New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant

PALAEO-06502; No of Pages 12 Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

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New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant Adrian M. Lister a,⁎, Wendy Dirks b, Amnon Assaf c, Michael Chazan d, Paul Goldberg e,h, Yaakov H. Applbaum f, Nathalie Greenbaum f, Liora Kolska Horwitz g a

Earth Sciences Department, Natural History Museum, London SW7 5BD, UK Centre for Oral Health Research, School of Dental Sciences, Newcastle University, Newcastle upon Tyne NE2 4BW, UK Prehistoric Man Museum, Kibbutz Ma'ayan Baruch, Israel d Department of Anthropology, University of Toronto, 19 Russell St., Toronto, ONT M5S 2S2, Canada e Department of Archaeology, Boston University, 675 Commonwealth Ave., Boston, MA 02215, USA f Department of Radiology, Hadassah-Hebrew University Medical Center, Jerusalem 91230, Israel g National Natural History Collections, Faculty of Life Sciences, The Hebrew University, Jerusalem 91904, Israel h Eberhard Karls University Tübingen, The Role of Culture in Early Expansions of Humans (ROCEEH), Rümelinstr. 23, D-72070 Tübingen, Germany b c

a r t i c l e

i n f o

Article history: Received 11 February 2013 Received in revised form 3 May 2013 Accepted 5 May 2013 Available online xxxx Keywords: Elephas hysudricus Elephas maximus Asian elephant 'Ain Soda (Jordan) Ma'ayan Baruch (Israel) Middle Pleistocene

a b s t r a c t We describe new fossil remains of elephant (Elephas cf. hysudricus) from archaeological sites in the Levant: Ma'ayan Baruch (Israel) and 'Ain Soda (Jordan). Both sites date to the Middle Pleistocene based on stone artefacts typical of Levantine Late Acheulian assemblages. The elephant remains show ‘primitive’ dental features reminiscent of E. hysudricus from the Plio-Pleistocene of the Siwaliks (northern India), the species thought to be ancestral to Asian elephant E. maximus. Regionally, the new fossils are chronologically intermediate between an earlier (ca. 1 Ma) record of Elephas sp. from Evron Quarry (Israel), and Holocene remains of E. maximus from archaeological sites in NW Syria, Turkey, Iraq and Iran. It is unclear at present whether this represents continuity of occupation or, more plausibly, independent westward expansions. © 2013 Elsevier B.V. All rights reserved.

1. Introduction The ancestry of the living Asian elephant Elephas maximus L. is poorly understood. While the generic name Elephas was formerly applied to many different kinds of fossil elephant, only a few fossil species are now included within Elephas sensu stricto (Maglio, 1973). Of these, the Pleistocene species E. hysudricus of the Indian subcontinent and E. hysudrindicus of SE Asia are clearly, from their morphology, closest to the ancestry of the living species. However, the history of these species, their temporal and geographical extent, and the mode of transformation of one or both of them into the modern species, are poorly known. Elephas maximus is today restricted to the Indian subcontinent and SE Asia. In historical times, however, its range extended eastward to the Pacific coast of China, and westward to the Levant (Shoshani and Eisenberg, 1982; Sukumar, 2012). Until recently, earlier fossil evidence of Elephas s.s. in the western extremity of the distribution was restricted to an Early Pleistocene molar from Evron Quarry (Israel), referred to Elephas sp. by Tchernov et al. (1994). ⁎ Corresponding author. Tel.: +44 207 942 5398. E-mail address: [email protected] (A.M. Lister).

This article describes new fossil remains from the Levant that are referable to Elephas and are of Middle Pleistocene age: two elephant teeth found at Ma'ayan Baruch (Israel), and three partial molars from 'Ain Soda (Jordan). Other Elephas specimens from the region are revised, and the place of all of this material in the evolutionary history of the genus is assessed.

2. Materials 2.1. Ma'ayan Baruch The Late Acheulian locality of Ma'ayan Baruch is a large, open-air site at the northern end of the Hula Valley (Israel) (Fig. 1). The locality comprises numerous small find spots as well as three dense concentrations of lithic artefacts that were exposed by ploughing in the ‘Hamara’ fields of Kibbutz Ma'ayan Baruch. The artefacts lie within and on top of a terra rossa soil. Since the 1960s, some 8000 artefacts, predominantly handaxes, have been collected from an area of ca. 0.3 km2 (Stekelis and Gilead, 1966; Gilead, 1977; Ronen et al., 1980; Grosman et al., 2008). The ‘Hamara’ find locality has yielded a few bone (probably proboscidean) and tusk fragments (Stekelis and

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Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013

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Fig. 1. Map showing location of Pleistocene sites mentioned in the text (Letters) and Holocene Middle Eastern sites (Numbers) with the type of elephant remains found. Map

Site

Country

Material

1 2 3 4 5 6

Turkey Turkey Turkey Turkey Turkey Turkey

Tusk Tusk Bone Bone & tooth Tooth Bone

7 8 9 9

Ulu Burun (Kaş) shipwreck Acemhoyuk Sirkeli Tepe Gavur Lake Swamp Zincirli Chatal Hoyuk (Amuq Valley) Tel Tayinat Tel Atchana-Alalakh Minet el Beida Ras Shamra-Ugarit

Turkey Turkey Syria Syria

10 11 12 13 14 15 16 17 18 19 20 21 22

Kamid el-Loz Mishrife/Qatna Arslantepe Tel Sabi Abyad El Qitar Munbaqa Emar Tel Sheikh Hamad Chagar Bazar Nimrud Nuzi Babylon Haft Tepe

Lebanon Syria Turkey Syria Syria Syria Syria Syria Syria Iraq Iraq Iraq Iran

Tusk Bone & tusk Tooth Bone, tooth & tusk Bone Bone Bone Bone Bone & tooth Bone Bone & tooth Bone Tusk Bone & tusk Bone Bone Bone & tusk

Note: Tusk = whole tusks or sawn but otherwise unworked sections of the tusk.

