First record of a fossil monkey (Primates, Cercopithecidae) from the Late Pliocene of Serbia

Journal of Human Evolution 137 (2019) 102681

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First record of a fossil monkey (Primates, Cercopithecidae) from the Late Pliocene of Serbia Predrag Radovi
c a, *, Joshua Lindal b, Zoran Markovi
c c, Sanja Alaburi
c c, Mirjana Roksandic b a b c

National Museum Kraljevo, 2 Trg Svetog Save, 36000 Kraljevo, Serbia Department of Anthropology, University of Winnipeg, 515 Portage Avenue, Winnipeg, Manitoba, R3B 2E9, Canada Natural History Museum in Belgrade, 51 Njegoseva, 11000 Belgrade, Serbia

a r t i c l e i n f o

a b s t r a c t

Article history: Received 29 May 2019 Accepted 16 September 2019 Available online xxx

The cercopithecid fossil record of the Balkan Peninsula extends from the Late Miocene to the Early Pleistocene, but to date no fossils of non-human primates have been identified in Serbia. Here we report the identification of two primate teeth from Ridjake, a rich paleontological site in western Serbia. NHMBEO 042501 is an upper third molar with heavy occlusal wear and taphonomic weathering. NHMBEO 042502 is a well-preserved lower third molar with only minor damage to the cusps and root apices. We performed an analysis of non-metric traits and made bivariate comparisons of crown linear measurements in order to assess the taxonomic affinity of the molars. Both show typical papionin occlusal patterns and relatively large overall sizes. In combination with the early Villafranchian (MN16) age of the site, we attribute both Ridjake primate fossils to cf. Paradolichopithecus sp. This represents the first identification of a non-human primate in Serbia, and the first identification of any primate in the Neogene period of Serbia. Along with recent hominin discoveries, the Ridjake fossils contribute to the growing primate fossil record in Serbia, and indicate the need for increased research into fossil primates in the country. © 2019 Elsevier Ltd. All rights reserved.

Keywords: Papionini MN16 Ridjake

1. Introduction Cercopithecids have a long fossil record in the Balkans that extends from the Late Miocene to the Late Pleistocene. The western Eurasian colobine genus Mesopithecus is well known in the Late Miocene and Early Pliocene (MN11 to MN15) of Greece, Bulgaria, Romania and Northern Macedonia (e.g. Eronen and Rook, 2004 and references therein; Koufos et al., 2003; Radovi
c et al., 2013; Lazaridis et al., 2018). Dolichopithecus, a Eurasian colobine monkey larger in size than Mesopithecus, is also documented in the Pliocene (MN15) of Greece and Bulgaria (Spassov and Geraads, 2007; Koufos, 2009). The cercopithecine genus Macaca dispersed € hler et al., into Europe from Africa in the Late Miocene (MN13; Ko 2000; Alba et al., 2014), and resided there until the Late Pleistocene (Eronen and Rook, 2004). According to the current consensus (e.g. Delson, 1974, 1980; Szalay and Delson, 1979; Alba et al., 2008, 2011) all fossil macaques from Europe belong to the lineage of the

* Corresponding author. E-mail address: [email protected] (P. Radovi
c). https://doi.org/10.1016/j.jhevol.2019.102681 0047-2484/© 2019 Elsevier Ltd. All rights reserved.

living Barbary macaque Macaca sylvanus. Although widely distributed in Europe during the Plio-Pleistocene, macaques are not very common in the Balkans, and the single (to our knowledge) published record comes from the Early Pleistocene of Greece (Symeonidis and Zapfe, 1976; Koufos, 2009). A large-bodied cercopithecine monkey of the genus Paradolichopithecus is recorded in the Early Pliocene of Romania (MN15; Delson, 1974; Szalay and Delson, 1979; Eronen and Rook, 2004), and is also known in the Early Pleistocene (MN17) of Romania and Greece (Necrasov et al., 1961; de Vos et al., 2002; Kostopoulos et al., 2018). Despite the fact that both colobines and cercopithecines ranged over much of Europe and the Balkans, no fossil monkeys have ever been recorded in Serbia. Here, we report fossil dental specimens referable to Papionini (Primates, Cercopithecinae) and originating from the Late Pliocene strata of Ridjake (Western Serbia).

