Mammalia, Cervidae) During the Middle Holocene in the Cave of Bizmoune (Morocco, Essaouira Region

Quaternary International xxx (2015) 1e14

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The last occurrence of Megaceroides algericus Lyddekker, 1890 (Mammalia, Cervidae) during the middle Holocene in the cave of Bizmoune (Morocco, Essaouira region)

* Philippe Fernandez a, , Abdeljalil Bouzouggar b, c, Jacques Collina-Girard a, Mathieu Coulon d a Aix Marseille Universite, CNRS, MCC, LAMPEA UMR 7269, 13094, Aix-en-Provence, France b Institut National des Sciences de l'Archeologie et du Patrimoine, Rabat, Morocco c Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, D-04103 Leipzig, Germany d Aix Marseille Universite, CNRS, LAMES UMR 7305, 13094, Aix-en-Provence, France article info abstract

Article history: During the course of archaeological test excavations carried out in 2007 in the cave of Bizmoune Available online xxx (Essaouira region, Morocco), seven archaeological layers yielding Pleistocene and Holocene artefacts and faunal remains were identified. In the layers C4, C3 and C2, respectively from the oldest to the most Keywords: recent, terrestrial Helicidae mollusk shells (Helix aspersa) were dated by 14C. These layers also contained Giant deer many fragments of eggshell, belonging to Struthio cf. camelus, associated with mammal remains such as Extinction Oryctolagus/Lepus, Gazella sp., Sus scrofa, Ammotragus lervia, Alcelaphus buselaphus, Equus sp., Pha- Holocene cochoerus aethiopicus and an undetermined Caprini. Among these remains, an incomplete mandible of North Africa Speciation Megaceroides algericus Lydekker, 1890 with M1 and M2 was found in layer C3. The 6641 to 6009 cal BP Palaeoecology time range attributed to this layer has provided the most recent date known so far for M. algericus. In this study, we review and contextualize the findings of this particular species both in time and space and discuss its systematic position. We describe the morphology of the typical pachyostosic mandibular bone with the teeth and compare the dimensions with existing data. The assumption of the combined development, on the one hand, of the pachyostosic phenomenon and on the other hand, of the body weight fluctuations and growth of antlers for cervids strongly affected by seasonality is not supported. In order to understand the origin and the extinction of M. algericus, we examined the AMS radiocarbon dates available in the literature and calibrated them with RenDateModel software. Comparisons are then made with sea surface temperatures (e.g. GISP2 d18O), eustasy and related environmental changes throughout the time span of this species. Based on these data a possible migration route by the Strait of Gibraltar connected with with eustatic rises in sea-level rises are discussed. The speciation-extinction processes for M. algericus and their correlations with climatic shifts on a long time-scale in North Af- rica (e.g. Heinrich events, 8200 cal BP event) are also considered. Finally, this new discovery in Bizmoune cave clearly shows that M. algericus lasted until the very end of the Epipaleolithic, around 6000 cal BP (middle Holocene), whereas this species was formerly not believed to have survived until the early Epipaleolithic (around 8000 cal BP). © 2015 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction Bizmoune is situated at an altitude of 260 m. The cavern is formed within Upper Cretaceous limestone with an entrance that faces The cave of Bizmoune (834032.9900W/3140010.0200N; WGS 84) southwest. It is roomy and well-lit, measuring some 15 m deep by is located about 20 km southeast of Essaouira (Mogador) and was 10 wide. Test excavations in the cave in 2007 revealed the presence discovered in 2005 by A. Bouzouggar (INSAP, Rabat) (Fig. 1). of North African Middle Stone Age (MSA), Late Stone Age (LSA) and Neolithic layers (Bouzouggar et al., 2010). In the present study, we ﬁ * Corresponding author. focus on the last giant deer species M. algericus identi ed from E-mail address: [email protected] (P. Fernandez). a layer dating to the middle Holocene. The evolution and the http://dx.doi.org/10.1016/j.quaint.2015.03.034 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved.

Fig. 1. Location of sites with M. algericus in the literature. Data from Table 1. paleoecological context of this species, which is exclusively 1.1. Megaceroides algericus in the Maghreb distributed in northwestern Africa during the Upper Pleistocene, are discussed here. The remains of M. algericus are rare and only M. algericus was identiﬁed for the ﬁrst time by Lydekker (1890) documented in 26 localities of Morocco and Algeria (Table 1, Fig. 1). as Cervus algericus in the late Pleistocene deposits at Hammam

Table 1 Inventory and dating of sites with Megaceroides algericus and their main bibliographic references (see Fig. 1 for location on map). LP ¼ Late Pleistocene; H¼Holocene.

Sites Former Countries Datation Main bibliographic references names

Aïn Tit Mellil Maroc LP Arambourg (1938a, 1938b); Hadjouis (1990); Camps (1993); Abbazzi (2004); Merzoug (2012) Ain-Taya Algerie LP Hadjouis (1990); Merzoug (2012) Ali Bacha Algerie LP Debruge (1907); Joleaud (1916); Arambourg et al. (1934); Vaufrey (1955); Camps (1993); Merzoug (2012) Anglade Guyotville Algerie LP Arambourg (1932, 1935); Arambourg et al. (1934); Hadjouis (1990); Abbazzi (2004); Croitor (2006); Merzoug (2012) Beni Segoual (n ¼ 4) Algerie LP Arambourg et al. (1934); Hadjouis (1990); Merzoug (2012)

Table 1 (continued )

