Journal of Human Evolution 131 (2019) 40e47

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Journal of Human Evolution

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News and Views New macaque remains from the Middle of Gruta da Aroeira ( karst system, Portugal)

* David M. Alba a, , Joan Daura b, Montserrat Sanz b, Elena Santos c, d, Almudena S. Yagüe b, Eric Delson e, f, g, h, a,Joao~ Zilhao~ i, j, k 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 Grup de Recerca del Quaternari (GRQ)-SERP, Departament d’Historia i Arqueologia, Universitat de Barcelona, c/ Montalegre 6, 08001, Barcelona, Spain c Departamento de Geodinamica, Estratigrafía y Paleontología, Facultad de Ciencias Geologicas, Universidad Complutense de Madrid, 28040, Madrid, Spain d Centro Universidad Complutense de MadrideInstituto de Salud Carlos III de Investigacion sobre la Evolucion y Comportamiento Humanos, 28029, Madrid, Spain e Department of Anthropology, Lehman College of the City University of New York, 250 Bedford Park Boulevard West, Bronx, NY, 10468, USA f Department of Vertebrate Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY, 10024, USA g The Graduate Center of the City University of New York, 365 Fifth Avenue, New York, NY, 10016, USA h New York Consortium in Evolutionary Primatology, New York, NY, USA i Departament d’Historia i Arqueologia, Universitat de Barcelona, c/ Montalegre 6, 08001, Barcelona, Spain j Institucio Catalana de Recerca i Estudis Avançats (ICREA), 08010, Barcelona, Spain k UNIARQ-Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras, Universidade de Lisboa, 1600-214, Lisbon, Portugal article info

Article history: although the material remains undescribed. Here we describe a ju- Received 1 November 2018 venile macaque mandible (ARO17-J8-Xa-934, hereinafter ARO 934) Accepted 21 February 2019 recovered in 2017 from a level that is correlated to Marine Isotope Stage 11 and radiometrically dated to 436e390 ka (Daura et al., 2017; see SOM S1 and SOM Fig. S1). We compare it with other fossil Keywords: papionins from Europe to provide a taxonomic assignment. Macaca Mandible Cercopithecoidea 2. Materials and methods Cercopithecidae Papionini Virtual 3D models of the deciduous teeth, the permanent teeth, and the mandibular bone of ARO 934 are provided in SOM Files S1eS3, respectively available from MorphoSource (https://www. morphosource.org). See SOM S2 for further details about the institutional repository of the specimen, CT scanning and image processing. Dental measurements of mesiodistal crown length (MD) and buccolingual crown breadth (BL) were taken to the 1. Introduction nearest 0.1 mm (see SOM S2 for further details). The measurements for the comparative sample are reported in SOM Table S1 (see also The Middle Pleistocene site of Gruta da Aroeira, in the Almonda SOM S2). Bivariate dental plots of BL vs. MD were employed to karst system (, central Portugal; Supplementary Online visually assess dental size and proportions, whereas analysis of Material [SOM] Fig. S1), has yielded a rich lithic assem- variance (ANOVA) and post hoc pairwise comparisons (Bonferroni) blage alongside a human cranium and associated faunal remains were performed to test for significant differences in MD, BL and (Trinkaus et al., 2003; Daura et al., 2017; Conde-Valverde et al., breadth/length index (BLI) for some teeth (see SOM S2 for details). 2018). The recovery of macaques in the 1997e2002 phase of the excavations is mentioned in the literature (Marks et al., 2002a, b; Trinkaus et al., 2003; Brugal, 2004; Marigoetal.,2014 ); the speci- 3. Results mens were assigned to Macaca sylvanus ?florentina (Cocchi, 1872), 3.1. Preservation and ontogenetic development

