Journal of Human Evolution 96 (2016) 139e144 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol News and views First record of macaques from the Early Pleistocene of Incarcal (NE Iberian Peninsula) * David M. Alba a, , Joan Madurell-Malapeira a, Eric Delson b, c, d, a, Víctor Vinuesa a, Ivette Susanna a, María Patrocinio Espigares e, Sergio Ros-Montoya e, Bienvenido Martínez-Navarro f, g, h a Institut Catala de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Valles, Barcelona, Spain b Department of Anthropology, Lehman College of the City University of New York, 250 Bedford Park Boulevard West, Bronx NY 10468, USA c Department of Vertebrate Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY 10024, USA d New York Consortium in Evolutionary Primatology, New York, NY, USA e Departamento de Ecología y Geología, Universidad de Malaga, Campus Teatinos, 29071 Malaga, Spain f ICREA, Barcelona, Spain g Institut Catala de Paleoecologia Humana i Evolucio Social, Campus Sescelades, Edifici W3, 43007 Tarragona, Spain h Area de Prehistoria, Universitat Rovira i Virgili, Avda. Catalunya 35, 43002 Tarragona, Spain article info Article history: papionins during the Late Miocene, with M. sylvanus being its Received 3 March 2016 basal-most extant species (Springer et al., 2012). Fossil evidence Accepted 12 May 2016 suggests in turn that early members of the macaque lineage Available online 11 June 2016 dispersed into Eurasia by the latest Miocene, roughly coinciding with the sea level drop associated with the Messinian Salinity Crisis Keywords: (Alba et al., 2014, 2015). Old World monkeys Cercopithecidae Although macaques were widely distributed across Eurasia Papionini during the Plio-Pleistocene, unlike in Asia they did not experience Macaca an impressive species radiation. There is consensus (e.g., Szalay and Pleistocene Delson, 1979; Delson, 1980; Alba et al., 2011, 2014) that all the Eu- ropean fossil macaques belong to the lineage of the extant M. sylvanus. The two earliest (latest Miocene) European macaque records (Kohler€ et al., 2000; Alba et al., 2014) are attributed to cf. Macaca sp., while the macaque fossil remains from Sardinia are recognized as an extinct, endemic insular species, Macaca majori Azzaroli, 1946 (Zanaga, 1998; Rook and O'Higgins, 2005). The remaining fossil macaque remains from Europe, following Szalay 1. Introduction and Delson (1979) and Delson (1980), are customarily attributed to various extinct subspecies of M. sylvanus (e.g., Alba et al., 2008, 1.1. Fossil European macaques 2011; Marigo et al., 2014). Due to the lack of complete cranial re- mains, these extinct subspecies are not particularly well charac- Macaques belong to the speciose and widely-distributed genus terized, being largely delimited on biochronological grounds Macaca Lacep ede, 1799 (Cercopithecidae: Papionini), which (Delson, 1980; Alba et al., 2008, 2011; Marigo et al., 2014): currently includes twenty one Asian species and a single North M. sylvanus prisca Gervais, 1859, Early Pliocene; M. sylvanus flor- African species, the Barbary macaque or Macaca sylvanus (Linnaeus, entina Cocchi, 1872, Late Pliocene to Early Pleistocene; and 1758), which was also introduced into Gibraltar in historical times M. sylvanus pliocena Owen, 1846, Middle and perhaps Late Pleis- (Fooden, 1976, 2007; Modolo et al., 2005; Masseti and Bruner, tocene. The local extinction of M. sylvanus in Europe during the Late 2009; Majolo et al., 2013; Roos and Zinner, 2015). According to Pleistocene may be attributable to a combination of different molecular analyses, macaques diverged from other extant ecological factors, such as climatic fluctuations, concomitant vegetation changes, and small population sizes, although human pressure and direct predation might have played major roles as * Corresponding author. E-mail address: [email protected] (D.M. Alba). well (Meloro and Elton, 2012; Elton and O'Regan, 2014). http://dx.doi.org/10.1016/j.jhevol.2016.05.005 0047-2484/© 2016 Elsevier Ltd. All rights reserved. 