Sites Former Countries Datation Main bibliographic references names

Berrouaghia Algerie LP Pomel (1892, 1893); Joleaud (1912, 1916); Hadjouis (1990); Merzoug (2012) Bizmoune Maroc LP/H Bouzouggar et al. (2010) Bouknadel Maroc LP Ennouchi (1953); Michel (1990, 1992); Hadjouis (1990); Merzoug (2012) Cap Carbon Bougie Algerie LP Pomel (1892, 1893); Joleaud (1912, 1916); Arambourg et al. (1934); Hadjouis (1990); Camps (1993); Merzoug (2012) Filﬁla Algerie LP Vaufrey (1955); Hadjouis (1990); Abbazzi (2004); Merzoug (2012) Grotte Rolland Algerie LP Marchand (1932); Camps (1974); Merzoug (2012) Hamman Meskhoutine Algerie LP Lydekker (1890); Joleaud (1914, 1916); Arambourg (1938a); Arambourg et al. (1934); Hadjouis (1990); Merzoug (2012) Kifan bel Ghomari Taza Maroc LP Doumergue (1917, 1936); Arambourg (1938a); Arambourg et al. (1934); Hadjouis (1990); Camps (1993); Merzoug (2012) La Mouillah Algerie LP Arambourg et al. (1934); Vaufrey (1955); Merzoug (2012) Les Bains romains (n ¼ 2) Algerie LP Ficheur and Brives (1900); Joleaud (1912, 1916); Arambourg et al. (1934); Hadjouis (1990); Camps (1993); Merzoug (2012) Les Phacocheres Les Allobroges Algerie LP Hadjouis (1990); Camps (1993); Merzoug (2012) Mugharet el Aliya Grotte d'Hercule Maroc LP Arambourg (1967); Camps (1993); Merzoug (2012) Oued Kerma Algerie LP Vaufrey (1955); Merzoug (2012) Pics des singes Algerie LP Joleaud (1912); Arambourg et al. (1934); Camps (1993); Merzoug (2012) Pointe Pescade Algerie LP Ficheur and Brives (1900); Arambourg (1931, 1932); Arambourg et al. (1934); Vaufrey (1955); Hadjouis (1990); Camps (1993); Merzoug (2012) Puits des Chaachas Algerie LP Vaufrey (1955); Camps (1993); Merzoug (2012) Sidi Saïd Algerie LP Betrouni (1997); Chaïd-Saoudi (2013) Sintes Guyotville Algerie LP Arambourg (1932, 1935); Arambourg et al. (1934); Vaufrey (1955); Hadjouis (1990); Camps (1993); Abbazzi (2004); Croitor (2006) Taforalt Grotte des Pigeons Maroc LP Roche (1963); Camps (1993); Wrinn and Rink (2003); Merzoug (2012) Tamar Hat Bougie Algerie LP/H Arambourg et al. (1934); Saxon et al. (1974); Hadjouis (1990); Camps (1993); Merzoug (2005, 2012) Taza I La Madeleine Algerie LP Arambourg et al. (1934); Delibrias et al. (1974); Hadjouis (1990); Camps (1993); Meier and Sahnouni (1995); Medig et al. (1996); Meier et al. (2003); Abbazzi (2004); Merzoug (2005, 2012); Croitor (2006)

Meskhoutine, near Guelma in Algeria. Lydekker's determination, In total, no more than six sites with remains of this species have based on a left maxilla, with P3 to M3, included features such as the been identified in Morocco: typical brachydonty of the squared crowns, the very marked in- - Aïn Tit Mellil (Arambourg, 1938a, 1938b; Abbazzi, 2004); ternal cingulum and interlobar columns. Unaware of the earlier - Kifan bel Ghomari or Taza (Doumergue, 1917, 1936; Arambourg work of Lydekker (as suggested by Joleaud, 1914), Pomel (1892) et al., 1934; Arambourg, 1938a); later named this species Cervus pachygenys after studying the ma- - Bouknadel (Ennouchi, 1953; Michel, 1990, 1992); terial from a cave near the city of Bejaïa (formerly Bougie) and from - Taforalt, formerly known as the Grotte des Pigeons (Roche,1963; the remains of a railway bed in Berrouaghia in Algeria. Pomel's Camps, 1993); sample consisted of teeth, mandible fragments, and antlers. In - Mugharet el Aliya (Arambourg, 1967); these sites, lower teeth showed interlobar columns and marked - Bizmoune (Bouzouggar et al., 2010). cingulum, as previously indicated for upper molars by Lydekker. In 1893, Pomel described the same species from a cave at Cap Carbon near Bougie and again from Berrouaghia, so we suspect, consid- 2. Systematic position of M. algericus ering the descriptions, that part of this material was the same as that published in 1892. Without morphological observations, The phylogenetic lineages of Pleistocene large-sized deer attrib- fl C. pachygenys was again brie y mentioned at les Bains-Romains in uted to the Megalocerini tribe, referred to as the so-called verticornis the Gulf of Bougie by Ficheur and Brives (1900), who also made group, as opposed to the giganteus group of “giant deer” (Azzarolli, reference to the site of Guyotville (perhaps the site currently known 1953, 1979), have been the subject of much controversy. as Sintes) and Pointe Pescade, while Debruge (1907) followed suit Ambrosetti (1967) and Azzaroli (1979) first related the European for the cave of Ali Bacha. Taking all these Algerian sites into verticornis group with Megaceroides algericus based on the shape of consideration and adding le Pic des Singes, Joleaud (1912) was the the frontal bone from skull fragments of this species published by fi rst to compare C. pachygenys to the genus Megaloceros and to Arambourg (1932, 1938). Although these two groups, including highlight the more evolved dental morphology of the former. Then, insular endemic Pleistocene Cervids from Crete, Sardinia, and Cor- he put C. pachygenys in synonymy with the type species of the sica, seem to represent two separate lineages, the nomenclature fi subgenus, C. (Megaceroides) algericus and discussed its af nities comprises different genera or subgenera to characterize the same with Megaloceros and Dama (Joleaud, 1914). Later in 1916, the same taxa (see Van der Made, 2006; Van der Geer et al., 2010; Vislobokova, author published an exhaustive and comparative overview of Cer- 2013). Consequently, there is still no consensus as to whether to use vids, focusing on the Megacerini tribe in Europe and Asia. He also either Megaloceros Brookes (Lister, 1993, 1994; Pfeiffer, 2002; Van “ ^ ” “ ascribed the few teeth and bones from the Golfe de Genes in Pise der Made and Palombo, 2006), which could be regarded as an in- ” “ ’ (monte Uliveto) and the breches de Nice, d Antibes et de dependent clade closely related to extant Dama (Lister et al., 2005; ” Gibraltar to C. (Megaceroides) algericus, making reference to the Hughes et al., 2006), Megaceroides Joleaud (Ambrosetti, 1967; drawings of Cuvier (1835). This material was probably subse- Azzaroli, 1979; Azzaroli and Mazza, 1992, 1993; Abbazzi and quently lost, as no further mention of it occurs in the literature after Masini, 1997; Abbazzi et al., 1999; Abbazzi, 2004)orPraemegaceros the study of Joleaud (1916). Portis (Geraads, 1986; Croitor and Bonifay, 2001; Croitor, 2006), the