1.1. Geographical and geological setting Ridjake is a village in the municipality of Vladimirci (Ma cva District of western Serbia), located about 55 km SW from Belgrade

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 and about 20 km SE from Sabac (Fig. 1a). The Ridjake paleontological site (44
390 2800 N, 19
450 5000 E) was revealed by accident during construction activities by the local landowners. The geology of the site was first studied in 2004 by Petar Steji
c as a part of a project conducted by the Geological Survey of Serbia (Steji
c, 2009). The fossil assemblage derives from red clayish sands which fill paleokarst depressions of the Middle to Upper Triassic Leli
c Fm; a simplified lithologic section following Steji
c (2009) is presented in Figure 1b. Starting in 2005, several excavation campaigns were conducted at Ridjake by the team of the Natural History Museum in Belgrade. Numerous in situ remains of large mammals were observed upon the first visits to the site. Sieving of the red fossiliferous sediment revealed an extremely rich assemblage of vertebrate remains; for example, a modest ~20 kg sample yielded over 1000 fish, amphibian, reptile, bird, and mammal fossil bones and teeth. In the several subsequent campaigns, a total of more than 1 t of sediment was screened. Although the last samples were collected in 2014, sorting of fossil remains from the residue is still in progress. Hundreds of thousands of osteological and odontological specimens have been separated so far, and this sheer abundance of fossilized remains places the locality of Ridjake as one of the richest vertebrate fossil sites in the world. The largest sieve fraction contained mostly remains of large mammals such as carnivores, artiodactyls and perissodactyls, but also the two isolated primate molars described in this paper. Finer fractions contained both small non-mammalian and mammalian vertebrate remains. Among the latter, the most numerous were Chiroptera, Insectivora and Rodentia.

The following mammalian taxa have been identified so far: Keramidomys mohleri, Estramomys simplex, Blackia sp., Spermophilinus sp., Glis minor, Muscardinus pliocaenicus, Dryomimus sp., Glirulus pussilus, Prospalax priscus, Micromys sp., Apodemus dominans, Apodemus cf. atavus, Miomys occitanus, Pliomys ucrainicus, Baranomys longidens, Allocricetus bursae, Mesocricetus sp., Tapirus, Lagomorpha, Mustelidae, Cervidae, Suidae, Bovidae, Insectivora, Chiroptera. This is the first mammalian faunal list available for the site. Furthermore, Ðuri
c and Radosavljevi
c (2014) also identified the following fossil herpetofauna at Ridjake: Pelobates sp., Ranidae gen. et sp. indet., Bufonidae gen. et sp. indet., Lacerta viridis, Lacerta cf. L. agilis, Lacerta sp., Lacertidae indet., Anguinae gen. et sp. indet., Elaphe quatuorlineata, Zamenis longissimus, Zamenis paralongissimus, Coronella austriaca, Hierophis viridiflavus, Telescopus cf. T. fallax, cf. Telescopus sp., Colubridae indet., Malpolon cf. M. monspessulanus, Vipera berus complex, Natrix sp., and Vipera ammodytes. The recorded small mammalian association places the locality of Ridjake in the early Villafranchian (MN16), representing the first discovery of this type of fauna from Serbia. Further excavations at Ridjake would probably increase the number of identified species (especially of large mammals) and would thus facilitate a more precise correlation with other sites in the Peri-Paratethyian region (Markovi
c, 2008). Additional research is needed in order to obtain reliable data on the site's lithostratigraphy and deposition modes involved. At this stage of research it is apparent that the material had been transported from a wider area of the surrounding karst plateaus, probably via alluvial-fluvial action. This is indicated by the observed fragmentation of fossil remains as well as