* Corresponding author. ARO 934 is a juvenile mandible that preserves the symphysis E-mail address: [email protected] (D.M. Alba). and most of both mandibular corpora, as well as most of the left https://doi.org/10.1016/j.jhevol.2019.02.003 0047-2484 D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47 41 ramus except for the coronoid process and the mandibular condyle labially due to damage to the mandibular bone (Fig. 1b, c). The left I2 (Fig. 1aee). The left gonial area and the inferior borders of the left crown is only very partially preserved due to damage in that area of and right corpora are also damaged. The symphysis is well pre- the mandible. Internally, the right and left C1 and P3 germs are served, but a fragment of the mandibular bone is missing on the left located well within the corpus below the dP3, whereas the right I2 is side (mesial to the dP3 roots). The right dP3eM1 and left dP4eM1 obliquely located between the P3 and the I1 germs (Fig. 2; SOM Files series are preserved fully erupted (Fig. 1f, g), whereas the left dP3 is S1 and S2). The right and left P4 and M2 are also inside their crypts, broken away close to the cervix, so that only its socketed roots are the former below the dP4 roots and the latter distally from the M1 preserved (see also Fig. 2 and SOM Files S1 and S2). The deciduous roots (Fig. 2; SOM Files S1 and S2); the right M2 germ is visible in canines and incisors are not preserved, except for the root and inferior view due to the damaged mandibular bone (Fig. 1b). basal-most crown portion of the left dI1 and the apical portion of None of the permanent teeth had completed their development the left dC1 root (which is visible due to damage to the mandibular at the time of death. Even the apices of the mesial and distal roots of bone). Mesially from the latter, there is an empty remnant of the the right and left M1, which are completely erupted and in occlu- left dI2 alveolus. Virtual reconstructions from CT images (Fig. 2; sion, are still open (albeit very close to completion). The develop- SOM File S2) reveal that germs of the permanent incisors, canines, ment of the remaining permanent teeth is comparable between premolars and second molars are inside their crypts, although the antimeres, being more advanced in the incisors (crown completed, apex of the right I1 crown was apparently beginning to erupt above and basal-most portion of the root formed), followed by the pre- alveolar level, and the left I1 and right I2 crown apices were very molars (crown completed but roots beginning to develop just close to the alveolar level as well (they are visible in occlusal view; below the cervix) and the M2 and C1 (crown almost completed but Fig. 1a). Externally, the crowns of these teeth tooth are partly visible cervix not yet completely formed). In macaques, as in most

Figure 1. aee) Photographs of the juvenile mandible of Macaca sylvanus cf. pliocena ARO 934 from Gruta da Aroeira, in occlusal (a), basal (b), frontal (c), right lateral (d), and left lateral (e) views. feh) Photographs of the ARO 934 cheek tooth series (f, g) compared with that of an extant male specimen (AMNH 17409) of Macaca sylvanus sylvanus of com- parable developmental stage (h): f, h) right dP3eM1; g) left dP4eM1 series. 42 D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47

Figure 2. Digital renderings of the juvenile macaque mandible ARO 934 from Gruta da Aroeira, in left lateral (a), right lateral (b), frontal (c), oblique (d), and basal (e) views. The mandibular bone is shown in semitransparent ocher, whereas tooth loci are denoted with different colors. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)

catarrhines, the first permanent tooth to erupt is the M1, followed transverse torus up to the dP3edP4 level. The dental arcade is by the I1 (Smith, 1994; Smith et al., 1994), and available dental vaguely U-shaped, with minimally diverging postcanine eruption times for extant macaques enable us to infer an age at (dP3eM1) toothrows. The corpora are relatively shallow, although death 2 years and most likely closer to 2.5 years (SOM S3). their depth (from alveolar plane to the inferior-most margin), due to damage, can only be measured at the level of the right dP3 (16.2 mm) and the left dP4 (16.0 mm). Postcanine toothrow 3.2. Description length is 23 mm on the right side (dP3eM1). The mandibular corpora widen buccally just behind the M1, at the level of a wide e Mandible The mandible (Figs. 1a e and 2) is approximately 64 mm and long retromolar space (where the M2 crypt is located within long in the anteroposterior direction and 41 mm broad at the the mandibular bone) that is only partly overlapped by a anterior origin of the rami. Once some diagenetic distortion somewhat oblique anterior margin of the ramus. (particularly of the right corpus and the partially preserved right ramus) is discounted, the general shape of the mandible is Deciduous dentition See measurements in Table 1. The dP3 crown comparable to that of Macaca sylvanus (Linnaeus, 1758) of similar (Fig. 3a) displays a suboval to subtriangular occlusal profile, being ontogenetic age (SOM Fig. S2)dexcept perhaps for a longer much longer than wide, and broader distally than mesially. This retromolar space and less vertically oriented anterior margin of tooth possesses two (mesial and distal) roots. The trigonid is the left ramus (which are likely attributable to a slightly more higher than the talonid, and there is some dentine exposure on advanced developmental stage in the Aroeira specimen). The the four main cusps (albeit more restricted on the metaconid). symphysis is 20.5 mm high between infradentale and gnathion; The protoconid is much more mesially located than the anteriorly it is slightly convex in lateral view, whereas posteriorly metaconid, both cusps being linked by a somewhat curved and the planum alveolare is not very steep and extends until middle obliquely aligned protolophid. A long preprotocristid descends of the dP3; there is a moderately developed but distinct inferior mesiolingually until the mesial-most end of the crown (where no D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47 43