3. Cranial morphology and pachyostosis of M. algericus and of some other Artiodactyla

M. algericus is still known most entirely from cranial elements, the post-cranium being practically unknown (Lydekker, 1890; Pomel, 1893; Joleaud, 1914, 1916; Arambourg et al., 1934; Arambourg, 1938a, 1938b). This material includes antlers from Phacocheres (Hadjouis, 1990), together with the original Algerian material from the sites of Filfila, Taza I (formerly La Madeleine) and Guyotville and from the Moroccan site of Aïn Tit Mellil (Abbazzi, 2004; Croitor, 2006). The general description of M. algericus by Abbazzi (2004) pointed out several distinctive features, including; mandibular pachyostosis; expanded tympanic bullae that could be related to an open environment; the strongly ossified skull with flattened and very divergent antler beams with a basal segment without tines; small praeorbital fossa, quasi-absence of ethmoidal fissures; very short muzzle and reduced size, intermediary between that of Dama dama and Cervus elaphus. We know little about the mandibular pachyostosis which is the characteristic feature of M. algericus, but is a more or less developed trait in other giant deers. For example, the former name of S. pachyosteus was derived from the peculiar, very thick horizontal ramus, as shown by the index of robustness (Van der Made and Tong, 2008, Fig. 13). In M. giganteus the mandibular pachyostosis is characterized by the deposition of fibro lamellar bone from the first stage of adult life onwards (Lister, 1994). In this latter species as well as C. elaphus and Rangifer tarandus there is no significant evidence of additional bone material for long bones (Sander and Andrassy, 2006). In the lower Miocene giraffoid Lorancameryx pachyostoticus of Spain, however, the diaphyses, especially those of the front limbs, are affected by this pachyostosic phenomenon. It could be connected to the development of the frontal appendages that appeared on species at the beginning of the Miocene and continued to expand regularly in several lineages (e.g. Cervidae, Climacoceratidae, Lago- merycidae, Bovidae, Giraffidae, Antilocapridae, Hoplitomerycidae, Palaeomerycidae) (Morales et al., 1993). These last authors have emphasized that horn cores in bovids, ossicones in giraffids and antlers in cervids could maintain homeostasis with a good balance fl between circulatory system, skeleton mass and body weight uc- Fig. 2. Bizmoune Mandible scan views of M. algericus (BM07e58, C3) from NextEngine © tuations in species strongly affected by seasonality. From this point 3D Scan .A¼ anterior face; B ¼ posterior face; C ¼ antero-lingual face; D ¼ occlusal of view the development of the mandibular pachyostosis in face (photography from P. Fernandez). M. giganteus could be an advanced physiological adaptation for the

Please cite this article in press as: Fernandez, P., et al., The last occurrence of Megaceroides algericus Lyddekker, 1890 (Mammalia, Cervidae) during the middle Holocene in the cave of Bizmoune (Morocco, Essaouira region), Quaternary International (2015), http://dx.doi.org/ 10.1016/j.quaint.2015.03.034 P. Fernandez et al. / Quaternary International xxx (2015) 1e14 5 fast growth of huge antlers during relatively short foraging seasons being incomplete, as well as the parastylid (Fig. 2D-7). The (Croitor, 2006). This is also the only example where the mandibles of buccal side of the protoconid is more rounded than that of the males seem to be more pachyostotic than those of females, ruling hypoconid, which has a “V” shape, especially on M/2 (Fig. 2D-8). out dietary adaptation as an explanation. It is true that in feral On M/1, a low isolated ectostylid is present within the interlobal populations of the Northern Hemisphere, C. elaphus (Clutton-Brock space (Fig. 2D-9). Another antero-buccal column, with a and Albon, 1982; Okarma, 1984), Capreolus capreolus (Cederlund damaged upper part, also rises from a basal cingulum (Fig. 2D- and Lindstrom,€ 1983)orOdocoileus virginianus (Carroll and Brown, 10). On M/2 the morphology is much the same with the pres- 1977) for example, males are often heavily affected by weight fluc- ence of these two well-developed columns (Fig. 2D-11 and 2D- tuations and the metabolic syndrome of under-nutrition. This syn- 12). The main differences are the much deeper valley between drome has been described at length by Bassano and Mussa (1998) the paraconid and the metaconid (Fig. 2D-13), and the entoconid and linked both to territorial control and seasonality (e.g. males (Fig. 2D-14) which is closed and separated from the entostylid fighting for females; energy deficit in winters). Nevertheless, none of on M/2 (Fig. 2D-15). these modern species have shown evidence of correlation between Biometric data for dentition of M. algericus are scarce. Never- the development of mandibular pachyostosis and the growth of theless, we undertook a comparison of the Bizmoune material, antlers and weight fluctuations during their evolution. In addition, based on a review of the available data in original publications in Pleistocene lineages of cervids subject to marked seasonality, (Pomel, 1893; Joleaud, 1916; Hadjouis, 1990) and the study of greater differences in the development of pachyostosis should be Abbazzi (2004, Table 1) from specimens stored in the National expected between males and females, but this has not been yet Museum of Natural History of Paris. Datasets for each measurement demonstrated (Lister, 1994) with the exception of M. giganteus. are small for M. algericus, which is why we derived confidence in- Moreover, we do not yet know whether the annually shed cranial tervals from the median (IC with a ¼ 0.05) in Table 2 using R appendages were present in both sexes in the earliest forms of software (R Version 2.14.0). Due to the fact that the Bizmoune cervids or only in males (Van der Made and Tong, 2008; Davis et al., mandible is damaged, the biometric comparison of the mandibular 2011). Given the current state of knowledge, we cannot reach a clear bone is difficult, but the thickness below the anterior and posterior conclusion concerning the respective relationship between the part of M/2 respectively indicates a rather large-sized specimen evolution of the seasonal development of antlers, bone mandibular (Table 2, measurements 9 and 10). As described previously, the microstructure, and the physiological processes involved in the anterior part of the horizontal ramus below M/1 is of reduced size development of mandibular pachyostosis of M. algericus. (Table 2, measurement 11), while the height of the anterior part below M/2 is large compared to the only measured specimen from 4. Material and method Berroughia (Table 2, measurement 12). In addition, the measurements of the Bizmoune molars, some of which correspond to the The terminology, the limits and the subdivisions of the three maximal limit of variation, confirm that they belong to a large in- € main chrono-cultural periods used here follow Linstadter et al. dividual. This is the case for the maximum length and to a lesser (2012). The Upper Paleolithic, including the Iberomaurusian cul- degree for the maximum width measurements which appear to be ture, which corresponds to the last part of the Pleistocene. The significantly larger for both the M/1 and M/2 from Bizmoune in Epipaleolithic and Neolithic both occur within the Holocene period comparison with other specimens (Table 2, measurements 1, 6, 7 (see Fig. 3 for the limits). and 8). We used the RenDateModel 5.0.0.1 calibration software (Lanos and Dufresne, 2012) and radiocarbon dates are presented either 5.2. Archaeological records and dating of M. algericus in the 14 as conventional C age (ka BP), or as calibrated ages on the cal- sequence of Bizmoune endar scale (cal BP), corrected by the IntCal09 calibration curve from Reimer et al. (2009). In order to show the distinctive One of the aims of this article is to focus on the lifespan of the mandibular cylindrical volume typical of M. algericus, we used the species M. algericus, hitherto only biochronologically attested in © e NextEngine 3D scanner (Fig. 2A C). Maghreb from the beginning of the Late Pleistocene until the first part of the Holocene and only dated by scarce indirect radiocarbon 5. Results dates (Table 1, Fig. 3). During the 2007 field season, our preliminary dig in Bizmoune was focused on the elongated west to 5.1. Morphometrical features of M. algericus from Bizmoune east excavation trench at the entrance of the cave over an area of 2 m long and 1.5 m wide. The sequence, which never reached The Bizmoune mandible is damaged but clearly shows that bedrock, is then more than 2 m deep. The seven individual sedi- the anterior part of the horizontal ramus below M/1 is cylin- mentological units can be identified based in different color and drical (Fig. 2A) and becomes regularly thicker and deeper in the texture of the sand matrices. The association of 14C dates with posterior part below M/2 (Fig. 2B and C). The early diagnosis of mammal remains, lithics, and ceramic artefacts provided signifi- the maxilla made by Lydekker (1890) also brought to light the cant chronostratigraphic information for each layer (Table 3). The characteristic brachydonty, the very marked internal cingulum Helix aspersa land snail shells used for the AMS radiocarbon and accessory columns. All these morphological features are samples were collected from three successive layers, from the most well represented on the lower jaw (Hadjouis, 1990; Croitor, recent to the oldest: C2 (Rabat-288, 4283 ± 130 ka BP), layer C3 2006). In the Bizmoune specimen, the lower molars are very (Rabat-289, 7467 ± 172 ka BP) and layer C4 (Rabat-290, brachydont with vertical striations on the enamel, which is 10,865 ± 208 ka BP). The dates are well ordered and consistent particularly thick on the buccal side, especially on M/2 (Fig. 2D- with the stratigraphy. To avoid age anomalies due to bulk samples 1) with a well-developed cingulum bulge on the lingual side of of shells that might come from different layers, each dated sample this tooth (Fig. 2C). On the buccal side of the molars, the consisted of the shell of a complete individual. Nevertheless, like cingulum is represented by a very thin basal convexity (Fig. 2D- the well-known “old wood effect” on bulk samples of charcoal, 2). On M/1, the metaconid and entoconid (Fig. 2D-3 and 2D-4) modern studies based on terrestrial snail shells recovered from are inflated on the lingual side and the same is true for the limestone areas have shown 14C anomalies (producing old radio- metastylid and the entostylid (Fig. 2D-5 and 2D-6); the latter carbon ages) as the result of the ingestion of old carbonate