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100 100 km 10 km

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Ridjake 44

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Figure 1. Geographical and geological setting of the Ridjake locality. a, location of the Ridjake site (red star) in Serbia (country abbreviations: AL ¼ Albania, BA ¼ Bosnia and Herzegovina, BG ¼ Bulgaria, HR ¼ Croatia, HU ¼ Hungary, ME ¼ Montenegro, NMK ¼ North Macedonia, RO ¼ Romania); b, simplified lithologic section of Ridjake (adapted from Steji
c, 2009) e 1. Massive to layered limestones (LandinianeUpper Triassic); 2. Paleokarst depressions filled with red clayish sands and dark-yellow sand lenses, with abundant vertebrate fauna (Pliocene); 3. Limestone regolith within sandy matrix (Quaternary?); 4. Red to yellow thin-bedded sands (Quaternary); 5. Dark brown loessoid-deluvial deposits with carbonate concretions (Quaternary). The red star marks the approximate stratigraphic position of the Ridjake molars. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)

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by their heterogeneous composition, with the taxa apparently originating from different habitats including forms characteristic of savannah (bovids, large carnivores), steppe (voles, rabbits), and damp forests (tapirs). Numerous long bone fragments bear distinctive rodent incisor gnawing marks, indicating that these bones were exposed on the surface before burial and fossilization (Dirkmaat and Passalacqua, 2012). 2. Materials and methods The fossil specimens described and studied in this paper include two isolated teeth curated at the Natural History Museum in Belgrade (Serbia): NHMBEO 042501 and NHMBEO 042502 (Fig. 2). Dental terminology largely follows Delson (1973, 1975). Measurements were taken using digital calipers to the nearest 0.1 mm. Bivariate comparisons were made using Microsoft Eronen and Rook, 2004 for Windows. Dental measurements for the Ridjake specimens and the comparative sample are reported in Supplementary Online Material (SOM) Table S1. 3. Systematic paleontology Order: Primates Linnaeus, 1758. Semiorder: Haplorhini Pocock, 1918. Suborder: Anthropoidea Mivart, 1864. Infraorder: Catarrhini Geoffroy Saint Hilaire, 1812. Family: Cercopithecidae Gray, 1821. Subfamily: Cercopithecinae Gray, 1821. Tribe: Papionini Burnett, 1828. Genus: Paradolichopithecus Necrasov et al., 1961.
ret, 1929 Species included: Paradolichopithecus arvernensis Depe (type species); P. geticus Necrasov et al., 1961; Papio sushkini Trofimov, 1977; P. gansuensis Qiu et al., 2004. cf. Paradolichopithecus sp. (Fig. 2)

3.1. Description and comparisons NHMBEO 042501 represents an isolated right upper molar, which preserves a damaged crown and a small part of the mesiobuccal root (Fig. 2aee). The tooth is clearly three-rooted, indicating an upper molar (Delson, 1973: 206). The enamel has an irregular,