Table 1 Permanent dentition See measurements in Table 1. The I1 (Fig. 3d, Dental measurements (in mm) of the deciduous and permanent teeth of the juvenile e) is spatulate and high crowned. The crown is almost symmetrical macaque mandible ARO17-J8-Xa-934 from Gruta da Aroeira. in labial/lingual views (only minimally higher mesially than Tooth Right Left distally), and its unworn apex only displays very faint MD BL BLm BLd MD BL BLm BLd mammelons. The crown displays maximum mesiodistal length at its apex, and progressively tapers basally until becoming dP3 6.8 4.0 4.5 labiolingually wider at the base than longer at the apex. It is dP4 7.4 5.3 5.4 7.1 5.0 5.1 I1 4.5 5.6 4.6 5.5 slightly concave lingually but markedly convex labially. The root ee I2 3.6 5.0 is compressed mesiodistally and longer labially than lingually. C1 5.6 3.3 5.7 3.3 Even from the virtual reconstructions it is evident that the lingual P 5.7 4.0 5.7 4.0 3 side of the crown is devoid of enamel; the apical halves of the P4 5.8 4.9 5.9 4.7 M1 8.3 6.0 6.0 8.5 5.9 6.1 mesial and distal aspects of the crown are almost completely M2 9.5 6.5 6.6 9.4 6.6 6.4 covered by enamel, but from about crown midheight the Abbreviations: BL ¼ labiolingual/buccolingual breadth (perpendicular to maximum cementoenamel junction progressively becomes obliquely crown base dimension in the canine), in the deciduous premolars and the perma- oriented toward the labial aspect of the crown base. ¼ nent molars both at the mesial (BLm) and the distal (BLd) lophs; MD mesiodistal The I2 (Fig. 3f) is also high-crowned and apparently devoid of length (maximum crown base dimension in the canine). lingual enamel, with concave and convex lingual and labial profiles, respectively. This incisor is however smaller and much more distinct paraconid can be discerned), shaping a longer than broad asymmetrical than the I1, the crown being curved from distal to and steep mesial fovea. This fovea is mesially enclosed by a mesial and reaching maximum mesiodistal length at about mid- moderately developed mesial marginal ridge, but lingually open height. The mesial portion of the apical ridge shows a distinct due to the poor development of the premetacristid. The central mammelon, and after a gentle convexity it steeply descends fovea (talonid basin) is subquadrangular and longer than broad, distally, until reaching a cuspulid-like thickening of the enamel, much larger than the mesial fovea, but mostly located on the which is located on the distal aspect of the tooth at about crown lingual half of the crown due to the more peripheral position of midheight and separated from the lingual aspect of the incisor by a the lingual cusps. This fovea is lingually enclosed by the vertical groove. postmetacristid and the somewhat shorter pre-entocristid, and The C1 crown (Fig. 3g, h) is pointed but not very tall relative to distally by the bases of the distal cusps. Wear makes it difficult to the postcanine dentition, being comparable in labial crown height ascertain whether a continuous hypolophid was present, but a (10.1 mm on both sides) to the I2 (and, hence, slightly lower than tiny distal fovea is present on the distal-most portion of the the I1 crown). If the canine mesiodistal axis is defined based on its crown, mesially from the poorly defined distal marginal ridge. On maximum basal diameter, the crown displays a labiolingually the buccal crown wall, between the protoconid and hypoconid, compressed subelliptical to suboval (slightly broader mesially) there is a moderately developed buccal cleft that ends in a basal cross-section, and in labial/lingual views it displays a convex poorly-defined interconule at about crown midheight. The mesial contour and more markedly concave distal profile. The median lingual notch is quite shallow and terminates well before enamel cap extends much farther labially than lingually, where the crown midheight. cementoenamel junction apparently follows an inverted U-shape. The dP4 (Fig. 3b, c) shows a lesser degree of wear than the Two cristids descend from the crown apex in mesiolingual and preceding tooth, with noticeable dentine exposure only at the distobucal directions, separating the convex mesiolabial crown wall apices of the buccal cusps and the metaconid. The crown displays a from the slightly concave lingual and distal aspects of the crown. subrectangular (longer than broad) and slightly waisted occlusal The well-marked mesiolingual cristid abruptly curves dis- outline, being minimally broader at the distal than at the mesial tolingually at about crown midheight, until progressively dis- lophid. The lingual cusps are more peripheral but transversely appearing toward the crown base. The distal cristid, in turn, is less aligned with the corresponding buccal ones. The hypolophid is conspicuous and ends at the distolabial corner of the crown before somewhat broader than the protolophid due to the less buccal reaching its base, forming a small enamel thickening on the left but position of the protoconid compared to the hypoconid. The mesial not the right antimere. There are no distinct cingulids. This fovea is shallow and very restricted (much broader than long, morphology suggests a female individual. almost slit-like) and recurves around the mesial aspect of the The P3 (Fig. 3i, j) is a unicuspid and biradiculate tooth. The crown metaconid, being delimited by a mesially directed preprotocristid displays a suboval occlusal contour, being broader distally than and a slightly longer and obliquely directed premetacristid that mesially, with the single main cusp located slightly mesially and merge with the marked but only slightly mesially-protruding toward the buccal half of the crown. Three steep cristids run from mesial marginal ridge. The mesial end of the tooth is flattened by the crown apex approximately in mesial, distal, and lingual di- contact with the preceding dP3. The central fovea is much larger rections. The mesial and distal cristids delimit the slightly convex than the mesial fovea. Distally from the hypolophid there is a small lingual crown wall from the rather concave mesiolingual and distal distal fovea, delimited by the short posthypocristid and post- aspects of the crown. The mesiolingual portion of the crown dis- entocristid, and to some extent located over a distally-projected plays a sinuous occlusal contour and bears a small cingulid close to extension of the marginal ridge that protrudes from the crown the crown base. The remaining lingual aspect of the crown base is wall and is slightly tilted to lingual. There is a cuspule-like thick- indistinctly bulging below the aforementioned lingual cristid on ening of the enamel at the merging of the postentocristid and the the left side, whereas its right antimere more clearly displays a distal marginal ridge. On the buccal side, there are vertical short secondary cusp. The distal marginal ridge, which merges with the grooves flanking the mesial and distal protrusions of the marginal end of the distal cristid on the distobuccal corner of the crown, ridges, and the buccal cleft is similar to that of the dP3, although distally encloses a small (broader than long) and otherwise somewhat wider and with no distinct interconule. The median lingually-open distal fovea, terminating on a cuspule-like thick- lingual notch is somewhat deeper than in the dP3, but similarly ening of the enamel close to the distolingual corner of the crown. ends before reaching crown midheight. Enamel extends mesiobuccally just onto the mesial root to form a 44 D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47