Table 2 Measurements of the lower dentition of Megaceroides algericus. 1: Occlusal length. 2: Length at the neck. 3: Occlusal width (anterior lobe). 4: Width at the neck (anterior lobe). 5: Occlusal width (posterior lobe). 6: Width at the neck (posterior lobe). 7: Maximum length. 8: Maximum width. 9: Mandibular thickness below anterior part of M/2. 10: Mandibular thickness below posterior part of M/2. 11: Mandibular height below anterior part of M/1. 12: Mandibular height below anterior part of M/2. 13: Mandibular height below posterior part of M/2.

Rank Locality/Collection Side Measurements

1 2 3456 7 8 910111213

Bizmoune (BM07-58, C-3) s 27.00 32.50 22.50 26.00 37.00 a Berroughia 25.00 24.00 25.00 Filﬁla (Fil166)b s 27.00 Filﬁla (Fil167)b 32.70 Guyotvilleb (336) 30.30 Mandible Guyotvilleb (337) 28.60 Taza 1 d 31.00 (ex Madeleine)b Taza 1 s 32.80 b 1 (2015) xxx International Quaternary / al. et Fernandez P. (ex Madeleine) ICa¼0.05 [28.63e32.76]

Bizmoune s 19.70 10.90 13.30 12.60 13.80 19.70 13.80 (BM07-58, C-3) a Berroughia 17.00 Bougie/Coll. Joleaudc 17.00 Coll. Joleaudc 17.00 15.00 Coll. Univ. Lyonc 15.00 11.00 M/1 Filﬁla (Fil166)b d 17.00 15.70 13.00 17.00 13.00 Guyotville (337)b 18.10 18.00 13.40 18.10 13.40 Aïn Benian 16.50 (Ex Guyotville)d

eaeodsalgericus Megaceroides Taza 1 d 19.00 17.80 12.50 19.00 12.50 b (ex Madeleine) ICa¼0.05 [16.29e18.43] [11.82e14.57]

Bizmoune s 21.30 11.90 15.80 12.20 14.40 21.30 15.80 (BM07-58, C-3) c e Berroughia 20.00 16.00 14 Berroughiaa 20.00 16.00 Bougie/Coll. Joleaudc 20.00 16.00 Coll. Joleaudc 19.00 15.00

ydke,19 Mmai,Cervidae) (Mammalia, 1890 Lyddekker, Coll. Univ. Lyonc 18.00 14.00 Filfila (Fil167)b 18.10 17.70 13.60 18.10 13.60 M/2 Filfila (Fil166)b d 18.20 15.40 13.30 18.20 13.30 Guyotville (336)b 19.60 16.60 13.90 19.60 13.90 Guyotville (337)b 18.00 15.60 13.80 18.00 13.80 Aïn Benian (Ex Guyotville)d 19.00 14.80 Aïn Benian (Ex Guyotville)d 20.00 13.50 Taza 1 (ex Madeleine)b d 20.30 19.20 13.60 20.30 13.60 Taza 1 (ex Madeleine)b s 17.00 16.00 12.60 17.00 12.60 ICa¼0.05 [17.52e20.32] [15.21e18.28] [12.95e14.24] [18.49e19.86] [13.75e15.09] a From Pomel (1893, p. 41). b From Abbazzi (2004, Tab. 1). c From Joleaud (1916, Tab. IX). d From Hadjouis (1990, Tab. 3). The median interval confidence for measurements (IC with a ¼ 0.05) is derived from R software. uigtemdl ooeei h aeo imue(ooc,Esoiargo) utrayItrainl(05,http://dx.doi.org/ (2015), International Quaternary region), Essaouira (Morocco, Bizmoune of cave the in Holocene 10.1016/j.quaint.2015.03.034 middle the during laect hsatcei rs s enne,P,e l,Tels curneof occurrence last The al., et P., Fernandez, as: press in article this cite Please

Table 3 Faunal inventory, artefacts and radiocarbon dating related to cultural chronology for each layer of Bizmoune. Radiocarbon dates of H. aspersa are from LARATES (Laboratoire de Recherches et d’Analyses Techniques et Scien- tiﬁques, Rabat). They are presented both as conventional 14 C ages (ka BP) and calibrated ages (cal BP), corrected using IntCal09 calibration curve (Reimer et al., 2009) with the RenDateModel 5.0.0.1 calibration software (Lanos and Dufresne, 2012).

