can be identified only to the subfamily level. Genus ELLOBIUS Fischer von Waldheim, 1814 Ellobius sp. Forty molars (nine m1, nine m2, three m3, eleven M1, four M2, and four M3) of arvicoline rodents show typical traits of the genus Ellobius, or mole voles. The m1s possess three buccal and four lingual salient angles and have rather broad anterior caps (Fig. 2D). The dentine fields are broadly confluent and there is no crown cementum in the re-entrant angles. There are five extant species of Ellobius: E. talpinus (Pallas, 1770), E. tancrei Blasius, 1884, E. alaicus Vorontsov et al., 1969, E. fuscocapillus Blyth, 1843, and E. lutescens Thomas, 1897. Mole voles are not part of the recent fauna of Syria, but the geographically closest species E. lutescens occurs in northwestern Iran, eastern Turkey and southern Azerbaijan. It resembles our finds also morphologically more than to the other species. The m1 of E. lutescens possesses a broad anterior cap, whereas that of E. talpinus is narrow and that of E. fuscocapillus elongated. However, also E. pedorychus is rather similar to our finds. It was described by Bate (1937), from the Israeli Middle Pleistocene sites of Tabun C, D, and E. In the Middle Pleistocene, Ellobius is also represented in North Africa (Jaeger, 1988; Geraads, 2010; Stoetzel, 2013, etc.) by the following species: E. africanus Jaeger, 1988, E. atlanticus Jaeger, 1988, E. zimae Jaeger, 1988, and E. barbarus (Pomel, 1892). E. barbarus is more similar to E. talpinus, whereas E. africanus and E. atlanticus show resemblances to our finds in the anteroconid of m1. Thus, since the specimens from Hummal resemble various species – the extant E. lutescens, as well as the fossil E. pedorychus, E. africanus, and E. atlanticus – we tentatively refer them only to Ellobius sp. Genus MICROTUS Schrank 1798 Microtus sp. A single m1 fragment belongs to Microtus. It has no roots and there is crown cementum in the re-entrant angles. The dentine fields T4 and T5 are closed, the anterior cap has an ‘arvaline’ shape, i.e. it resembles that of Microtus arvalis. The enamel differentiation is Microtus-like. However, given the fragmentary preservation, the specimen is referred only to Microtus sp. Family MURIDAE, Illiger, 1811 Subfamily MURINAE Illiger, 1811 MURINAE indet. An M2 belongs to a member of the subfamily Murinae that is considerably larger than corresponding molars in other large-sized murines, like Rattus norvegicus (Berkenhout, 1769)
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and even larger than the corresponding molar of the Pest Rat Nesokia indica (Gray, 1830), which is more hypsodont and has fused tubercles (Niethammer, 1977). There are resemblances in size and shape to the M2 of Bandicota indica Gray, 1873. However, a reliable assignment requires further comparisons with larger samples. Genus MUS Linnaeus, 1758 Mus sp. Based on its size and the lack of tubercle t9 (Reichstein, 1978), one left M1 can be identified as belonging to Mus. Considering the distribution of extant species, it is most probably M. macedonicus Petrov & Ruzic, 1983. Since, however, it is only one specimen we refer it to as Mus sp. Subfamily GERBILLINAE Gray, 1825 Genus MERIONES Illiger, 1811 Meriones sp.and Genus GERBILLUS Desmarest 1804 Gerbillus sp. Some 250 molars and molar fragments can be assigned to the subfamily Gerbillinae. Among them, eight m1 and M1 belong to Gerbillus and 60 to Meriones. Species of the latter genus have larger and more hypsodont molars, the lateral incision of enamel (linea sinuosa) is more oblique distally, the outline of m1 is less asymmetric, and the anterior cap is broader than in molars of Gerbillus. In morphology and size, the finds are consistent with Meriones tristrami and Gerbillus dasyurus. The frequency distribution of m1 length, measured at the base of the crown indicates that the gerbilline finds belong to more than two species. Considering the distribution of extant species and size, possible candidates would be Meriones libycus, M. vinogradovi, and M. crassus on one hand and Gerbillus dasyurus, G. mesopotamiae, G. henleyi or G. cheesmani on the other. Morphologically they can hardly be distinguished. Therefore, we refer the Gerbillinae from Hummal tentatively only to the genus level (Fig. 2B and C).