Figure 3. Digital renderings of the deciduous and permanent lower teeth of the juvenile macaque mandible ARO 934 from Gruta da Aroeira. aec) Deciduous premolars, from left to right in occlusal (mesial on top), buccal, and lingual views: right dP3 (a), right dP4 (b), and left dP4 (c). deh) Anterior permanent teeth, from left to right in labial, mesial, lingual, and distal views: right I1 (d), left I1 (e), right I2 (f), right C1 (g), and left C1 (g). iel) Permanent premolars, from left to right in buccal, occlusal (mesial on top), and lingual views: right P3 (i), left P3 (j), right P4 (k), and left P4 (l). mep) Permanent molars, from left to right in occlusal (mesial on top), buccal, and lingual views: right M1 (m), left M1 (n), right M2 (o), and left M2 (p). honing surface for the upper canine but does not extend along the semicircular distal marginal ridge that thickens slightly at the dis- mesial root as far as on male specimens. tobuccal corner of the crown without forming a distinct secondary The P4 (Fig. 3k, l) is a bicuspid tooth that displays a suboval cuspulid. There are no cingulids. (longer than broad and broader mesially than distally) occlusal Finally, the right and left M1 (Fig. 3m, n), which are only slightly contour. It presents a high trigonid and a much lower talonid. The worn but display some damage at several cusp apices, and the two main cusps are subequal in size, the protoconid being as high larger right and left M2 (Fig. 3o, p), which are preserved as uner- as, but more buccolingually compressed than, the metaconid. These upted tooth germs, resemble the smaller dP4 in overall shape, by cusps are united by a short cristid that separates the small, asym- displaying a subrectangular (longer than broad) and slightly metrical (lingually-directed) and mesially-open (there is no marked waisted occlusal contour with similarly broad mesial and distal mesial marginal ridge) mesial fovea from the deeper and more lobes. The mesial fovea of the lower molars is as restricted as that of extensive distal fovea. The latter is enclosed by a marked and the dP4, but more clearly crescent-shaped, and it is mesially D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47 45