Layers Inventory Anatomical Taxa Sample Larates Material Age ka cal BP (ka) Artefact Chronology and elements no. lab. no. BP cultures

1 BM07-S1 right fragment Oryctolagus/Lepus Pottery fragments; undiagnostic of scapula Late/Final Neolithic microlithic ﬂakes of silex BM07-S1 left M2 Gazella sp.

2 BM07-42 right dp4 Sus scrofa S1/C2 Rabat 288 Helix aspersa 4283 ± 130 [3337;2504] BM07-99 right maxillary Sus scrofa Bizmoune with M2 to M3 BM07-42 left dp4 Ammotragus lervia .Fradze l utrayItrainlxx(05 1 (2015) xxx International Quaternary / al. et Fernandez P. 3 BM07-58 left Megaceroides S1/C3 Rabat 289 Helix aspersa 7467 ± 172 [6641; 6009] hemimandibule algericus Bizmoune with M1 to M2 BM07-43 right M2 Ammotragus lervia BM07-29 right M2 Ammotragus Bladelets production lervia Late Epipaleolithic BM07-206 right incisive Ammotragus without pottery lervia BM07-S1 Phanlage III Alcelaphus buselaphus BM07-38 left prox. Alcelaphus metatarsal buselaphus eaeodsalgericus Megaceroides ± 4 BM07-70 left M1-2 Equus sp. S4/C4 Rabat 290 Helix aspersa 10,865 208 [ 11,294; 10,274] Primary and baked Early Epipaleolithic BM07-72 right M1-2 Equus sp. Bizmoune bladelets; bladelets core; few scrappers

5 BM07-285 right cheek thoot Equus sp. e 14

6 BM07-264 left Gazella sp. Tanged pieces; debitage Levallois; Middle Palaeolithic hemimandibule scrapers; convergent scrapers; (Aterian)

ydke,19 Mmai,Cervidae) (Mammalia, 1890 Lyddekker, with M1 to M3 Mousterian points; possible fragment of unifacial point

7 no faunal element

2 reworked BM07-483 left M Equus sp. BM07-484 M3 Phacochoerus aethiopicus BM07-206/38 Phal. II prox. Caprini indet. No number ostrich eggshell Struthio cf. camelus BM07-206/39 right M3 Ammotragus lervia 7 8 P. Fernandez et al. / Quaternary International xxx (2015) 1e14