Fig. 2. Selected small mammal remains from Hummal. A. Right p4 of Lepus sp. (MAMe3). B. Left m1 (invers) of Meriones sp. (GER1a1). C. Right m1 of Gerbillus sp. (GER1b6). D. Left m1 fragment (invers) of Ellobius sp. (MAMa11). Sélection de restes de petits mammifères de Hummal. A. p4 droite de Lepus sp. (MAMe3). B. m1 gauche (inversée) de Meriones sp. (GER1a1). C. m1 droite de Gerbillus sp. (GER1b6). D. Fragments de m1 gauche (inversé) de Ellobius sp. (MAMa11).
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4. Palaeoecology Based on the ecological requirements of the nearest living relatives, the recorded taxa suggest various open habitats, but also the presence of vegetation and wet areas, at least close to the site. Species of the genera Lepus, Ellobius, Meriones, Gerbillus, and Eryx live to various extents in grasslands, in steppes, semi-deserts, and deserts respectively (Harrison and Bates, 1991; Disi et al., 2001; Cos¸kun and Ulutürk, 2003; Krapp, 2003). If the referral of the large murid molar to Bandicota will hold true, this would indicate the occurrence of streams, other water bodies or swampy areas (http://www.iucnredlist.org/details/2541/0). Preliminary result using the coexistence approach (Mosbrugger and Utescher, 1997; Maul et al., in press) for a dataset of various recent close relatives of the recorded taxa, show higher precipitation values but similar temperatures in comparison with today. However, to obtain reliable results more detailed taxonomical identifications are required. Layer 17 as well as most of the other archaeological layers in Hummal were mainly formed during active episodes of the artesian spring (Le Tensorer et al., 2007) and therefore, the presence of water in direct vicinity of the site must be assumed. 5. Biostratigraphy In Plio- and Pleistocene Palaearctic sites, arvicolines generally allow best biostratigraphic estimation. At Hummal, these would be Ellobius und Microtus. Whereas Microtus today is nearly invariable element of faunas in the Near East, Ellobius is not a part of the extant fauna of the Levant. There are fossil records of Ellobius both from the Near East and northern Africa (for compilations see Tchernov, 1996; Stoetzel, 2013). All these finds are exclusively of Middle Pleistocene age. In Israel, the finds range from Late Acheulean (Oumm Qatafa) to Early Mousterian (Tabun D and Hayonim E). In North Africa, Ellobius is recorded only in the older part of the Middle Pleistocene, between c. 800 ka and 370 ka (Jaeger, 1988; Geraads et al., 2013). However, in the Levant, it is also present in slightly younger sites, as Tabun D, and C (Mousterian) (Bate, 1937), but not in the 780–700 ka old Gesher Benot Ya’aqov (Goren-Inbar et al., 2000). Thus, we could conclude for Hummal an age between early Middle Pleistocene and Early Mousterian, an age range between 780 and 165 ka. However, detailed studies of the Ellobius finds will provide further refinement of the biostratigraphic age of this assemblage. 6. Taphonomy According to the geological observation of the excavators, the layer experienced heavy sedimentary pressure. This is possibly one reason for the fragmentary nature of the remains. In addition, if the fragmentation is partly predepositional, it could suggest that the microvertebrates were accumulated as prey remains. The question of whether owls, diurnal raptors or mammalian carnivores are responsible for the accumulation is the subject of ongoing investigations. 7. Conclusion Preliminary investigation of the microvertebrates from layer 17 of unit G at Hummal provide taxonomic, biostratigraphic and palaeoecological information about the fauna and its palaeoenvironment. Our age estimate for Hummal layer 17 – from early Middle Pleistocene to Mousterian – is broadly consistent with the fact that layer 17 is subjacent to Yabrudian, Hummalian and
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Mousterian horizons. This estimate is based on the regional occurrence of Ellobius, which is not part of the extant fauna of Syria. However, the reason for the expansion and shrinking of the geographic distribution of Ellobius is thought to be quite complex. The geographic range of extant E. lutescens in eastern Turkey is restricted by topography and by the occurrence of Spalax rather than by temperature, rainfall or humidity (Cos¸kun and Ulutürk, 2003). It can be only speculated that ancient populations of Ellobius in Israel and Syria were relatively small and isolated. Probably, after their extirpation, recolonization was prevented because dispersal routes were blocked by competing Spalax and Nesokia. Current coexistence of Ellobius and other fossorial forms, such as Spalax, Nesokia and Myospalax is really marginal or even exclusive. The microfauna from Hummal is of great importance for the faunal history of this region since we have very few records from this area and time period. Unfortunately, excavations have been suspended due to armed conflict in the region. However, a detailed investigation of the microvertebrate material presented here, including a more extensive taxonomic analysis, is under way.