Figure 4. Bivariate dental plots of mesial buccolingual (BL) breadth vs. mesiodistal (MD) length (in mm) for the deciduous and permanent teeth of the juvenile macaque mandible

ARO 934 from Gruta da Aroeira, compared to those of other European Plio-Pleistocene Macaca spp. and Paradolichopithecus arvernensis:a)dP3;b)dP4;c)C1;d)P3;e)P4;f)M1;g) M2. The measurements for the comparative sample are reported in SOM Table S1. Empty symbols indicate mixed-sex samples, yellow and red symbols denote females and males, respectively. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.) 46 D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47 enclosed by a thicker mesial marginal ridge that constitutes a more Marigo et al., 2014). This has led some authors to refrain from protruding ledge-like extension than in the dP4. The M2 marginal providing subspecies assignments (Bona et al., 2016) or to use open ~ ridge is convex instead of straight as in the dP4 and M1, due to the nomenclature (Alba et al., 2008, 2016, 2018; Castanos et al., 2011). lack of an interproximal contact facet. The central fovea of the Our analyses do not substantiate previous suggestions (Szalay molars is similarly extensive to that of the dP4, but in the M1 it is and Delson, 1979) that M. s. pliocena tends to have relatively subsquare rather than subrectangular (slightly longer than broad) wider teeth than other subspecies for the M1 and M2, but indicate as in the dP4 and M2. The distal fovea of the M1 is comparable to larger M1 dimensions on average for both M. s. florentina (see also that of the dP4, except by the lack of any secondary cuspulid, Alba et al., 2011) and M. s. pliocena as compared to M. s. sylvanus. whereas that of the M2 further appears less tilted to lingual and However, the ample overlap among these subspecies and the Gruta slightly more extensive due to the greater distal extension of the da Aroeira specimen for all the lower cheek teeth inspected pre- ledge-like distal marginal ridge. The buccal mesial and distal ver- cludes an unambiguous taxonomic assignment. As noted by Marigo tical grooves are more conspicuous and higher in the M2 than in the et al. (2014), an attribution to the Middle to Late Plestocene M. s. M1 and dP4, and a fainter and short distal lingual groove can be pliocena may be favored on a chronological basis. However, while further discerned in the former molar flanking the ledge-like an assignment to either the extant M. s. sylvanus and the Early extension of the marginal ridge. The median buccal cleft is some- Pliocene M. s. prisca is easy to rule out on the same grounds, an what deeper and better developed in the molars than in the dP4, alternate attribution to the Late Pliocene and Early Pleistocene M. s. although similarly without interconules, and only in the M2 does it florentina is more difficult to dismiss until the differential diagnoses extend below crown midheight (even if without reaching the of the various extinct subspecies of M. sylvanus are further clarified. crown base). Although occlusal relief is somewhat more marked in Consequently, we tentatively assign the macaque mandible from the molars than in the dP4, relative to crown height the median Gruta da Aroeira to M. s. cf. pliocena on chronological grounds, lingual and buccal notches are similarly developed and do not reach pending more comprehensive analyses that could clarify further crown midheight. the alpha-taxonomy of extinct M. sylvanus subspecies. Middle Pleistocene macaque remains assigned to M. s. pliocena 3.3. Metrical comparisons are known from more than thirty localities across Europe, indi- cating a wide geographical range (Elton and O'Reagan, 2014: Ap- The dental size and proportions of both the deciduous (Fig. 4a,b) pendix A) and often being associated with temperate faunal and permanent (Fig. 4ceg) teeth indicate that ARO 934 fits well assemblages and human occupation sites (Szalay and Delson, with a female individual of M. sylvanus subsp., both extant and 1979)das is the case of Gruta da Aroeira (Trinkaus et al., 2003; extinct, being smaller than Paradolichopithecus spp. and larger than Daura et al., 2017; Conde-Valverde et al., 2018). The extinction of Macaca majori Azzaroli, 1946 (see SOM S4 for further details). Some M. s. pliocena in the Late Pleistocene was likely the result of complex minor differences relative to