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rough surface, indicating post-mortem erosion (Martínez and
rez-Pe
rez, 2004). The tooth surface (especially the exposed Pe occlusal dentine and roots) is dotted with dull black deposits; these can be produced by several different agents and processes, among which manganese dioxide staining is common in cave settings
ndez-Jalvo and Andrews, 2016: 156). Red sediment and (Ferna brown-red staining are also present on the surface, the color of
pez-Gonza
lez et al., which is likely due to ferric oxide deposition (Lo 2006). Much of the external morphology of the crown is obscured by weathering and heavy wear. The dentine is heavily exposed on all the cusps, especially on the lingual side, with the merging of wear facets (grade F according to the scoring system of Delson, 1973). There is also a distinct disto-lingual dentine exposure, corresponding to an accessory cusp (distoconule). The crown clearly shows a bilophodont arrangement of the four principal cusps e a hallmark feature of the molars of Old World monkeys (Swindler, 2002). It also shows a typical papionin (tribe Papionini, excluding Theropithecus) occlusal pattern, as indicated by the presence of a well-developed median lingual cleft that ends well before the cervix (Delson, 1973, 1975), obvious despite the enamel erosion (Fig. 2e). The degree of flare could not be quantified (e.g. the index of molar flare sensu Benefit, 1993) due to damage and heavy wear, but it is apparent nevertheless, especially on the lingual side (Fig. 2b, d) which shows characteristic bulging near the cervix (Delson, 1973; Benefit, 1993). The distal fovea (i.e. the primitive talon basin) is less developed than the central fovea (trigon basin). The mesial fovea is worn down to just a small patch of enamel surrounded by the exposed dentine, which, combined with the mesial breakage of the enamel, precludes the proper assessment of this feature. The tooth is most likely an M3 due to its apparent lack of a distal contact facet (although it is impossible to rule one out due to the wear), presence of distoconule, and visually apparent talon reduction. Metrically, the distal tapering is not pronounced (mesial or paraloph maximum BL breadth ¼ 11.2 mm, distal or metaloph maximum BL breadth ¼ 10.7 mm); however, the distal breadth is probably underestimated due to the enamel damage. The mesial portion of the enamel is damaged, so the true mesiodistal length could not be reliably taken (MD length > 11.9 mm). NHMBEO 042502 is a well-preserved right lower third molar (Fig. 2fej), missing only the apical half of the mesial and the tip of the distal root, with minor damage to the apices of the metaconid and entoconid. The tooth's surface is not eroded, and it presents patches of the red sediment and black deposits. The crown shows

Figure 2. Dental remains of cf. Paradolichopithecus sp. from Ridjake (Western Serbia). aee) Right M3 (NHMBEO 042501) in occlusal (a), mesial (b), buccal (c), distal (d), and lingual (e) views. fej) Right M3 (NHMBEO 042502) in occlusal (f), mesial (g), buccal (h), distal (i) and lingual (j) views.

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moderate wear, with restricted dentine exposures at the protoconid (subquadrangular facet), hypoconid (smaller, oval-shaped) and entoconid (smallest, point exposure); although the metaconid wear is obscured by the enamel damage, it was likely less worn than the protoconid. This corresponds to the wear grade C according to the system of Delson (1973). A marked mesial contact facet can also be observed. The fused single root under the mesial lobe (metalophid) is mesiodistally compressed and vertical (judging from the preserved upper portion). The roots under the central lobe (hypolophid) are fused with the root under the third lobe, thus comprising a single distally-inclined complex root with a triangular horizontal section. The morphology of the five-cusped crown is clearly papionin, as evidenced by the bilophodont occlusal pattern, low relief, and the moderate amount of buccal flare (e.g., Delson, 1973, 1975; Szalay and Delson, 1979; Jablonski, 2002). The degree of wear and apical damage precludes quantification of the flare; however, sloping of the crown walls from cusp apices towards roots and bulging near the cervix is obvious, especially on the buccal side (Fig. 2g, i). A pronounced distalward narrowing (tapering) of the crown behind the metalophid and the presence of a distinct third lobe are consistent with the specimen being a lower third molar (Delson, 1973, 1975). The third lobe shows a large and well-developed hypoconulid placed buccal to the midline, as well as a distinct sixth cusp (tuberculum sextum) set between the hypoconulid and the entoconid. The well-developed median buccal cleft and the less-developed mesial and distal clefts all end well before the cervix. The lingual side presents shallow median and distal notches, as is expected for the dentally typical Papionini (Delson, 1975). The mesial fovea (trigonid basin) is relatively shallow, restricted and oval-shaped, with a much greater breadth than length. In comparison, the central fovea (talonid basin) is much deeper and more extensive, showing a subrectangular shape. The distal fovea is even deeper (representing the most basal point of the occlusal surface), but it is less extensive than the central fovea, and it displays a subtriangular shape. The crown dimensions are: MD length ¼ 16.4 mm; metalophid BL breadth ¼ 11.5 mm; hypolophid BL breadth ¼ 9.8 mm; hypoconid BL breadth ¼ 7.7 mm. In sum, both specimens show typically papionin occlusal relief with low cusps, clefts set well before the cervical line and moderately expressed flare (Delson, 1973; Kullmer et al., 2011). Furthermore, the wear patterns are also typical for papionins in which the lingual cusps of the upper and the buccal cusps of the lower molariform teeth wear faster (Delson, 1973; Frost and Kullmer, 2008). NHMBEO 042501 does not show any observable evidence of an interconulus in the lingual projection (Fig. 2e), but the absence of this trait can not be entirely dismissed due to the poor preservation of the enamel. As seen on the buccal side of the much better preserved lower molar from Ridjake (Fig. 2h), there is a distinctive, though not strongly expressed, interconulid (Type 3 according to Hlusko, 2002). The interconulus is an accessory cuspule (style) which may be present at the base of the lingual groove between the protocone and hypocone in maxillary molars of some cercopithecid species; the interconulid is a similar feature, located on the buccal sides of mandibular molars (between the protoconid and hypoconid) (Swindler, 1976; Hlusko, 2002; Monson and Hlusko, 2014). These features are most commonly interpreted as remnants of the primitive mammalian lingual cingula (e.g., Saheki, 1966; Swindler, 1976). As demonstrated by Monson and Hlusko (2014), the interconulus has a significantly higher frequency and degree of expression in papionins than in other closely related taxa, and could thus represent a derived dental trait for the group; moreover, the trait was found to be most frequent in maxillary third molars. Both molars from Ridjake clearly display larger sizes when compared to the data for the extant Barbary macaque (M. sylvanus