140 D.M. Alba et al. / Journal of Human Evolution 96 (2016) 139e144 Despite the wide geographical and temporal distribution of 1999; Anton et al., 2005; Ros-Montoya et al., 2012). To date, no macaques during the European Plio-Pleistocene, their fossil record primate remains have been found in any of the more than fifteen is quite sparse, thereby suggesting that they were not very abun- fossiliferous infills discovered (Madurell-Malapeira and Martínez- dant, probably due to suboptimal ecological conditions (Meloro and Navarro, in press). Elton, 2012; Elton and O'Regan, 2014). Here we report the first The Early Pleistocene limestones of Crespia have yielded abun- primate find from the Early Pleistocene site complex of Incarcal (NE dant plant but only a few vertebrate remains (Ros et al., 2003; Iberian Peninsula): an upper molar identified as a macaque that Galobart et al., 2010). Most of the vertebrate fossils come from was found during the 2015 field season. the karstic infills of the Incarcal site complex, which consists of a series of funnel-shaped dolines filled with fossiliferous clays that 1.2. The Incarcal site complex originated due to the karstification (by dissolution and collapse) of the limestones (Ros and Palomar, 2003; Galobart et al., 2010). The The fossiliferous deposits from Incarcal (Crespia, NE Iberian fossil reported here comes from the most-extensively excavated Peninsula; Fig. 1) were discovered shortly after the Cal Taco car- site, Incarcal I, which has provided 1576 large mammal remains bonate quarry began to be exploited in 1967, and take their name (Table 1). The overrepresentation of carnivorans and the frequent from the acronym of the former excavation company (Industrias de recovery of bones in anatomical connection suggest that these Carbonato de Cal; Maroto et al., 2003a; Galobart et al., 2010). Initial karstic cavities functioned as vertebrate traps (Galobart, 2003a), in surveys resulted in the publication of plant macroremains from the agreement with the very short transportation distance inferred limestones of Crespia( Villalta and Vicente, 1972; Roiron, 1983), from the abundance of unfragmented long bones and the excellent from the lacustrine Incarcal Formation (Banyoles-Besalú Basin), as state of preservation of the fossils (Galobart, 2003b). However, well as the publication of abundant late Villafranchian vertebrate sedimentological evidence suggests that bones were transported fossils from the karstic infills developed in these limestones (Julia by water flow into the dolines (Canals and Galobart, 2003). This is and Villalta, 1984). Subsequent surveys and excavations between further supported by current natural ponds from nearby areas, 1984 and 1990 (Maroto and Galobart, 1991; Maroto et al., 2003b; which act as sinkholes that eventually collect animal remains from Galobart et al., 2010) and 1999e2005 (Galobart et al., 2010)were the surroundings after rainy episodes (Madurell-Malapeira and particularly focused on Incarcal I and V. Excavations were resumed Martínez-Navarro, in press). in 2009 by a team led by the Institut Catala de Paleontologia Miquel There are no known age differences among the various karst Crusafont and have continued to the present (Madurell-Malapeira infills (Galobart et al., 2010), although no chronometric dates are and Martínez-Navarro, in press). The fauna from Incarcal has available yet (Madurell-Malapeira and Martínez-Navarro, in been treated in a doctoral dissertation (Galobart, 1996), a journal press). On biochronological grounds, several studies suggested special issue (Galobart and Maroto, 2003), and several scattered an estimated age of ca. 1.5e1.4 Ma (millions of years ago) for the papers (Pons-Moya and Moya-Sol a, 1992; Anton and Galobart, Incarcal sites (Ros-Montoya et al., 2012; Madurell-Malapeira et al., Figure 1. Location map of the site of Incarcal (dot), close to the town of Crespia, and its surroundings; rivers and lakes are indicated in light gray, urban areas in dark gray, and roads in black. The inset in bottom left shows the situation of the depicted area (square) relative to the city of Barcelona (black dot) within the Iberian Peninsula. D.M. Alba et al. / Journal of Human Evolution 96 (2016) 139e144 141 Table 1 2. Description and comparisons Large mammal species from Incarcal I. Updated from Madurell-Malapeira and Martínez-Navarro (in press). The primate upper molar IN-I 1960 (Fig. 2) is temporarily Order Proboscidea housed at the Institut Catala de Paleontologia Miquel Crusafont Mammuthus meridionalis and later will be deposited at the Museu Arqueologic Comarcal de Order Perissodactyla Banyoles. The crown and the mesiobuccal root are completely Stephanorhinus hundsheimensis Equus altidens preserved, whereas the tip of the single lingual root and most of Order Artiodactyla the distobuccal root are missing. The crown shows a moderate Hippopotamus antiquus degree of wear (with very slight dentine exposure at the apices of Bison sp. the buccal cusps, but more extensive dentine exposure on the Bovidae indet. Megacerini indet. lingual ones) and two (mesial and distal) contact facets. It dis- Metacervocerus rhenanus plays a bilophodont and typically papionin occlusal pattern (e.g., Caproleus sp. Delson, 1973, 1975), as indicated by such features as the relatively Order Carnivora well developed median