Macaca sylvanus prisca Gervais, 1859 ecological, climatic and stochastic genetic factors, among which (SOM S4) are likely attributable to the small comparative sample anthropogenetic pressure might have played a role (Meloro and sizes for the latter subspecies. ANOVA comparisons indicate sig- Elton, 2012). As such, the factors enabling the co-occurrence of nificant differences among M. s. sylvanus, Macaca sylvanus pliocena macaques and humans in localities such as Gruta da Aroeira and the Owen, 1846, and M. s. florentina for both MD and BL in the M1 but potential interactions between humans and macaques during the not in the M2, with both M. s. pliocena and M. s. florentina displaying Pleistocene deserve further investigation from both paleoecological on average longer and broader M1 than M. s. sylvanus despite a and archaeological viewpoints. considerable overlap (SOM Fig. S3). The M1 of ARO 934 falls above the third quartile of the three investigated subspecies for MD, Acknowledgments probably due to its nearly unworn condition. This work has been funded by and the Agencia Estatal de 4. Discussion and conclusions Investigacion (Ministerio de Economía, Industria y Com- petitividad)eEuropean Regional Development Fund of the Euro- Macaques originated in Africa but are already recorded in pean Union (HAR2017-86509, CGL2015-65387-C3-2-P, MINECO/ Europe during the latest Miocene (Kohler€ et al., 2000; Alba et al., FEDER EU; and CGL2016-76431-P and CGL2017-82654-P, AEI/ 2014). This roughly coincided with a major mammalian turnover FEDER EU), the Ministerior de Economía y Competitividad that took place ca. 5.5e5.3 Ma and, probably thanks to the sea level (‘Ramon y Cajal’ contract RYC-2015-17667 to J.D., and ‘Juan de la drop associated with the Messinian Salinity Crisis (Alba et al., 2014), Cierva’ postdoctoral grant IJCI-2017-33908 to M.S.), the Generalitat involved multiple intercontinental dispersals between Africa and de Catalunya (CERCA Program, and 2017 SGR 00011 to J.D. and Europe (Gibert et al., 2013). Subsequently, macaques are widely M.S.), and the Fundacion Ramon Areces (postdoctoral grant for recorded across Europe from the Early Pliocene to the Late Pleis- your scientists of the Atapuerca Project, granted and managed by tocene (e.g., Elton and O'Regan, 2014), even if represented by the Atapuerca Foundation, to E.S.). Financial support for archeo- fragmentary and generally scarce remains. Except for the extinct logical fieldwork in the Almonda karst sites has been provided by M. majori from Sardinia (e.g., Rook and O'Higgins, 2005), Plio- the Municipality of Torres Novas and projects PTDC HIS-ARQ Pleistocene macaque remains are generally allocated to three 098164 and PTDC/HAR-ARQ/30413/2017, funded by FCT time-successive subspecies of the extant M. sylvanus, currently (Fundaçao~ para a Ciencia^ e Tecnologia, Portugal), with logistical represented in Africa by the nominotypical subspecies (Delson, support from Fabrica de Papel A Renova. We further thank Eileen 1974, 1980; Szalay and Delson, 1979; Alba et al., 2008, 2011, 2014, Westwig (Mammalogy Department, American Museum of Natural 2016, 2018; Marigo et al., 2014). A preliminary analysis of the History) for access to extant primate comparative material, and most complete facial specimen available, from the Middle Pleisto- Sergio Llacer for assistance with the processing of the CT scans. We cene of Albania, supports the current consensus that Plio- also acknowledge the Editor (Mike Plavcan), the Associate Editor, Pleistocene continental macaques belong to the extant species Sarah Elton and two anonymous reviewers for helpful comments (Shearer and Delson, 2012). In contrast, the various extinct sub- that helped us to improve a previous version of this paper. The mCT species of M. sylvanus are not particularly well characterized (Szalay images were obtained in the Laboratory of Microscopy of the and Delson, 1979; Delson, 1980; Alba et al., 2008, 2011, 2016; CENIEHeICTS (Spain) in collaboration with CENIEH staff. D.M.A. is D.M. Alba et al. / Journal of Human Evolution 131 (2019) 40e47 47 member of the consolidated research group 2017 SGR 116 of the hominin cranium from Gruta da Aroeira (Portugal). Proceedings of the National e Generalitat de Catalunya. Academy of Sciences USA 114, 3397 3402. Delson, E., 1974. 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