sylvanus) and various fossil papionin and colobine groups from Europe (Fig. 3). The only European papionin displaying a dentition this large is Paradolichopithecus, which occurs from the Middle Pliocene to the Early Pleistocene and is well known from the Balkans (see Eronen and Rook, 2004; Kostopoulos et al., 2018). Based on this, the two primate molars from Ridjake most likely belong to the genus Paradolichopithecus. However, two isolated molars (one of which is badly preserved) can not provide a basis for a firm taxonomic attribution, considering the fact that all papionin molars show similar morphologies (Szalay and Delson, 1979; Alba et al., 2014). There is still a possibility (however slight considering the stratigraphy and geography) that the teeth might belong to an as yet unknown species of Paradolichopithecus, or to an unknown papionin genus. This degree of uncertainty prompted us to attribute the specimens to cf. Paradolichopithecus sp., following the recommendations on open nomenclature provided by Bengtson (1988).

4. Discussion and conclusion Based on the morphological (i.e. typical papionin occlusal patterns), metrical (large dental sizes), biochronological (MN16) and

Figure 3. Bivariate plots of maximum mesial buccolingual breadth (in mm) versus mesiodistal length (in mm) for the dental remains from Ridjake compared with extant Macaca sylvanus sylvanus, and European fossil papionins (M. s. florentina, M. s. pliocena, M. s. prisca, M. majori and Paradolichopithecus arvernensis) and colobines (Mesopithecus pentelicus, Dolichopithecus rusciniensis). a, third upper molars (the arrow indicates that the true mesiodistal length of NHMBEO 042501 was higher); b, third lower molars. Data for the comparative sample were taken from NYCEP's PRIMO (Primate Morphometrics Online) database (http://primo.nycep.org) and from the literature (P. aff. arvernensis specimen LGPUT DFN3-150; Kostopoulos et al., 2018). The measurements used for these comparisons are reported in SOM Table S1.

