Quaternary International xxx (2015) 1e14 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint 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. 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 2 P. Fernandez et al. / Quaternary International xxx (2015) 1e14 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) 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 3 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