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Original article
Microvertebrates from unit G/layer 17 of the archaeological site of Hummal (El Kowm, Central Syria): Preliminary results Les microvertébrés de l’unité G/couche 17 du site archéologique de Hummal (El Kowm, Syrie centrale) : résultats préliminaires Lutz Christian Maul a,*, Krister T. Smith b, Georgy Shenbrot c, Angela A. Bruch d, Fabio Wegmüller e, Jean-Marie Le Tensorer e a Senckenberg Research Station of Quaternary Palaeontology, Weimar, Germany Senckenberg Research Institute and Natural History Museum, Frankfurt/Main, Germany c Ben-Gurion University of the Negev, Midreshet Ben-Gurion, Israel d ROCEEH, Senckenberg Research Institute and Natural History Museum, Frankfurt/Main, Germany e Institute for Prehistory and Archaeological Science, University of Basel, Basel, Germany b
Available online 14 November 2015
Abstract We present a first look at the microvertebrate fauna of the Middle Pleistocene site of Hummal in Central Syria. Some 2000 microvertebrate remains (1200 mammalian; 230 reptilian; 600 unidentified) were found in unit G/layer 17 by screen-washing sediments in an area of 4 m2. The following taxa have been identified: Reptilia: Agaminae indet., Gekkota indet., Lacertidae indet. (2–3 taxa), Eryx sp., Natricinae indet.; and Mammalia: Crocidurinae indet., Chiroptera indet., Lepus sp., Arvicolinae indet., Ellobius sp., Microtus sp., Murinae indet. (large form), Mus sp., Meriones sp., Gerbillus sp. The presence of Ellobius indicates a Middle Pleistocene age of the fauna. This genus does not occur in Syria today, but is recorded in Israel in Late Acheulean to Early Mousterian sites. In North Africa, the occurrence of this genus is restricted to the early part of the Middle Pleistocene. The ecological requirements of the nearest living relatives of the recorded taxa indicate mainly open habitats, but also the presence of vegetation and wet conditions, at least close to the site. Lepus, Ellobius, Meriones, Gerbillus, and Eryx live to various extents in steppes, semideserts, and deserts. Various extant species of large murids that come into consideration occupy a variety of wooded habitats, grassland and savannah and require the presence of water. Hummal offers the most
* Corresponding author. E-mail address: [email protected] (L.C. Maul). http://dx.doi.org/10.1016/j.anthro.2015.10.010 0003-5521/# 2015 Elsevier Masson SAS. All rights reserved.