Gilead, 1966:12), but it has not been possible to identify them to genus due to their extreme fragmentation. No excavations have been conducted at the Ma'ayan Baruch locality, but in 1974 and 1977 one of the authors (AA) recovered 28 artefacts from the walls of a naturally formed trench ca. 0.1 m deep, that had been created as a result of local high-velocity winter runoff. At the base of this trench, partly embedded in the north wall, a large elephant tooth was found lying on top of yellowish sediment. A small fragment of a second tooth was found in the eastern part of the same trench — the direction of the water flow (Ronen et al., 1980). The large tooth was removed by consolidating it in a block of sediment

cradled in a fibreglass jacket (see Suppl. S1). No other osteological remains were found in their vicinity. The soil surrounding the large elephant molar shows micromorphological features typical of an oxisol (Marcelino et al., 2010), and micromorphological analysis confirmed that the sediment in which the tooth was embedded had been waterlogged (see Suppl. S2). Researchers (e.g. Gilead, 1977; Bar-Yosef, 1994; Bar-Yosef and Belmaker, 2011) have grouped Ma'ayan Baruch with other Late Acheulian assemblages, such as Oumm Qatafa D1, that are dominated by cordiform bifaces with few ovates, pointed bifaces and cleavers. Analyses agree that all the Ma'ayan Baruch lithic assemblages display

Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013

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a high degree of standardization and show careful control of symmetry. As early as 1966, Stekelis and Gilead argued that they should be considered the product of a single industry. Ronen et al. (1980) reported some variation between artefacts recovered from the natural trench where the elephant teeth were found (and an assemblage recovered from a second trench), and artefacts recovered from the ‘Hamara’ field, with the trench lithics showing varying degrees of abrasion, alteration and rolling. They suggested that the artefacts recovered from the lowermost part of the trenches had been swept down the walls by the runoff water. A new study of the bifaces from the natural ‘elephant’ trench is presented in Suppl. S3, and concludes that there are no significant signs of long-distance transport on any artefacts in the sample currently available for analysis. Most artefacts are in very fresh condition and two bifaces show a clear difference in patina between the two faces suggesting that that were recovered from an undisturbed context. Although a number of handaxes in the trench were recovered lying in horizontal position, others were found in vertical orientation. Dating of the Ma'ayan Baruch find locality has relied on geological correlations with overlying or underlying basalts and travertines. The terra rossa soil in which the artefacts are found interdigitates with the Kefar Yuval Travertine, as shown by the travertine coating on many of the artefacts (Schwarcz et al., 1980). Heimann and Sass (1989) postulated that this travertine began to accumulate ca. 1 Ma and continued until ca. 25 ka. Based on pollen content, Horowitz (1979) estimated the age of the Kefar Yuval travertine as ~ 0.15 Ma, an age corroborated by radiometric measurements (Th-230/U-234) that gave an age of 189 ± 49 ka (Gur et al., 2002). Direct U-series dates on the Kefar Yuval travertine gave ages in excess of 350 ka, but this age was rejected as too old, the result of contamination (Schwarcz et al., 1980; Horowitz, 2001:561). As summarised by Horowitz (2001:559–560), the Kefar Yuval Travertine is overlain, unconformably, by the Ma'ayan Baruch basalt dated at 73 ± 14 ka (Seidner and Horowitz, 1974) and is underlain by the Hasbani and Dalwe basalts dated to ca. 1 Ma. Hence, the Ma'ayan Baruch Acheulian occurrence is older than 189 ± 49 ka BP but younger than 1 Ma. New chronological work is required to accurately date this travertine. Palaeomagnetic analysis was carried out on four soil samples taken from the block of matrix after removal of the elephant tooth at the Paleomagnetic Laboratory of the Institute of Earth Sciences, The Hebrew University of Jerusalem, by the late Prof. Hagai Ron. The original orientation of the block was reconstructed based on the field diaries of AA. Although this is certainly not an ideal way to sample for palaeomagnetism, all four samples were correctly oriented and gave similar Normal signals (Suppl. S4). Soil samples were also taken from the inner part of the block for OSL dating, but all proved to be unusable since the block had been exposed to light for several years since it was removed from the trench. Based on the geological context, the palaeomagnetic signal — with some reservation, and most clearly the character of the associated lithic artefacts, it may be concluded that the Ma'ayan Baruch elephant teeth are Late Acheulian in age. The time span covered by the Late Acheulian in the Levant is currently under debate. We support a time range of ca. 500–220 ka based on a compilation of currently available chronometric ages, as published in Porat et al. (2002). Other researchers however (e.g. Gopher et al., 2010; Bar-Yosef and Belmaker, 2011), constrain the Late Acheulian to ca. 600–400/350 ka, with the Acheulo– Yabrudian industries as a later and separate phase spanning the period 400/350–250/220 ka. 2.2. 'Ain Soda 'Ain Soda is an open-air site located in the wetlands of the Azraq Basin in eastern Jordan. Excavations, co-directed by Gary Rollefson, Philip Wilke and Leslie Quintero, were initiated in 1977 as an archaeological field school for students from San Juan College (Farmington,