(Goodfriend and Stipp, 1983; Goodfriend, 1987). However some 5.3. Chronological range of M. algericus in Northern Africa “minute” taxa (i.e. Vallonia, Pupilla, Euconulus and Succineidae) have demonstrated their high potential for dating a variety of Recently, several studies were undertaken using Uranium series, Quaternary sediments and provided reliable 14C ages (Brennan and luminescence or AMS dating for the Middle (MSA) and Late Stone Age Quade, 1997; Pigati et al., 2004). This is also true for genera such as (LSA) in order to provide high resolution dating records for the Catinella, Columella, Discus, Gastrocopta and Succinea, regardless of Maghreb (Barton et al., 2005; Mercier et al., 2009; Richter et al., 2010; the local geologic substrata (Pigati et al., 2010). The use of the land Clark-Balzan et al., 2012; Jacobs et al., 2012; Barton et al., 2013). snail shell carbonates appears to be a good alternative and is Unfortunately, according to published studies, there are no radio- preferable to organic soil 14C analyses for dating strata, as soil or- carbon dates conducted directly on bones or teeth of M. algericus ganics are generally easily contaminated by subsequent root where it is present. Here we discussed for each site the chronological growth, as detailed by Goodfriend et al. (1999). In this study, we range of this species starting from all the available dates related to assumed that H. aspersa was a preferable material for 14C dating as well identified stratigraphic units with the RenDateModel software we did not have any guarantee of the preservation quality of the previously mentioned. As shown in Fig. 3, in the stratified context of bone organics on which the accuracy of 14C dating depends Tamar Hat (Algeria), charcoal gave the latest occurrence of (Stafford et al., 1990). For example, trial samples of bone selected M. algericus with ages of 10,350 ± 375 and 12,450 ± 480 ka BP from for dating may fail the pre-treatment chemistry analysis (e.g. samples ALG-5, 2-99 and ALG-4,1-98 respectively (Brahimi,1969 and protein preservation) (Barton et al., 2005). More importantly, no sample description in ; Rahmouni et al., 1970). Later, new dates on well-preserved bones were available in layer C2, just above the samples MC-817 (Layer 9) and MC-822 (Layer 84/5) from the earliest mandible recovered in layer C3, or in underlying layer C4, to pro- to the oldest part of Tamar Hat, gave a time interval of 16,100 ± 360 vide relative stratigraphic radiocarbon ages. and 20,600 ± 500 ka BP, including coherent successive dates within Layer C1, the most recent layer, yielded fragments of pottery and that period (MC-812; MC-818; MC-820) (details in Saxon et al., 1974, microliths bladelets and rare blades as well as a broken Oryctolagus or p. 50). In Algeria, at Grotte Rolland, M. algericus could be associated Lepus scapula and a Gazella sp. tooth (Table 3). Layer C2 also provided with the date of 13,330 ± 280 ka BP obtained from marine shells fragments of pottery and undiagnostic lithic artefacts associated with (Camps, 1974, p. 273). In that same country, at Taza I, charcoal in A. lervia and a wild form of S. scrofa, which appears in North Africa at probable association with M. algericus provided an age range from the beginning of the Late Pleistocene (Geraads, 1982; Geraads et al., 12,700 ± 220 (GIF-2111) to 11,340 ± 220 ka BP (GIF-2110) (sample 2010) as mentioned for example in Doukkala II (Michel and description in Delibrias et al.,1974). In addition, a bone fragment from Wengler, 1993; Stoetzel et al., 2007; Steele, 2012). In layer C2, the an undetermined species from layer B and a charcoal in layer C were age obtained on H. aspersa is comprised between 3337 and dated respectively to 16,100 ± 1400 ka BP (GIF-6800) and 2504 cal BP and related to the Late/Final Neolithic (see chronological 13,800 ± 130 ka BP (GIF-6799) (Meier and Sahnouni, 1995; Medig reviews in Linstadter€ et al., 2012). Layer C3, where the M. algericus et al., 1996). More recently in the oldest layer of Taza I, a radio- mandible was unearthed, covers an age range of 6641 to 6009 cal BP, carbon age with the unique mention of “<39 ka BP” was given for a clearly related to the very Late Epipaleolithic. The mandible is asso- bulk sample of several mammal bones (Meier et al., 2003). In the ciated with A. buselaphus, a species that undoubtedly arrived earlier paleokarst of Sidi Saïd (Tipasa, Algeria), M. algericus as well as other from East Africa (Vrba, 1997) and for which there is no definite evi- taxa typical of the Late Pleistocene were discovered without a dence before the Late Pleistocene (Geraads et al., 2010). The Barbary stratified context (Chaïd-Saoudi, 2013) and associated with a sheep, A. lervia is recorded with the latter species, which is not un- maximum age of 38,130 ± 1320 ka BP on Ribbed Mediterranean usual given its temporal extension (Ouchaou, 2000; Ouchaou et al., Limpet shells Patella ferruginea has been given by 14C(Betrouni, 2003). Layer C3 is characterized by bladelets technology without 1997). The recent study using ESR (Electron spin resonance) on un- pottery. Just below this level, in layer C4, the 14C date indicates an age gulate tooth enamel samples from layers 9 and 10 of Mugharet el range of 11,294 to 10,274 cal BP, corresponding to the Early Epi- Aliya, where Arambourg (1967) identified M. algericus, yielded paleolithic, but only a couple of Equus sp. teeth were recovered. This respectively a time range of 35e60 ka for layer 9 and of 60e100 ka for species is also represented in layer C5 by a single tooth. Layers C4 and layer 10 (Wrinn and Rink, 2003). Finally, M. algericus was identified C5 provided some side scrapers, primary and backed bladelets with a by Arambourg in Taforalt in layer VIII (Roche, 1963, p. 152). Unfor- bladelet core as well as pieces esquillees, related to the Iber- tunately, this material from the Late Pleistocene was never published omaurusian industry (LSA). The production in C4 and C5 could be or further mentioned by Arambourg and may be lost. The study of the related to the very late extension of the Iberomaurusian until the faunal remains from new excavations at Taforalt is in progress and Early Epipaleolithic, as indicated in several studies (Barton et al., there is still no evidence of M. algericus according to E. Turner (comm. 2005; Bouzouggar et al., 2008; Linstadter,€ 2008; Linstadter€ et al., to P. Fernandez). 2012; Barton et al., 2013). In layer C6, the presence of Gazella sp. (BM07-264) is of no significant biochronological value and layer C7 6. Discussion contains no faunal remains. Nevertheless, in these two layers, the lithic industry has been referred to as Aterian with typical tanged 6.1. Where: possible routes of dispersal for M. algericus pieces, Levallois debitage, side scrapers, Mousterian points and a possible fragment of a foliate point that indicate the Late North Af- On the basis of the descriptions of a cervid found in the “breches rican MSA (Bouzouggar et al., 2010). de Nice, d’Antibes et de Gibraltar” (Cuvier, 1835) and related to

Fig. 3. Available radiocarbon ages BP and calibrated associated with M. algericus (see detailed comments in Section 5.2 and 5.3). AMS dates were calibrated using IntCal09 calibration curve (Reimer et al., 2009) and the RenDateModel 5.0.0.1 calibration software (Lanos and Dufresne, 2012). Solid lines under probability density distributions stand for calendar age range (2s) before 1950 and bounds are given just below. The Oxygen isotopic curve GISP2 d18O is drawn starting from the bi-decadal original dataset available at http:// depts.washington.edu/qil/datasets/gisp2_20yr.txt according to the work of Grootes et al. (1993), Meese et al. (1994), Stuiver et al. (1995), Grootes and Stuiver (1997) and Stuiver and Grootes (2000). Gaps in the GISP d18O record are due to the lack of original data respectively for periods of 22,020e22,040, 27,480e27,560 and 37,060e37,120 cal BP. (YD) ¼ Younger Dryas from Penaud et al. (2010) and Heinrich events H1 and H2 sensu Stanford et al. (2011). Relative sea level curves from 19 ka BP (modiﬁed from Collina-Girard, 2001). Calibration after Bard et al. (1990) and three core drillings in coral reefs (A: Tahiti; B: Barbades; C: New Guinea; MWP-1A and B: MeltWater Pulse 1 and 2) from Bard et al. (1996). Chrono- cultural framework is taken from the work of Linstadter€ et al. (2012).