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detailed Pleistocene portrait yet of hominin palaeoenvironment in an area that, at the end of the Pleistocene, became known as part of the Fertile Crescent. # 2015 Elsevier Masson SAS. All rights reserved. Keywords: Microvertebrate fauna; Palaeoenvironment; Levant; El Kowm; Hummal; Unit G
Résumé Dans cet article, nous présentons un premier aperçu de la faune des microvertébrés du site Pléistocène moyen de Hummal en Syrie centrale. Environ 2000 restes de microvertébrés (1200 mammifères ; 230 reptiles ; 600 non identifiés) ont été récoltés par tamisage à l’eau des sédiments d’une surface de 4 m2 dans l’unité G/couche 17. Les taxons suivants ont été identifiés : Reptilia : Agaminae indet., Gekkota indet., Lacertidae indet. (2–3 taxa), Eryx sp., Natricinae indet. ; et Mammalia : Crocidurinae indet., Chiroptera indet., Lepus sp., Arvicolinae indet., Ellobius sp., Microtus sp., Murinae indet. (grande espèce), Mus sp., Meriones sp., Gerbillus sp. La présence de Ellobius indique un âge Pléistocène moyen de la faune. Ce genre n’est pas connu aujourd’hui en Syrie, mais est représenté en Israël dans les sites de l’Acheuléen supérieur au début du Moustérien. En Afrique du Nord, ce genre est limité à la partie la plus ancienne du Pléistocène moyen. Les exigences écologiques espèces apparentées les plus proches des taxons enregistrés indiquent surtout des habitats ouverts, mais aussi la présence de végétation et de conditions humides, au moins à proximité du site. Lepus, Ellobius, Meriones, Gerbillus et Eryx vivent à des degrés divers dans les steppes, les semi-déserts et les déserts. Les diverses espèces existantes de grands muridés occupent une grande variété d’habitats boisés, prairies et savanes et ont besoin de la présence d’eau. Hummal offre le cadre le plus détaillé du paléoenvironnement humain au Pléistocène dans une région qui, à la fin du Pléistocène, est considérée comme une partie du Croissant fertile. # 2015 Elsevier Masson SAS. Tous droits réservés. Mots clés : Faune de microvertébrés ; Paléoenvironnement ; El Kowm ; Hummal ; Unité G
1. Introduction Microvertebrates are rare in Syrian Pleistocene sites. That is unfortunate, since these kinds of fossils are particularly suitable for palaeoecological reconstruction and biostratigraphical inference. The small scale of typical terrestrial microvertebrate habitats allows inferences about palaeoenvironment very close to the site of deposition. The suitability of small mammals, in particular, for biostratigraphic age estimation results from their rather high evolutionary rates; for this reason, the identified evolutionary stages allow detailed age determination. In other parts of the Near East, particularly in Israel, the Pleistocene record of microvertebrates is much more dense (for a compilation see Tchernov, 1996 and references therein). On the other hand, there is only one sufficiently large sample is known from Syria: Late Pleistocene Douara Cave near Palmyra (Payne, 1983). We report on a concentration of microfaunal remains unearthed in a small excavation area (unit G, layer 17) of the archaeological site of Hummal. These remains occur together with numerous crushed and fragmented large bones (Wegmüller, 2011; Wegmüller et al., 2012). The site of Hummal is located in the El Kowm oasis in the desert steppe of Central Syria, a key area for the Palaeolithic in the Levant (Jagher and Le Tensorer, 2011; Le Tensorer et al., 2011; Jagher this volume). The sequence begins in the Lower Palaeolithic and concludes in the Upper
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Palaeolithic (Le Tensorer et al., 2011). In this paper, we present preliminary results of our investigations of the microvertebrates from Hummal and their palaeoecological and biostratigraphical implications. 