3

New Mexico, USA). 'Ain Soda (Rollefson et al., 1997a, 1997b, 2006; Quintero et al., 2004), like the neighbouring sites of 'Ain el-Assad (Rollefson, 1983), C-Spring (Copeland and Hours, 1989) and those in the Al-Jafr basin (Rech et al., 2007), all reflect the distribution and movement of Middle Pleistocene hominins in the desert interior of Eastern Jordan. The 'Ain Soda site lies on the edge of a large pool, originally created by a spring fed by an underground aquifer, bringing water from as far away as Jebel Druze in southern Syria and Zarqa in western Jordan. In the Pleistocene, the site lay along the shore of what was once a large lake. Four trenches (1–4) were excavated along the northern, southern and western edges of the 'Ain Soda pool. Altogether some 60 m2 were sampled (Rollefson et al., 1997a, 1997b). The site yielded evidence of in situ Epipalaeolithic/Late Upper Palaeolithic, Early Mousterian and Late Acheulian occupations. In places the sediments were waterlogged due to the high water table (Quintero et al., 2004). Analysis of the artefacts from the Late Acheulian layers revealed an extremely high proportion (> 90%) of bifacial tranchet cleavers (Quintero et al., 2004:3, 2005; Wilke et al., 2005; Rollefson et al., 2006), similar to that found in Late Acheulian localities in the Al-Jafr basin (Rollefson et al., 2006). However, compared to sites in the Mediterranean region, such as the Late Acheulian of Tabun Cave Layer E, 'Ain Soda has a restricted range of artefacts, a high ratio (3:2) of flake tools to bifaces, and a slightly higher frequency of bifaces which are also larger and narrower (relative to length), with extensive use of Levallois technology. The 'Ain Soda locality has been identified by the excavators as a butchering site (Rollefson et al., 2006). The faunal preservation was good in both the Late/Final Acheulian and Mousterian deposits (Quintero et al., 2004), despite the fact that the site was an open-air locality and the silt dunes at the edges of the pool contain salts which are detrimental to bone preservation. Two trenches yielded Mousterian artefacts associated with aurochs (Bos primigenius) and equid remains. Two other trenches produced Late Acheulian artefacts together with faunal remains including those of rhinoceros (Stephanorhinus cf. hemitoechus), Equus hydruntinus, and an extinct elephant identified as Elephas cf. hysudricus (Dirks et al., 1998; Rollefson et al., 2006; the elephant remains were incorrectly listed in Rollefson et al., 1997a as Elephas planifrons). The three elephant teeth were recovered from the south trench (termed the “Elephant Trench”), and were found nearly at water level. Given the uniqueness of the elephant remains, the teeth were taken to the Mammoth Site of Hot Springs, South Dakota, USA and three sets of polyurethane casts made. Subsequently, the original teeth were lost in transit. Fortunately the casts survive and these were used in the current study in conjunction with notes and photographs made during direct observations of the teeth by WD. No radiometric dates are available for the site of 'Ain Soda, so dating of the elephant teeth is based on the characteristics of the associated lithic artefacts, which indicate a Late Acheulian age. As such, it falls within the same general age range as the Ma'ayan Baruch elephant remains. 3. Analytical methods 3.1. Measurements Measurement of elephant molars follows Maglio (1973), modified by Lister (2012: 207). Talons (‘x’) and platelets (‘p’) are not included in lamellar counts. Lamellae (‘plates’) are numbered ‘l1’, ‘l2’, etc., counting from the natural anterior end of the tooth, or ‘L1’, ‘L2’ etc., counting from the posterior end. Comparative data of modern Elephas maximus teeth is from original measurements of material at The Natural History Museum and other UK collections (see Acknowledgements). Roth and Shoshani (1988) also provide useful comparative data, but their lamellar counts have not been used as they included the vestigial structures here

Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013

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3.2. CT scanning Even after cleaning and consolidation, the occlusal surface of the large molar from Ma'ayan Baruch was poorly preserved. To facilitate examination and measurement, a CT scan was made of the tooth following Roth (1989). We used a Philips Brilliance 64ME (dual energy) CT scanner (Radiology Department, Hadassah Hospital, Jerusalem) to obtain contiguous CT slices in a coronal (transverse) planes passing from root to occlusal surface, and parasagittal (vertical) planes parallel to the vertical plane of the molar (for complete set of scans, see Suppl. S5). Scans were made at two settings: (i) 140 kV, 0.67 mm × 0.33 mm at 550 mas and (ii) 140 kV, 0.9 mm × 0.45 mm at 250 mas. Abbreviations: FAD: First Appearance Datum; LAD, Last Appearance Datum; NHM, Natural History Museum, London; UMZC, University Museum of Zoology, Cambridge; P, plate (lamellar) number; LF, lamellar frequency; H, crown height, HI, hypsodonty index; W, crown width; L, crown length; e, enamel thickness; M3, upper third molar; M3, lower third molar; CT, computed tomography. 4. Morphological and metric study of Maya'an Baruch specimens

A 240 Crown height (H), mm

termed talons and platelets, so the relationship of their scores to ours, which exclude these structures, is unclear.

220 200 180 160 140 120 100 65

B

The specimen MB1 (Fig. 2) is a very large elephantid molar from the left side, almost certainly an upper and very probably, from its size (Fig. 3), an M3. The preserved specimen represents the posterior 9 or 10 lamellae, the anterior part of the molar being partly lost. The

Fig. 2. The large molar (left M3) MB1, Elephas cf. hysudricus, from Ma'ayan Baruch, in medial (above) and occlusal (below) views. Anterior is to the left.

75

80

85

90

95

100

10 9 8 7 6 5 4 3 60

4.1. The larger specimen (catalogue MB1)

70

Crown width (W), mm

Lamellar Frequency (LF)

4

70

80

90

100

110

120

Crown width (W), mm Fig. 3. Dimensions of the upper molar teeth from Ma'ayan Baruch, 'Ain Soda, Siwalik Elephas hysudricus, and modern E. maximus. In both graphs, molar width (horizontal axis) is an index of the tooth size. A: crown height (H) against molar width. B: Lamellar Frequency (LF) against molar width. Blue diamonds: E. maximus M3; green triangles: E. maximus M2; red squares: 'Ain Soda M2 or M3; black circle: Ma'ayan Baruch M3; blue dashed box: M3 range of E. hysudricus from Maglio (1973); green dashed box: M2 range of E. hysudricus from Maglio (1973). In B, logarithmic regression lines have been fitted to the E. maximus M3 (blue) and M2 (green). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

interpretation of the specimen is challenging because of damage and likely distortion, but has been greatly aided by the CT scans (Fig. 4; Suppl. S5). The identification of the tooth as an upper is based, firstly, on the angle of the occlusal surface to the vertical plane of the lamellae. It is not perpendicular to the lamellae as in a lower, but forms an obtuse angle to them. Posterior to the occlusal region, the surface of the specimen tilts rootward for the last 3–4 lamellae (L3 to p in Fig. 2a), forming a ‘tented’ shape with the occlusal surface — again like an upper. This interpretation depends, however, on the exposed surface being natural and undistorted. Most parts of the exposed enamel appear naturally rounded through wear, suggesting that this is approximately the natural occlusal surface. The lamella labelled L1 in Fig. 2a shows an unworn and only slightly broken peak of a ‘digit’, positioned rootwards of the occlusal area, again consistent with the slope of the unworn posterior part of an upper. However, the two lamellae immediately behind the proposed occlusal surface (L2–L3) appear to have naturally worn enamel too; this is difficult to interpret since their apices are rootward of the main worn (occlusal) area. Conceivably they were part of the occlusal surface but have been moved rootward through crushing or slippage. In medial view, the lamellae converge from bottom to top of the crown, generally considered characteristic of lower rather than upper molars. The convergence is seen in the vertical CT sections (Fig. 4a,b) and is considered genuine, a separate phenomenon from the distorted orientation of some lamellae discussed below. However, the lamellae are quite straight and not S-shaped as in M3. Moreover a similar degree of overall convergence can be seen in some M3 — e.g. in E. maximus UMZC H.4692 (Fig. 5).

Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013

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Fig. 4. Selected CT scans of the Ma'ayan Baruch upper molar MB1. A–B, vertical scans; C–D, transverse scans. A: scan 95, close to midline of the tooth; B: scan 120, parasagittal scan between midline and medial edges of the tooth; C: scan 163, close to top of the crown; D: scan 133, about half-way down crown.

The posterior medio-lateral narrowing of the crown suggests we are dealing with the last molar (M3). The medial sides of the posterior lamellae are complete, and are positioned increasingly toward the tooth mid-line as we move posteriorly across the last three plates. The lateral sides are too broken to tell. In transverse scans (Fig. 4c, d), L3 is much narrower than L4–6; its medial and lateral ends both seem to be complete, and both are closer to the midline than in L4– 6. Moreover, the transverse CT scans suggest that the successive displacement of the medial edges is not due to slippage of the lamellae

in the medio-lateral plane, because homologous points (idiosyncrasies of enamel rings and wiggles) seem to line up antero-posteriorly between successive lamellae. The narrowing of the crown across its posterior three lamellae thus suggests an M3. However, anterior of L4, there seems to be no further increase in lamellar width (Table 1); this is unexpected as the widest part of an M3 is normally more anteriorly placed, with a longer zone of narrowing toward the posterior end. However, the morphology of the Ma'ayan Baruch tooth does not fit a typical M2 either. In an M2, the last true lamella and talon can be narrowed, but not as much as in the Maya'an Baruch tooth. Another diagnostic difference between M3 and M2 is the root, which tapers posteriorly in M3 but is widest at the very back of the tooth in M2. Unfortunately, the CT scans show that there is hardly any root left in the Maya'an Baruch tooth, so this feature cannot be determined. Overall, the significant narrowing across the preserved posterior three lamellae better fits an M3. The narrowness of the most posterior preserved lamella, seen on the CT scans (Fig. 4c,d), suggests that it is probably very close to the natural back end of the tooth. Additionally, the crown base of the posterior three lamellae rises in the direction of the crown apex. This is visible on the medial side of the tooth though not on the lateral side where the lamellae are very crushed. Such a trend in the crown base is common in elephant molars, and confirms that the last lamellae in the Maya'an Baruch specimen are close to, or at, the natural posterior end of the tooth. The vertical CT scan, Fig. 4b, suggests that the posterior ‘platelet’ (a reduced lamella analogous to the

Table 1 Measurements of Ma'ayan Baruch molar MB1. p = platelet, L1, L2, L3 etc. = position of lamella starting from posterior end. P, lamellar (plate) formula; L, crown length; LF, lamellar frequency across L3-L5; W, crown width (no cement); H, crown height.

Fig. 5. Left M3 of E. maximus, UMZC H.4692 in medial view, showing features similar to the large Ma'ayan Baruch molar; specifically: lamellae strongly convergent from base to apex, and posterior end lacking a long taper. Although unusual, these features in a demonstrable M3 corroborate the attribution of the Ma'ayan Baruch tooth to an upper third molar. Because of the distortion this specimen has not been plotted in Fig. 3b.

P L LF W H

p

L1

L2

L3

L4

L5

L6

?42 75

?65 108

?75 140

~90 150

~103 146

~103 Worn

~105 Worn

−10p >273 4.76

Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013

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talon of more anterior teeth; Lister and van Essen, 2003) is preserved behind L1, so the tooth is essentially complete posteriorly. Lamellae immediately in front of L1 (seen especially in the vertical CT sections, Fig. 4a) are quite high, suggesting that the posterior tapering of crown height was not pronounced in this specimen. Although posterior tapering is typical of third molars in elephantids, lesser tapering similar to the Maya'an Baruch example can be found in some M3 and even M3 of E. maximus (e.g. UMZC H.4692: Fig. 5), with very rapid lowering of crown height only in the posterior-most one or two lamellae. The preserved unworn ‘digit’ at the apex of the penultimate lamella (L1), mentioned above, if more or less in its correct place (as suggested by vertical CT scan, Fig. 4b), confirms a lowering of crown height, since its apex is rootward of those of more anterior lamellae. However, the restriction of tapering to the last few lamellae means this feature cannot be used to distinguish M2 from M3 in this specimen. The strongest evidence for post-mortem distortion of this tooth is the orientation of the lamellae seen in medial and lateral views. The wide spacing and backward sloping of the lamellae in the middle region of the crown gives an unnatural appearance. First, the sloping of the lamellae becomes less marked toward the back of the crown, rather than more marked as would be the case with the natural sloping of an M3. Second, the odd backward slope of the plates from L5 forward is not seen in the vertical CT scan (compare medial view Fig. 2a with midline scan Fig. 4a), suggesting that it is in large part due to crushing or movement of the lateral sides of the tooth. Overall, the form of the tooth suggests that it may have been partly liquified post-depositionally, and re-set in a slightly distorted shape. Spaces between the enamel ridges, that should have been filled with dentine or cement, are either hollow or appear to be filled with consolidated sediment. The tooth may have been partly dissolved – starting with the softer cement and dentine – so the remaining enamel plates were somewhat mobile – accounting for their odd orientation plus some possible up–down displacement. This interpretation is consistent with evidence for long-term waterlogging of the sediment (see above and Suppl. S2). Measurements of the molar are given in Table 1, but because of the damage and distortion of the crown, some discussion of the validity of the measurements is necessary.

4.1.1. Lamellar number and crown length Seven lamellae are clearly demarcated on the medial side of the molar, but the vertical CT scan shows additional lamellae in the damaged anterior portion, with a total of 8 lamellae clearly visible and the remains of probably two more, progressively dropped rootwards, in the crushed anterior end (Fig. 4b). These two lamellar remnants can just be seen on the medial side of the tooth itself. The total of 10 or so preserved lamellae is, however, unlikely to represent the original count, lamellae having been lost anteriorly through natural wear in life or breakage post-mortem. There are no preserved roots allowing us to assess position in relation to the original anterior end (cf. Sher and Garutt, 1987). However, in the vertical CT scans (Fig. 4b), even the anterior-most preserved plate has a significant height of crown remaining, suggesting that the loss of more anterior lamellae is due to breakage, not wear in life. The preserved length of the crown, 273 mm, is therefore also less than the original. Among Siwalik E. hysudricus and modern E. maximus, maximal recorded lengths for M3 are 300–340 mm, for a width of 90–100 mm (Maglio, 1973; Roth and Shoshani, 1988; Suppl. S6). With an estimated original width of 115–120 mm, the Maya'an Baruch molar, if of similar length/width proportion, could have been as much as 380 mm long when complete. Assuming a lamellar frequency of 4.76 as measured on the least-distorted region of the preserved molar (Table 1), original plate count can be estimated at ca. 18. The significance of this value is discussed below.