“C. algericus”, Joleaud (1916) was the first to raise the question of et al., 2006). Consequently, the competition between M. algericus the possible movement of this species to Northern Africa through and many African bovids and equids, all grazers or mixed feeders the Strait of Gibraltar. From this point of view, Artiodactyla such as may have been limited. However during the Holocene, the North Cervids, Proboscideans, Hippopotamidae and Suidae are very good African distribution of grazers or open habitat species was pre- swimmers (Schüle, 1993; Sondaar and Van der Geer, 2005) and we dominately affected by the disappearance of two species of Gazella believe that M. algericus as well as C. elaphus and S. scrofa could (G. tingitana, G. atlantica), three species of Equus (E. algericus, easily have crossed the Strait of Gibraltar during the Pleistocene E. mauritanicus, E. melkiensis), and three other hypsodont large- (Geraads, 2010). Other Lower and Middle Pleistocene mammal sized species (Camelus sp., Bos primigenius, Syncerus antiquus) species may have made similar crossings, in both directions, be- (Faith, 2014). According to this author, their common paleoeco- tween Europe and Africa (Geraads, 1980, 1982, 2006, 2008, 2010; logical adaptions (e.g. typical grazers or mixed feeder) imply that Agustí et al., 1986; Palmquist and Arribas, 2001; Gibert et al., environmental mechanisms likely played a role and that the ex- 2003; Aouraghe, 2006; Stoetzel, 2013). During cold periods (i.e. tinctions could reflect severe environmental changes rather than MIS 12, 10 and 6), global sea levels dropped significantly several human impact. M. algericus could have also been affected by such times (Rohling et al., 1998; Waelbroeck et al., 2002). As an example, events but the precise extinction chronology of all these taxa need in Fig. 3, at the beginning of Heinrich event 1 sensu Stanford et al. to be known. Lastely, the very low frequency of the number of (2011), the sea level in the Mediterranean Basin was individuals of M. algericus in Tamar Hat and Taza I indicate occa- around 120 m below the current 0 sea-level, resulting in the sional and opportunistic human hunting, and possibly scavenging emergence of islands in the Strait of Gibraltar (Lambek and Bard, of this species. By this period the antlers and bones might have 2000; Collina-Girard, 2001). Consequently, it would have been been used as raw material for “manufactured” products (Merzoug, possible to cross the sea between Europe and Africa by island 2005, 2012; Merzoug and Sari, 2008). hopping, over a distance of less than 10 km (Collina-Girard, 2009). Finally, a single fragment of M. algericus maxillary bearing only the 6.3. When: climatic events during the lifespan of M. algericus indication “Cyrenaica” (Libya) and conserved in the British Museum also deserves a mention (Arambourg, 1938a, 1938b; Ennouchi, The date of the initial arrival of M. algericus in Algeria and 1953; Hadjouis, 1990; Michel, 1990). However a hypothetical Morocco still remains unknown (Geraads, 2010) and direct arrival from the LibyaneEgyptian or SicilianeTunisian routes needs radiocarbon dates of this species are needed. The few available to be supported by more evidence. Lastly, the relatively small dates in stratigraphic contexts, vary considerably with no gap in geographic distribution and limited time-range of M. algericus may the record of this species within the time period between around be related to the short lifespan of the species, as is often the case for 25,000 to 8000 ka cal BP (Fig. 3). Throughout that time, especially large African mammal fossils (Vrba and DeGusta, 2004). Heinrich events 2 and 1, major decreases in sea surface temperatures and deep-water changes appear to be strongly related to 6.2. How: palaeoecological conditions for the settlement of phases of greater aridity in northwest Africa and Iberia, as docu- M. algericus mented by pollen analysis (Fletcher and Sanchez-Go ni,~ 2008; Rofes et al., 2014). Paleoclimatic records from the Alboran and Whatever the initial parent population discussed above for Atlantic Sea cores indicate sharp oscillations with decreases in M. algericus, chances for speciation greatly increase when vicariant marine and terrestrial temperatures and increasing aridity, espe- or dispersal events occur in small or isolated populations sub- cially during the two last Heinrich events, mainly marked by more jected to new strong selective pressure. Compared to the southern or less important incursions of polar waters into the Mediterra- European peninsulas of the Balkans, Italy and Iberia, the Maghreb nean Basin (Cacho et al., 1999; Hemming, 2004; Moreno et al., exhibits the highest rate of endemic taxa (13.7%), across 855 2005). In the western part of the Mediterranean, sea surface modern species including mammals (with the exception of bats), temperatures were cooler than in the adjacent Atlantic, leading for amphibians, reptiles, butterflies and odonates (Husemann et al., example to the abundance and quasi endemism of the cold water 2014). In this region, the high rates of endemism associated with coccolithophore species Emiliania huxleyi until the end of Heinrich a relatively moderate number of species, generally of Atlan- event 1. This may be related to the restriction in the communi- ticeMediterranean and Saharan origin, suggest that the Maghreb cation between the Atlantic and the Mediterranean through the was an important differentiation and speciation refuge area during Strait of Gibraltar (Flores et al., 2010). However, many other the Pliocene and Pleistocene. In contrast, the mixed faunal ele- ecological marine factors, such as the system of currents in the ments from the Balkan peninsula present an image of a biogeo- Strait of Gibraltar, salinity, and nutrient cycling, may also have graphical melting-pot influenced by a higher number of species played a major role in the evolution of the Mediterranean marine coming from multiple directions at different times (Fernandez, fauna, regardless of sea surface temperatures and climate (Emig 2009; Husemann et al., 2014). In view of the above observations and Geistdoerfer, 2004). Around 14,500 cal BP, just after the end concerning the first appearance of M. algericus in North Africa (i.e. of Heinrich 1 (Phase 3 described by Stanford et al., 2011), until Taza I, Sidi Saïd, Mugharet el Aliya), the speciation process of this approximately 5500 cal BP, the so-called African Humid Phase is species must have occurred before at least 25,000 cal BP (Fig. 3). marked by low dust influx indicating more or less closed vegeta- Since the first settlement, its geographic isolation in Northern tion cover, particularly in the Sahara region, which was charac- Africa (Morocco, Algeria) suggests an allopatric speciation from an terized by a much wetter climate than at present (COHMAP unknown population dispersal. It may be assumed that adaptions Members, 1988; Combourieu-Nebout et al., 1999; Hughes et al., to new ecological conditions led to significant morphological 2011). However, short arid continental episodes have been iden- specialization, as previously observed for the species (e.g. ossified tified during the Early Holocene in the Maghreb (Wengler and skull, pachyostosis, specialized teeth, reduced size …). For Vernet, 1992). For example, malacological records in the “Blirh example, if the presence of interlobal columns and cingulum fluvio-lacustrine formation” in Ksabi Basin (Morocco) have shown development are not believed to reflect the diet of Cervids, the two phases of marked desiccation (respectively at 10,500 and brachydonty of M. algericus suggests a diet based on non-abrasive 9280 cal BP) (Limondin-Lozouet et al., 2013). The penultimate food as opposed to the high-crown type more adapted to coarse record of M. algericus ends precisely with the cooling event of grass forage (i.e. browsers vs grazers) (Palombo, 2005; Croitor 8200 cal BP (the so-called 8.2 ka Event), which is clearly identified