2. Material and methods All microvertebrate remains mentioned here originate from an area of 4 m2 in layer 17 of unit G at Hummal (for details see Wegmüller, 2011; Wegmüller et al., 2012). This layer is situated c. 12.5 m below the surface. It is c. 10–15 cm thick and consists of dark grey to black compact clay. The precompactional thickness of the sediment layer was presumably greater. The layer is also rich in bone fragments of large mammals but poor in artefacts. It conformably overlies layer 18, a level very rich in artefacts. All layers superjacent to layer 17 are of Yabrudian, Hummalian or Mousterian age. Generally speaking, the lower part of the profile (layers 15–23) has yielded archaic artefacts, some of which are comparing to those from Ubeidiya in Israel (Wegmüller, this volume). More than 2000 microvertebrate remains was extracted by screen-washing of the sediments. Sixty percent of the remains clearly belong to small mammals, and ten percent to small reptiles, while 600 bones are of small vertebrates but cannot be identified more precisely. Many small mammal bone fragments are part of post-cranial elements. In general, these remains are unsuitable for identification to a fine taxonomic level. The incisors of rodents were merely identified as belonging to the order Rodentia. In the following descriptions, lower case denotes lower teeth and upper case denotes upper teeth. Dental morphological nomenclature of murines is after Vandebroek (1961–1962), that of arvicolines after van der Meulen (1973). 3. Taxonomy Class REPTILIA Laurenti, 1768 Order SQUAMATA Oppel, 1811 Suborder IGUANIA Cope, 1864 Family AGAMIDAE Fitzinger, 1826 Subfamily AGAMINAE Fitzinger, 1826 sensu Macey et al., 2000 AGAMINAE indet. Twenty-three cranial specimens, including maxillae (Fig. 1), ectopterygoids, quadrates, dentaries and coronoids, as well as a number of presacral and caudal vertebrae, are attributed to an agamine lizard. Caudal vertebrae lack autotomy planes, as in all members of Acrodonta (Etheridge, 1967), and the cheek tooth series includes two pleurodont tooth loci in the maxilla followed by a series of acrodont teeth, which is frequently interpreted as an apomorphy of Agamidae. The remains furthermore show characters of the African/West Asian clade Agaminae (sensu Macey et al., 2000), which have been interpreted as apomorphies (Maul et al., 2011): facial process (as indicated by its broken base) sharply folded medially along an oblique, posterodorsally trending axis to create a distinct anterodorsally directed surface behind the external nares; two pleurodont tooth loci in both maxilla and dentary; and acrodont teeth simple, triangular and unicuspid (also known in Hydrosaurus and Amphibolurinae). Presently available evidence on the identity of the Hummal agamid is therefore meagre but suggestive of Agaminae. Most members of that clade – especially those taxa found in the area
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today, including species of Trapelus (Wagner et al., 2011) – are considered ‘‘semiclimbers’’ and typical inhabitants of warm, temperate environments (Moody, 1980). Suborder GEKKOTA Cuvier, 1817 GEKKOTA indet. The most abundant squamate taxon at Hummal is a single species of gecko. Prefrontals and frontals are particularly common, but quadrates, compound bones, and dentigerous elements (Fig. 1) are also present. Derived gekkotan characters found in this species (see Estes et al., 1988; Gauthier et al., 2012) include: azygous frontal, subolfactory processes of frontal meet and fuse below olfactory tracts, and fused Meckelian groove. Comparative specimens of Near Eastern geckos are lamentably rare, precluding a more precise identification at this time, despite the propitiousness of the material. Suborder LACERTIFORMES Estes et al., 1988 Family LACERTIDAE Gray, 1825 LACERTIDAE indet. (at least two species) Forty-nine cranial bones, especially maxilla and dentary fragments (Fig. 1) but also frontals, prefrontals, an ectopterygoid (?), a jugal, postorbitofrontals, a quadrate, and a coronoid, are referred to Lacertidae. The frontals are ayzgous; the orbital margins show a narrow band where the supraocular osteoderms articulated. The postorbitofrontal is medially expanded, showing partial closure of the supratemporal fenestra. The exterior surfaces of the cranial bones (generally fused osteoderms) are sculptured. Jaws are the most abundant elements, and the posterior teeth are bicuspid. Tooth morphology and jaw size suggest that at least two species are represented. Suborder SERPENTES Linnaeus, 1758 Family BOIDAE Gray, 1825 Genus ERYX Daudin, 1802 Eryx sp. Three trunk vertebrae, a cloacal vertebra, and ten caudal vertebrae are attributed to a single species of the Old World sand boa clade Eryx. The trunk vertebrae are anteroposteriorly short, as in Boidae, and have low, restricted neural spines and somewhat depressed neural arches, as in Erycinae (Hoffstetter and Rage, 1972). The most diagnostic, and curiously also the most abundant, specimens are caudal vertebrae (Fig. 1) that evince the bifurcated neural spine and