4.1.2. Crown width The maximum preserved width of the crown, 105 mm, was measured on L5 and L6. This is considered to be a true measurement because the medial ends of the lamellae appear undamaged, and while the lateral ends are damaged, the transverse CT scan shows three subequal enamel loops, with the centre of the middle loop at the preserved midline of the tooth, suggesting that little or nothing has been lost at the lateral side. No external cement is preserved and a molar of this size typically would have had 10–15 mm of cement (medial and lateral sides combined), indicating an original width of 115–120 mm. A value of 115 mm has been plotted on the graph (Fig. 3b). 4.1.3. Crown height The maximum preserved crown height, 150 mm measured just behind the occlusal surface on L3, is certainly not the original maximum of the tooth. Sher and Garutt (1987) showed that in elephantid M3s, there is a ‘zone of maximum (unworn) crown height’ in the central region of the tooth, with lower heights anterior and, especially, posterior to this, due to the posterior taper of the tooth. The maximum height of the Maya'an Baruch tooth would have been in the region that has been naturally worn, and is therefore not measurable. 4.1.4. Lamellar frequency Because of the severe damage to the anterior part of the preserved tooth, and the distortion to the middle portion discussed above, the region of lamellae L3–L5 is considered to give the most accurate estimate of original lamellar frequency. Lamellar frequency measured on all seven plates clearly visible on the medial side (i.e. including those considered to be distorted) gives a value of 4.54. The three lamellae L3–L5 give 4.76, and this is taken as the best estimate. 4.1.5. Enamel thickness This can be measured at a few points on the specimen, and also on the vertical and transverse CT scans, and gives values in the region of 2.8–3.0 mm. 4.1.6. Individual age Roth and Shoshani (1988: Fig. 7) presented a scheme of dental eruption versus age in known-age Asian elephants. In the Maya'an Baruch M3, all but the posterior four lamellae are in wear, and the M2 would have been naturally lost. Assuming an original lamellar count of 18 (see above), 14/18 or 78% of the lamellae were in wear, translating to an age of around 50 years in Roth and Shoshani's (1988) scheme. 4.2. The smaller specimen (MB2) This specimen is a crushed part of an elephantid molar crown (Fig. 6). It comprises six lamellae which are uncemented and no longer in their orderly alignment. The lamellae are apparently from the middle part of a molar, but their identity as upper or lower, left or right, is uncertain. Their apices are unaffected by wear in life, so the specimen represents a molar all or part of which was unerupted at death. The large size of the tooth – the widest lamella measures 96 mm without cement – indicates an upper or lower M2 or M3. Enamel thickness is ca. 3.5 mm. If the larger, worn molar from Maya'an Baruch is an M3 as suggested above, the smaller, unworn specimen cannot be from the same individual. If the larger specimen were an M2, the smaller tooth could be part of the unfinished M3 or M3 of the same individual. The lamellae appear to be either hollow or sediment-filled like the larger molar, suggesting a similar history of erosion or dissolution.

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7

Fig. 6. The small elephant molar from Ma'ayan Baruch, MB2. A: lateral view, B: apical view.

4.3. Generic identification

5.2. Upper molar M92450

Possible candidate taxa for the Maya'an Baruch teeth are Elephas, Palaeoloxodon, Mammuthus, and conceivably Loxodonta (although the latter genus has never been definitively identified outside Africa). Characters for the separation of these genera are given in Maglio (1973), Albayrak and Lister (2012) and elsewhere. The lack of a loxodont lamellar sinus rules out Loxodonta, while the hypsodont crowns rule out Mammuthus rumanus or M. meridionalis. All Mammuthus, moreover, have relatively unfolded enamel. Several features, however, point strongly to identification as Elephas. The pattern of wear of the lamellae, seen in the transverse CT scans of the large tooth (Fig. 4c,d), progresses from a row of small rings at the apex, to three subequal rings, which then fuse to form the lamella as the tooth wears, a typical configuration for Elephas. The transverse CT scans (Fig. 4c,d) also show the enamel band strongly folded into a series of tight loops along the lamella, again typical for Elephas. The smaller specimen also shows a row of small, equal digits at the apex, that would have worn into small rings, while the unfinished base of each lamella, seen in ‘root’ view (although there are no roots), shows strong, tight folding of the enamel band. Neither tooth shows any sign of the typical features of Palaeoloxodon, where the enamel folds are concentrated into a major fold in the midline of the tooth, flanked by smaller folds on either side of it, and additional minor (and not so strongly plicated) folds. The early wear pattern in Palaeoloxodon also typically shows a long central enamel loop flanked by two subcircular loops at the lateral and medial sides, not seen in the Maya'an Baruch molars. Palaeoloxodon is also characterised by relatively narrow crowns, unlike the markedly wide crowns of the Maya'an Baruch specimens.

This specimen represents the anterior part of a left upper molar, from its size M2 or M3. Its width, greater than the range of modern E. maximus M2 (Fig. 3), and the beginning of distinct curvature from front to back, makes M3 more likely, but as above, M2 cannot be excluded. Only the first three plates are worn, so the unworn plates behind give the true maximum crown height of the molar. Similarly the measured crown width is in the maximal region. If M3, the individual was in its early 30s at death; if M2, in its mid-teens.

5. Morphological and metric study of 'Ain Soda specimens The sample comprises three partial molars (Fig. 7), incomplete but well-preserved and undistorted. Measurements are given in Table 2.

5.3. Lower molar M92451 This is a small segment of a left lower molar, of uncertain position in the crown. From its width and evident curvature (concave laterally) it is probably M3, but M2 cannot be excluded. The occlusal surface of the piece is in very early wear. The two upper molars M92449 and M92450, from their respective crown widths and wear stages, cannot be from the same individual. It is not excluded, however, that the lower molar M92451 could be from the same individual as one of the uppers. 5.4. Generic identification Specimens M92449 and M92450 show subequal enamel rings in early wear, with no sign of the Palaeoloxodon features described above. In M92449 the second plate shows fusion of the rings into a longer lateral and shorter medial loop, commonly seen in early to mid wear of E. maximus (Albayrak and Lister, 2012). This specimen further shows the beginnings of strong enamel folding, and in all three specimens the ends of the enamel loops, seen in medial and lateral views of the teeth, are longitudinally grooved, an expression of strong enamel folding commonly seen in Elephas and Palaeoloxodon, but not Mammuthus or Loxodonta. The 'Ain Soda sample is therefore referred to Elephas.