Please cite this article in press as: Fernandez, P., et al., The last occurrence of Megaceroides algericus Lyddekker, 1890 (Mammalia, Cervidae) during the middle Holocene in the cave of Bizmoune (Morocco, Essaouira region), Quaternary International (2015), http://dx.doi.org/ 10.1016/j.quaint.2015.03.034 P. Fernandez et al. / Quaternary International xxx (2015) 1e14 11 from the GISP2 curve (Fig. 3). Cooler than the Younger Dryas, References which probably entailed the drying of the Sahara Desert (Carlson, 2013), this cold event corresponds to the main Holocene abrupt Abbazzi, L., 2004. Remarks on the validity of the generic name Praemegaceros Portis 1920, and an overview on Praemegaceros species in Italy. Rendiconti Fisiche climatic reversal. It was caused by the catastrophic meltwater of Accademia dei Lincei 115e132. S. 9, 15. the glacial lakes Ojibway and Agassiz in northeastern North Abbazzi, L., Masini, F., 1997. Megaceroides solilhacus and other deer from the middle America, which slowed the Atlantic meridional overturning cir- Pleistocene site of Isernia La Pineta (Molise, Italy). Bollettino della Societa Paleontologica Italiana 35 (2), 213e226. culation (so-called AMOC) and cooled the global climate (Cronin Abbazzi, L., Croitor, R., David, A., 1999. Megaceroides obscurus (Azzaroli, 1953) from et al., 2007). Did this specific event of 8200 cal BP, characterized early Pleistocene sites of Eastern Moldova. Acta zoologica cracoviensia 42, by arid and cooler conditions in North Africa (Alley et al., 1997), 377e392. ‘‘ ’’ play a major role in the extinction of M. algericus? Some key Adkins, J., deMenocal, P.B., Eshel, G., 2006. The African humid period and the record of marine upwelling from excess 230Th in Ocean Drilling Program Hole studies have pointed out that there had been an underlying trend 658C. Paleoceanography 21, PA4203. http://dx.doi.org/10.1029/2005PA001200. toward climatic deterioration since at least 8500 cal BP that lasted Agustí, J., Moya-Sola, S., Pons-Moya, J., 1986. Venta Micena (Guadix-Baza Basin, until around 8000 cal BP (Rohling and Palike,€ 2005; Adkins et al., South-eastern Spain): its place in the Plio-Pleistocene mammal succession in Europe. Geologica Romana 25, 33e62. 2006). Alley, R.B., Mayewski, P.A., Sowers, T., Stuiver, M., Taylor, K.C., Clark, P.U., 1997. Holocene climatic instability: a prominent, widespread event 8200 yr ago. Geology 25, 483e486. Ambrosetti, P., 1967. Cromerian fauna of the Rome area. Quaternaria 9, 267e284. 7. Conclusion Aouraghe, H., 2006. Histoire du peuplement paleolithique de l'Afrique du Nord et dynamique des interactions entre l'homme et son environnement. Comptes The estabishment of M. algericus as an endemic species in North Rendus Palevol 5, 237e242. Arambourg, C., 1931. Observations sur une grotte a ossements des environs d'Alger. Africa was achieved at least before Heinrich event 2 around Bulletin de la Societ e d'Histoire naturelle de l'Afrique du Nord 22, 169e196. 24 ka cal BP. A typical browser, M. algericus was not directly in Arambourg, C., 1932. Note preliminaire sur une nouvelle grotte a ossements des competition for resources with the grazers, mixed feeders or the environs d'Alger. Bulletin de la Societe d'histoire naturelle d'Afrique du Nord, Alger 23, 154e162. only other browser, Camelus sp., also found in Holocene in North Arambourg, C., 1935. La grotte de la Carriere Anglade a Guyoville (Departement Africa. It is clear that M. algericus had perfectly adapted to new d'Alger). Bulletin de la Societ ed ’histoire naturelle d’Afrique du Nord, Alger 26I, ecological conditions with specialized morphological features such 15e22. as mandibular pachyostosis that are not only related to the seasonal Arambourg, C., 1938a. Mammiferes fossiles du Maroc. Memoires de la Societe des Sciences Naturelles du Maroc 46, 1e74. body weight fluctuations and growth of antlers in cervids. Although Arambourg, C., 1938b. La faune fossile de l'Aïn Til Mellil (Maroc). Bulletin de la speciation and sub-speciation can be documented during the Societ edePr ehistoire du Maroc n1e4, 97e101, 12eme annee . ’ Quaternary, there is little evidence to accurately link specific short- Arambourg, C., 1967. APPENDIX A: observations sur la faune des Grottes d Hercule pres de Tanger, Maroc. In: Howe, B. (Ed.), The Palaeolithic of Tangier, Morocco: term climatic events with mammal speciations (Lister, 2004; Excavations at Cape Ashakar, 1939-1947, vol. 22. American School of Prehistoric Barnosky, 2005). This is also true for extinction processes, as Research, Peabody Museum, Harvard University, pp. 181e186. exemplified by M. algericus in this study. The long gap between the Arambourg, C., Boule, M., Vallois, H., Verneau, R., 1934. In: Masson (Ed.), Les grottes paleolithiques des Beni Segoual (Algerie), Archives de l'Institut de Paleontologie so-called 8.2 ka event and the last record of M. algericus at Biz- humaine, vol. 13, p. 242. Mem . moune between 6641 and 6009 ka cal BP, suggests that this species Azzaroli, A., 1953. The deer of Weybourne Crag and forest bed of Norfolk. Bulletin of successfully faced sporadic events throughout its lifespan. Never- the British Museum (Nat. Hist.), A: Geology 2 (1), 1e96. Azzaroli, A., 1979. Critical Remarks on Some Giant Deer (Genus Megaceros Owen) theless, the long-term climatic changes associated with the quasi from the Pleistocene of Europe. Palaeontographia Italica, 71. In: Nuova Serie, absence of dispersal out of the Maghreb may have affected the basic vol. XLI, pp. 5e16. Pisa. demographic parameters (i.e. survival rates) of M. algericus. In this Azzaroli, A., Mazza, P., 1992. On the Possible Origin of the Giant Deer Genus Meg- aceroides. Rendiconti Fisiche Accademia dei Lincei, pp. 23e32. S.9, 3. sense, the initial population could have crossed the Strait of Azzaroli, A., Mazza, P., 1993. Large early Pleistocene deer from Pietrafitta lignite Gibraltar and probably split regularly in North Africa into many mine, Central Italy. Palaeontographia Italica 80, 1e24. geographical isolated cohorts until extinction. Finally, the possi- Bard, E., Hamelin, B., Fairbanks, R., Zindler, A., 1990. Calibration of the 14C timescale e bility of hunting pressure by Iberomaurusians should not be over the past 30 000 years using mass spectrometric U Th ages from Barbados corals. Nature 345, 405e410. completely discounted, in particular for small-sized populations, Bard, E., Hamelin, B., Arnold, M., Montaggioni, L., Cabioch, G., Faure, G., Rougerie, E., but such evidence is still lacking. 1996. Deglacial sea-level record from Tahiti corals and the timing of global meltwater discharge. Nature 382, 241e244. Barnosky, A.D., 2005. 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