complicated accessory processes typical of erycine boids (Rage, 1974; Szyndlar and Rage, 2003; Smith, 2013). The more distal caudals are anteroposteriorly very short and tall, and the zygosphene-zygantral articulations are absent, as in Eryx and Bransapteryx (Szyndlar, 1994). As in most Eryx, a direct connection between the pre-zygapophyses and post-zygapophyses has been lost (Szyndlar, 1994; Szyndlar and Schleich, 1994). Further detailed studies of Eryx, incorportating the most recent results on phylogenetic structure within the taxon (Reynolds et al., 2014), will be necessary for an apomorphy-based identification of the Hummal species, but its attribution to Eryx appears secure. Only Eryx jaculus is known from Syria today. This species has a patchy distribution in North Africa, the Levant, and southeastern Europe, but does not presently occur in the hyperarid Syrian plains in which Hummal is situated. Szyndlar and Schleich (1994) previously described uppermost Miocene material from the south of Spain as Eryx cf. jaculus, postulating a formerly circum-Mediterranean distribution. Family COLUBRIDAE Oppel, 1811 sensu Lawson et al., 2005
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Subfamily NATRICINAE Bonaparte, 1838 NATRICINAE indet. Thirty elongate, delicate vertebrae appear to represent a single species of Natricinae. They all possess at least small hypapophyses, which on more posterior trunk vertebrae are short, pointed, and posteriorly directed. Class MAMMALIA Linnaeus, 1758 Some 700 post-cranial bone fragments are typical in size and shape of small mammals, but they have not been identified more precisely. Order SORICOMORPHA Gregory, 1910 Family SORICIDAE Fischer von Waldheim, 1814 Subfamily CROCIDURINAE Milne-Edwards, 1872 CROCIDURINAE indet. Despite its incompleteness, including loss of the processus articularis, a mandible with m1 and m2 and three isolated teeth clearly belong to a white-toothed shrew (subfamily Crocidurinae). Possible candidates are Suncus etruscus (Savi, 1822), Crocidura leucodon (Hermann, 1780), C. suaveolens (Pallas, 1811), C. katinka Bate, 1937, and C. caspica Thomas, 1907. However, for the moment, we cannot assign the fragments to a particular species or genus. Order CHIROPTERA Blumenbach, 1779 CHIROPTERA indet. Three molar fragments display characters diagnostic of bats: high cusps, crowns oblique in a longitudinal direction, and a strong cingulum. However, because of the fragmentary nature of the specimens, they cannot currently be referred to a particular taxon. Order LAGOMORPHA Brandt, 1855 Family LEPORIDAE Fischer von Waldheim, 1817 Genus LEPUS Linnaeus, 1758 Lepus sp. Several cheek teeth (two P2, two p3, two M) are characteristic of leporids. The diagnostic p3 (Fig. 2A) and P2 indicate the presence of the genus Lepus. The occlusal pattern with small buccal and lingual folds is similar to that of L. capensis Linnaeus, 1758, and the size of the teeth also
Fig. 1. Selected small reptile remains from Hummal. A. Partial left maxilla of Agaminae indet. (HERP3-c2). B. Partial left dentary of Gekkota indet. (HERP2-f4). C. Partial right dentary of Lacertidae indet. (HERP2-a10). D. Distal caudal vertebra of Eryx sp. (HERP1-a11). Sélection de restes de petits reptiles de Hummal. A. Portion de maxillaire gauche d’Agaminae indet. (HERP3-c2). B. Portion dentaire gauche de Gekkota indet. (HERP2-f4). C. Portion dentaire droite de Lacertidae indet. (HERP2-a10). D. Vertèbre caudale distale d’Eryx sp. (HERP1-a11).
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matches this species, which is smaller than L. europaeus Pallas, 1778. However, there is another small species, L. tolai Pallas, 1778, which is distributed in central Asia, westwards up to the Caspian Sea, and in north-eastern Asia. Due to the small number of specimens in Hummal and overlap in size, we tentatively refer the material to Lepus sp. Order RODENTIA Bowdich, 1821 RODENTIA indet. There are some 200 incisors of rodents, which have not been referred to the family or genus level. Family CRICETIDAE Fischer von Waldheim, 1817 Subfamily ARVICOLINAE Gray, 1821 ARVICOLINAE indet. Five molar fragments, consisting only of one or two prisms, can be identified only to the subfamily level. Genus ELLOBIUS Fischer von Waldheim, 1814 Ellobius sp. Forty molars (nine m1, nine m2, three m3, eleven M1, four M2, and four M3) of arvicoline rodents show typical traits of the genus Ellobius, or mole voles. The m1s possess three