5.1. Upper molar M92449

6. Morphometric comparison and specific identification of all material

This is the anterior part of a right upper molar, from its size M2 or M3. Its width, greater than the range of modern E. maximus M2 (Fig. 3), and its distinct curvature from front to back (concave medially, convex laterally), makes M3 more likely, but given the very large size of the Maya'an Baruch M3 (Fig. 3b), M2 cannot be excluded. Only the first three plates are worn, so the unworn plates behind give the true maximum crown height of the molar. Similarly the measured crown width is in the maximal region. If M3, the individual was in its early 30s at death; if M2, in its mid-teens.

The most important variables for specific identification within the Elephas hysudricus–E. maximus lineage are lamellar number and crown height, both of which increase between the two species. The roughly estimated original lamellar number of 18 for the Maya'an Baruch M3 compares to observed ranges of 12–17 in E. hysudricus (Maglio, 1973, n = 10) and 21–26 in E. maximus (n = 8; Suppl. 6), uppers and lowers pooled. If anything, the figure of 18 may be an overestimate because the estimated crown length on which it was based is greater than for any Elephas molar ever recorded. Overall,

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Fig. 7. Elephant molars, Elephas cf. hysudricus, from 'Ain Soda. A: right upper molar M92449 in occlusal and lateral views; B: left upper molar M92450 in occlusal and lateral views; C: left lower molar M92451 in occlusal and medial views.

therefore, lamellar number appears more consonant with E. hysudricus than E. maximus. All of the 'Ain Soda teeth are too incomplete to allow estimation of original lamellar number. In this situation, lamellar frequency (LF), the number of lamellae in a 10 cm length of tooth, can be used as a proxy, provided the size-related nature of lamellar frequency is taken into account (Lister and Joysey, 1992). LF is inversely related to molar size, here represented by crown width (W). In Fig. 3b, LF is therefore regressed against width (W), with a logarithmic fit applied to the modern comparative sample, extrapolated to encompass the large widths of the fossil specimens. Modern M3 and M2 are both plotted, because of the uncertain positional identity of the 'Ain Soda specimens. The graph shows both the two 'Ain Soda M2/3 plotting clearly below the trends of the modern sample, especially that of M3, indicating lower LF even taking their large size into account, and implying an originally lower lamellar number (P) than in modern E. maximus. The Maya'an Baruch M3 similarly falls below the extrapolated modern LF/W trend, corresponding to its lower estimated lamellar number than the living species. Hypsodonty cannot be estimated for the Maya'an Baruch tooth, but in the two 'Ain Soda M3s it clearly falls below the modern sample in a plot of crown height (H) versus width (W) (Fig. 3a). This corresponds to hypsodonty indices (H/W) of 1.35 and 1.44 (Table 2), compared to a range of 1.98–2.71 in E. maximus (n = 12; Suppl. S6).

Table 2 Measurements of 'Ain Soda molars. Abbreviations as in Table 1, plus: HI = hypsodonty index. Width measurements (W) exclude cement, but are plotted (Fig. 3) with an allowance of 5 mm for missing cement. LF of the lower molar measured at base (medial and lateral averaged) only (see Lister, 2012). Specimen

Position

P

L

LF

W

H

HI

M92449 M92450 M92451

RM2/3 LM2/3 Lm2/3

x8– x6– −4–

>164 >120 >95

5.82 5.35 4.73

89 92 ≥87

135 131 ≥110

1.44 1.35 –

The LF and H values of the 'Ain Soda specimens, lower than modern E. maximus, fall, however, within the range of E. hysudricus M3 from the Siwaliks (Fig. 3b; data from Maglio, 1973). The LF of the Maya'an Baruch M3 also falls within this range, with a value somewhat below those of 'Ain Soda which can be accounted for by its very large size and the inverse relation of LF to tooth size (Lister and Joysey, 1992). All of the studied molars, therefore, are at an evolutionary level comparable to Siwalik Plio-Pleistocene E. hysudricus. Because of the limited nature of the material, and the lack of characters outside the dentition, they are referred to Elephas cf. hysudricus. 7. Discussion and conclusions 7.1. Significance of the Levantine Pleistocene fossils in the evolution of the Asian elephant The most numerous fossil remains from the lineage of the Asian elephant have come from the Indian subcontinent, especially the Siwalik Group of India and Pakistan. Of the two Elephas species known from the Siwaliks, E. planifrons Falconer & Cautley, 1845 and E. hysudricus Falconer & Cautley, 1845, the latter is clearly, from its cranial and dental morphology, closest to the ancestry of the living species E. maximus L. (Maglio, 1973). A related fossil species, E. hysudrindicus Dubois, 1908, is known from insular SE Asia. Finally, the more distantly related Palaeoloxodon namadicus (Falconer & Cautley, 1846), originally and still sometimes named Elephas namadicus, also occurs in the Pleistocene of the Indian subcontinent. In order to assess the position and significance of the new Levantine finds in the context of Asian elephant evolution, existing evidence on the LAD of E. hysudricus and FAD of E. maximus will be reviewed (see also Vidya et al., 2009: supplementary information 10). This evidence is complicated by both stratigraphic and dating uncertainties, and issues of the correct identification of remains. In the Siwalik sequence, E. hysudricus is characteristic of the Pinjor Formation; the appearance of the species, and the base of the Formation, are currently placed at 2.6–2.7 Ma (Nanda, 2002). The upper limit of the Pinjor is time-transgressive but its youngest date, and

Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013

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that of E. hysudricus in the Siwaliks, is placed at 0.6 Ma (Nanda, 2002, 2008). Based on available published information, Chauhan (2008) and Nanda (2008) note the persistence of E. hysudricus into the postSiwalik (Middle to Late Pleistocene) Narmada (= Narbada) and Godavari beds of Peninsular India, although Nanda (2008, p. 9) cautions that “the post-Siwalik faunal lists provided by various workers are not always supported by descriptions, line diagrams or photographs”. According to Chauhan (2008), the Lower Narmada group (Middle Pleistocene, i.e. ca. 780–125 ka) yielded both E. hysudricus and E. namadicus. Cameron et al. (2004) indicate a date of >236 ka for Narmada fauna associated with Late Acheulian industry. Chauhan (2008, p. 27) further indicates that the Upper Narmada Group (early Late Pleistocene) ‘is thought to yield’ both E. hysudricus and P. namadicus, while Nanda (2008) lists E. hysudricus in the Late Pleistocene (ca. 125–12 ka) of Peninsular India and the Indo-Gangetic region. Other authors, however, such as Deraniyagala (1955), Khatry (1966), Maglio (1973) and Badam (1979) list only E. namadicus for Narmada and Godavari, implying that E. hysudricus was extinct, at least regionally, by the end of Pinjor times (i.e. 0.6 Ma). The earliest date for E. maximus is also problematic. Badam (1988, cited by Chauhan, 2008) is of the opinion that the species is not found in the older part of the Narmada sediments (Middle Pleistocene), but is present only in the Late Pleistocene of the Narmada and other Deccan fluvial systems. Nanda (2008), similarly, lists the species as present in the Late Pleistocene of Peninsular India and the Indo-Gangetic region. On the available evidence it must be admitted that the date and mode of the transition from E. hysudricus to E. maximus within the Middle to Late Pleistocene is uncertain (Dennell, 2004; Vidya et al., 2009: supplementary information 10; Sukumar, 2012). If the identification of both E. hysudricus and E. maximus in the Late Pleistocene is correct, this could reflect either (i) strict chronological co-occurrence, implying earlier speciation of E. maximus elsewhere, followed by dispersal into the Indian range of E. hysudricus (cf. Eurasian Mammuthus: Lister et al., 2005); or (ii) that the remains are of different ages within the Late Pleistocene and that E. maximus chronologically replaced E. hysudricus. However, the persistence of E. hysudricus beyond the Middle Pleistocene still remains to be rigorously demonstrated. The likely age of Maya'an Baruch and 'Ain Soda (in the range ca. 500–220 ka), and the correspondence of their dental morphology to E. hysudricus, make them potentially the youngest dated remains attributable either to that species, or at least to a transitional form not yet at the level of E. maximus. If E. maximus evolved anagenetically from E. hysudricus, this suggests 500 ka as a maximal age for the transition. However, it cannot be excluded that E. maximus arose earlier in another part of the E. hysudricus range (presumably further east), so that the Levantine population evidenced at Maya'an Baruch and 'Ain Soda represents a relict, more primitive population. At present, since we lack dated early remains of E. maximus from the Indian subcontinent or SE Asia, it is impossible to choose between these options. Vidya et al. (2009) showed that the two major mitochondrial DNA clades within modern E. maximus originated 1.6–2.1 Myr ago, the mitochondrial split therefore probably occurring within E. hysudricus and plausibly in isolated populations during a period of refugial contraction in the regions of Myanmar and India/Sri Lanka. They further suggest that the current complex pattern of distribution of the mtDNA clades in E. maximus resulted from a series of contractions and expansions during the climatic oscillations of the Quaternary, implying a complex distributional history for the living species. An alternative, intriguing possibility is that the two palaeo-species recognised on morphology, E. hysudricus from mainland south and southeast Asia, and E. hysudrindicus from insular southeast Asia, could be the origin of the two clades in modern E. maximus, so that the latter represents a ‘hybrid’ of the two forms. Vidya et al. (2009) consider this less likely because of the Late Pleistocene age assigned to fossils of

9

E. hysudrindicus. However, as summarised here, the age of this material, and the date of origin of the species, are poorly-constrained, so its contribution to the modern species remains a possibility. The earliest known Elephas in the Levant comprises an upper molar from Evron Quarry (Fig. 8) reported by Lister in Tchernov et al. (1994) and dated to between ca. 1.0 and 0.78 Ma (Ron et al., 2003). Its size (crown width 73 mm including cement) is too large for dP4 and makes identification as M1 highly probable. In this case its morphology is closer to E. hysudricus than E. maximus (Table 3), with some caution because of damage to the specimen (Tchernov et al., 1994). The plate count of 10 assumes the preserved anteriormost root is the true anterior root. Crown height cannot be measured at its highest point (at the posterior end in an M1), but the value of 87 mm on plate 6 suggests that a true maximum in the range of 108–111 (E. hysudricus) is much more likely than 127–142 (E. maximus) (see Table 3). The LF value for the Evron tooth is valid and unaffected by the breakage, and corroborates a morphology more primitive than the living species. Other records of ‘Elephas’ in the Levant are not supported by detailed morphological study. Pliocene remains from Bethlehem formerly referred to E. planifrons (Hooijer, 1958) are now considered probably to pertain to Mammuthus (Markov, 2012). Bate (1937:222) listed a tusk fragment from Tabun Layer E (Israel) as ‘Elephas sp.’, but the identity of this and other tusk fragments from Tabun in the NHM collection cannot be determined beyond Elephantidae indet. A rolled, partly mineralized posterior crown fragment of an elephantid upper molar (cf. right M1 or M2) from the Late Acheulian locality of Oumm Zinat (Israel), listed as Elephas by Horwitz and Tchernov

Fig. 8. Elephant molar, cf. Elephas hysudricus, from Evron Quarry. A: occlusal view, B: lateral view.

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Table 3 Comparison of M1 from Evron Quarry with E. hysudricus and E. maximus. Evron data from Tchernov et al. (1994); Siwalik E. hysudricus from Maglio (1973: Table 23); modern and subfossil E. maximus (Supplementary S6). Numbers in brackets are sample sizes. Taxon

P (uppers & lowers pooled)

LF (uppers)

H (uppers)

Evron E. hysudricus M1 E. maximus M1

10 9–10 (5) 12–15 (7)

7.5 5.5–8.2 (2) 7.8–10.5 (6)

>87 108–111 (2) 127–143 (4)