buccal and four lingual salient angles and have rather broad anterior caps (Fig. 2D). The dentine fields are broadly confluent and there is no crown cementum in the re-entrant angles. There are five extant species of Ellobius: E. talpinus (Pallas, 1770), E. tancrei Blasius, 1884, E. alaicus Vorontsov et al., 1969, E. fuscocapillus Blyth, 1843, and E. lutescens Thomas, 1897. Mole voles are not part of the recent fauna of Syria, but the geographically closest species E. lutescens occurs in northwestern Iran, eastern Turkey and southern Azerbaijan. It resembles our finds also morphologically more than to the other species. The m1 of E. lutescens possesses a broad anterior cap, whereas that of E. talpinus is narrow and that of E. fuscocapillus elongated. However, also E. pedorychus is rather similar to our finds. It was described by Bate (1937), from the Israeli Middle Pleistocene sites of Tabun C, D, and E. In the Middle Pleistocene, Ellobius is also represented in North Africa (Jaeger, 1988; Geraads, 2010; Stoetzel, 2013, etc.) by the following species: E. africanus Jaeger, 1988, E. atlanticus Jaeger, 1988, E. zimae Jaeger, 1988, and E. barbarus (Pomel, 1892). E. barbarus is more similar to E. talpinus, whereas E. africanus and E. atlanticus show resemblances to our finds in the anteroconid of m1. Thus, since the specimens from Hummal resemble various species – the extant E. lutescens, as well as the fossil E. pedorychus, E. africanus, and E. atlanticus – we tentatively refer them only to Ellobius sp. Genus MICROTUS Schrank 1798 Microtus sp. A single m1 fragment belongs to Microtus. It has no roots and there is crown cementum in the re-entrant angles. The dentine fields T4 and T5 are closed, the anterior cap has an ‘arvaline’ shape, i.e. it resembles that of Microtus arvalis. The enamel differentiation is Microtus-like. However, given the fragmentary preservation, the specimen is referred only to Microtus sp. Family MURIDAE, Illiger, 1811 Subfamily MURINAE Illiger, 1811 MURINAE indet. An M2 belongs to a member of the subfamily Murinae that is considerably larger than corresponding molars in other large-sized murines, like Rattus norvegicus (Berkenhout, 1769)
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and even larger than the corresponding molar of the Pest Rat Nesokia indica (Gray, 1830), which is more hypsodont and has fused tubercles (Niethammer, 1977). There are resemblances in size and shape to the M2 of Bandicota indica Gray, 1873. However, a reliable assignment requires further comparisons with larger samples. Genus MUS Linnaeus, 1758 Mus sp. Based on its size and the lack of tubercle t9 (Reichstein, 1978), one left M1 can be identified as belonging to Mus. Considering the distribution of extant species, it is most probably M. macedonicus Petrov & Ruzic, 1983. Since, however, it is only one specimen we refer it to as Mus sp. Subfamily GERBILLINAE Gray, 1825 Genus MERIONES Illiger, 1811 Meriones sp.and Genus GERBILLUS Desmarest 1804 Gerbillus sp. Some 250 molars and molar fragments can be assigned to the subfamily Gerbillinae. Among them, eight m1 and M1 belong to Gerbillus and 60 to Meriones. Species of the latter genus have larger and more hypsodont molars, the lateral incision of enamel (linea sinuosa) is more oblique distally, the outline of m1 is less asymmetric, and the anterior cap is broader than in molars of Gerbillus. In morphology and size, the finds are consistent with Meriones tristrami and Gerbillus dasyurus. The frequency distribution of m1 length, measured at the base of the crown indicates that the gerbilline finds belong to more than two species. Considering the distribution of extant species and size, possible candidates would be Meriones libycus, M. vinogradovi, and M. crassus on one hand and Gerbillus dasyurus, G. mesopotamiae, G. henleyi or G. cheesmani on the other. Morphologically they can hardly be distinguished. Therefore, we refer the Gerbillinae from Hummal tentatively only to the genus level (Fig. 2B and C).
Fig. 2. Selected small mammal remains from Hummal. A. Right p4 of Lepus sp. (MAMe3). B. Left m1 (invers) of Meriones sp. (GER1a1). C. Right m1 of Gerbillus sp. (GER1b6). D. Left m1 fragment (invers) of Ellobius sp. (MAMa11). Sélection de restes de petits mammifères de Hummal. A. p4 droite de Lepus sp. (MAMe3). B. m1 gauche (inversée) de Meriones sp. (GER1a1). C. m1 droite de Gerbillus sp. (GER1b6). D. Fragments de m1 gauche (inversé) de Ellobius sp. (MAMa11).