(1989), is of uncertain identity but shows features suggestive of Palaeoloxodon sp. 7.2. Ecology of Levantine Pleistocene proboscideans The ecology of Elephas hysudricus is essentially unknown, although its dental morphology – with moderate development of hypsodonty and lamellar number – suggests a mixed-feeder taking both browse and graze. In the southern Levant, there is overlap in the chronological range of several proboscidean taxa in the late Early to Middle Pleistocene. E. cf. hysudricus at Evron Quarry (1.0–0.78 Ma) co-occurs with Stegodon (Tchernov et al., 1994), while Gesher Benot Ya'akov (Israel), dated to slightly younger than 0.78 ka (Goren-Inbar et al., 2000), has yielded both Stegodon and the earliest occurrence in the region of the straight-tusked elephant Palaeoloxodon (Tchernov and Shoshani, 1996). Eurasian Palaeoloxodon is a migrant from Africa, a derivative of P. recki (formerly Elephas recki). The Gesher Benot Ya'akov elephantid skull was identified as the European species P. antiquus (Goren-Inbar et al., 1994; Shoshani et al., 2001), but has been considered a possible P. recki by Saegusa and Gilbert (2008). Possible contemporaneity of Mammuthus during this interval also cannot be excluded (M. meridionalis at 'Ubeidiyeh, Israel; M. trogontherii at Latamne, Syria: Lister, 2004). Lister (2004) discussed the ecology of Palaeoloxodon in Eurasia and its niche separation from the Mammuthus lineage, the latter moving from browser/mixed feeder M. meridionalis toward a more grazing adaptation (M. trogontherii) after the entry of the browsing/ mixed-feeding Palaeoloxodon. However, on current evidence it is difficult to ascertain if any of the three genera were precisely contemporaneous (and hence competed) at any time in the Levant. The next known occurrence of Palaeoloxodon in the region (as P. antiquus) is in the Late Acheulian beginning ca. 500 ka. Records include Holon (Davies and Lister, 2007), Revadim (Rabinovich et al., 2012), Oumm Zinat (Horwitz and Tchernov, 1989; see above), and most recently the Zuq Fawqani find locality near to Ma'ayan Baruch (see Suppl. S7). Although the Late Acheulian Palaeoloxodon remains span the same general time period as the Elephas finds from Ma'ayan Baruch and 'Ain Soda, their remains are not found in the same sites, so it is unclear if they coexisted and presumably exploited different habitats or resources (niche separation). Alternatively, they may have been separated chronologically, a possibility given the lengthy time span of the Late Acheulian and the lack of refinement in dating the localities. 7.3. Elephas in the Holocene of the Levant It is clear from archaeological and documentary evidence that in relatively recent times the distribution of Elephas maximus extended much further west than it does today. However, the western limit of the distribution has been unclear, with authors varyingly placing it in Iraq, NE Syria or SE Turkey (e.g. Hofman, 1974; Shoshani and Eisenberg, 1982; Sukumar, 1989, 2012; Santiapilli and Jackson, 1990). Faunal remains of elephants from Holocene archaeological sites in Southwest Asia have been summarised by several researchers (Miller, 1986; Caubet and Poplin, 1987, 2010; von den Driesch, 1996; Albayrak and Lister, 2012), and by Becker (2005, 2008) who has also

reviewed the ancient written and iconographic records dealing with elephants. Fig. 1 presents an updated map of these finds. Some researchers have supported the idea that a living E. maximus population inhabited the Euphrates–Tigris River region in the late Holocene (e.g. Miller, 1986; Becker, 2005, 2008). The disappearance of this population has been linked to human predation and hunting, changing climate (aridification) and/or diminishing natural habitats, especially deforestation (Miller, 1986; Sukumar, 2012). The last appearance has been dated to the early 1st millennium BC, this being the latest written reference to the hunting of live elephants in north Syria by the Assyrian king Shalmaneser III (858–824 BC) (Winter, 1973: 265; Miller, 1986; Moorey, 1994; Becker, 2005, 2008). However, other researchers view the same finds as remains of live animals and/or raw material that originated in the Indian subcontinent and were traded, sent as tribute, or dispatched for help in military campaigns to areas in the west (Deraniyagala, 1955; Winter, 1973; Colon, 1977; Vila, 2010). Reconstructing the Holocene history of E. maximus in Southwest Asia is further complicated by imprecise species identifications, as in the case of material from the site of Kamid el Loz, Lebanon (Bökönyi, 1985), or problematic contexts, for example the contested Early Bronze Age date for elephant remains from Ras Shamra (Ugarit) in Syria (Hooijer, 1978; Moorey, 1994:118; Caubet and Poplin, 1987, 2010). The Elephas cf. hysudricus remains from the Middle Pleistocene sites of Ma'ayan Baruch and 'Ain Soda, spanning the period 500–220 ka, are chronologically intermediate between cf. Elephas sp. from Evron Quarry, dated to between ca. 1.0 Ma and 0.78 Ma, and the mid-Holocene E. maximus of the Near East. However, available data are too scanty to assess whether this represents continuity of occupation, independent westward expansions from further east, or importation of some or all of the Holocene material (see above). The Pleistocene records do, however, provide a precedent for the natural expansion of Elephas as far as the Near East. While not proving the existence of an indigenous Holocene population, it makes it at least ecologically plausible. Supplementary data to this article can be found online at http:// dx.doi.org/10.1016/j.palaeo.2013.05.013. Acknowledgements Special thanks to Ms. Gali Beiner for her invaluable and painstaking restoration work on the elephant teeth from the Ma'ayan Baruch trench, and to Mr. Vladimir Nakhlin for the photography of the Ma'ayan Baruch and Evron Quarry teeth and lithics. We are grateful to Prof. G.O. Rollefson and Dr. Leslie Quintero for the permission to study and publish the 'Ain Soda remains and to Gary Sawyer (American Museum of Natural History) and Malon Anderson (Mammoth Site of Hot Springs, South Dakota) for the assistance with preparation of the casts. We would like to acknowledge the contribution of the late Prof. Hagai Ron, who undertook the palaeomagnetic analysis of the Ma'ayan Baruch sediment block, and Dr. Naomi Porat (Geological Survey of Israel) for trying to date the Ma'ayan Baruch sediment block using OSL. Thanks also to Yael Ebert for assistance with the palaeomagnetic results. For access to modern Asian elephant molars we thank Roberto Portela-Miguez (Natural History Museum, London), Mark Carnall (Grant Museum, UCL, London), Matthew Lowe and Ann Charlton (University Museum of Zoology, Cambridge), Milly Farrell (Hunterian Museum, Royal College of Surgeons, London) and Natasja den Ouden (Naturalis, Leiden). Work on the Ma'ayan Baruch specimens was funded by grants from the Canadian Social Sciences and Humanities Research Council to MC. References Albayrak, E., Lister, A.M., 2012. Dental remains of fossil elephants from Turkey. Quaternary International 276–277, 198–211. Badam, G.L., 1979. Pleistocene Fauna of India, with Special Reference to the Siwaliks. Deccan College Postgraduate and Research Institute, Pune, India.

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Please cite this article as: Lister, A.M., et al., New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant..., Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.05.013