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4. Palaeoecology Based on the ecological requirements of the nearest living relatives, the recorded taxa suggest various open habitats, but also the presence of vegetation and wet areas, at least close to the site. Species of the genera Lepus, Ellobius, Meriones, Gerbillus, and Eryx live to various extents in grasslands, in steppes, semi-deserts, and deserts respectively (Harrison and Bates, 1991; Disi et al., 2001; Cos¸kun and Ulutürk, 2003; Krapp, 2003). If the referral of the large murid molar to Bandicota will hold true, this would indicate the occurrence of streams, other water bodies or swampy areas (http://www.iucnredlist.org/details/2541/0). Preliminary result using the coexistence approach (Mosbrugger and Utescher, 1997; Maul et al., in press) for a dataset of various recent close relatives of the recorded taxa, show higher precipitation values but similar temperatures in comparison with today. However, to obtain reliable results more detailed taxonomical identifications are required. Layer 17 as well as most of the other archaeological layers in Hummal were mainly formed during active episodes of the artesian spring (Le Tensorer et al., 2007) and therefore, the presence of water in direct vicinity of the site must be assumed. 5. Biostratigraphy In Plio- and Pleistocene Palaearctic sites, arvicolines generally allow best biostratigraphic estimation. At Hummal, these would be Ellobius und Microtus. Whereas Microtus today is nearly invariable element of faunas in the Near East, Ellobius is not a part of the extant fauna of the Levant. There are fossil records of Ellobius both from the Near East and northern Africa (for compilations see Tchernov, 1996; Stoetzel, 2013). All these finds are exclusively of Middle Pleistocene age. In Israel, the finds range from Late Acheulean (Oumm Qatafa) to Early Mousterian (Tabun D and Hayonim E). In North Africa, Ellobius is recorded only in the older part of the Middle Pleistocene, between c. 800 ka and 370 ka (Jaeger, 1988; Geraads et al., 2013). However, in the Levant, it is also present in slightly younger sites, as Tabun D, and C (Mousterian) (Bate, 1937), but not in the 780–700 ka old Gesher Benot Ya’aqov (Goren-Inbar et al., 2000). Thus, we could conclude for Hummal an age between early Middle Pleistocene and Early Mousterian, an age range between 780 and 165 ka. However, detailed studies of the Ellobius finds will provide further refinement of the biostratigraphic age of this assemblage. 6. Taphonomy According to the geological observation of the excavators, the layer experienced heavy sedimentary pressure. This is possibly one reason for the fragmentary nature of the remains. In addition, if the fragmentation is partly predepositional, it could suggest that the microvertebrates were accumulated as prey remains. The question of whether owls, diurnal raptors or mammalian carnivores are responsible for the accumulation is the subject of ongoing investigations. 7. Conclusion Preliminary investigation of the microvertebrates from layer 17 of unit G at Hummal provide taxonomic, biostratigraphic and palaeoecological information about the fauna and its palaeoenvironment. Our age estimate for Hummal layer 17 – from early Middle Pleistocene to Mousterian – is broadly consistent with the fact that layer 17 is subjacent to Yabrudian, Hummalian and
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Mousterian horizons. This estimate is based on the regional occurrence of Ellobius, which is not part of the extant fauna of Syria. However, the reason for the expansion and shrinking of the geographic distribution of Ellobius is thought to be quite complex. The geographic range of extant E. lutescens in eastern Turkey is restricted by topography and by the occurrence of Spalax rather than by temperature, rainfall or humidity (Cos¸kun and Ulutürk, 2003). It can be only speculated that ancient populations of Ellobius in Israel and Syria were relatively small and isolated. Probably, after their extirpation, recolonization was prevented because dispersal routes were blocked by competing Spalax and Nesokia. Current coexistence of Ellobius and other fossorial forms, such as Spalax, Nesokia and Myospalax is really marginal or even exclusive. The microfauna from Hummal is of great importance for the faunal history of this region since we have very few records from this area and time period. Unfortunately, excavations have been suspended due to armed conflict in the region. However, a detailed investigation of the microvertebrate material presented here, including a more extensive taxonomic analysis, is under way.
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