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geographic (Balkans) grounds, the Ridjake molars are here assigned to cf. Paradolichopithecus sp. The genus Paradolichopithecus currently includes four large-sized papionin species, two of which have been named based on the material coming from the PlioPleistocene of Europe: the type species P. arvernensis and P. geticus. The record of P. arvernensis includes specimens such as ze (MN17; Depe
ret, 1929) and Vialette (MN16; those from Sene Delson, 1973) in France, La Puebla de Valverde (MN17; Aguirre and Soto, 1978) in Spain, and Vatera (MN17; de Vos et al., 2002) in Greece. Several partial crania, mandibles and postcranial elements attributed to P. geticus come from Valea Graunceanului, also known unceanu (MN17/MNQ1; Necrasov et al., 1961; Terhune et al., as Gra 2013) in Romania, and some authors (e.g. Delson and NicolaescuPlopsor, 1975; Szalay and Delson, 1979; Kostopoulos et al., 2018) have suggested that the species should be synonymized with P. arvernensis. The earliest Balkan record of the genus comes from the Early Pliocene (MN15) site of Malushteni in Romania (Delson, 1974; Szalay and Delson, 1979). The Asian record includes P. sushkini from Tadjikistan (Trofimov, 1977; Maschenko, 1994) and P. gansuensis from China (Qiu et al., 2004). Based on the presence of some baboon-like cranial features (e.g. large molars relative to premolars; lacrimal fossa positioned within lacrimal bone), Maschenko (1994, 2005) argued that Paradolichopithecus should be reassigned to Papio. However, Paradolichopithecus is usually regarded as the sister taxon to Macaca, which is supported by craniofacial features such as the presence of a maxillary sinus, small anterior dentition and a rounded muzzle dorsum, the lack of a clear anteorbital drop in the midline profile, and the absence of maxillary fossae (Szalay and Delson, 1979; Strasser and Delson, 1987; Delson and Frost, 2004; Rae, 2008; O'Shea et al., 2016). With a body mass (Delson et al., 2000) comparable to the largest living baboons (e.g. Mandrillus sphinx, Papio ursinus, Theropithecus gelada), Paradolichopithecus is easily distinguished from the much smaller Macaca, but it cannot be readily distinguished from Procynocephalus, another large-sized fossil cercopithecine from East Asia (Delson and Nicolaescu-Plopsor, 1975; Jablonski, 2002; van der Geer and Dermitzakis, 2008). Based on the strong similarities in the dental and gnathic remains available at the time, Simons (1970) suggested Paradolichopithecus might be synonymous with Procynocephalus. Others (e.g. Szalay and Delson, 1979; Jablonski, 2002) were more cautious and proposed retaining the two separate genera. However, the recently described cranium from Dafnero-3 (Northern Greece) shows affinities with both the Chinese Procynocephalus and the Eurasian Paradolichopithecus, adding further support for the synonymy of the two genera (Kostopoulos et al., 2018). The paleoenvironmental data from the Plio-Pleistocene of Europe indicates that the genus Paradolichopithecus inhabited both humid, closed environments (MN15 of Romania; MN16 of France and Spain) and more arid, open savannah woodlands (MN16 of Spain; MN17 of Spain, France and Greece) (Eronen and Rook, 2004). Dental microwear analyses conducted on the maxillary dentition of P. aff. arvernensis from Dafnero-3 have suggested that this large papionin monkey may have consumed hard food items, consistent with the hypothesis of the Early Pleistocene spread of grasslands (Plastiras et al., 2017). The postcranial morphology indicates that P. arvernensis practiced a form of terrestrial quadrupedal locomotion similar to that of modern baboons (Jablonski, 2002; van der Geer and Sondaar, 2002; Ting et al., 2004). The paleoenvironment inhabited by the Ridjake papionins is hard to define at this moment, but there is a possibility that it was a mixture of closed (forest) and open (grassland) habitats. In order to provide a more precise paleoecological reconstruction, additional geological research into the processes involved with the formation of the site's fossiliferous strata is needed.

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The identification of cf. Paradolichopithecus sp. at Ridjake marks the first discovery of a fossil non-human primate in Serbia. Along with the recent finds of fossil hominins (e.g. Roksandic et al., 2011; Roksandic, 2016; Radovi
c et al., 2019), the papionin teeth described in this paper add to the growing record of fossil primates from this country. The discovery confirms that primates were present in this region during the latest part of the Neogene period (i.e. early Villafranchian age). This is in agreement with the well-established presence of fossil cercopithecoids in the Mio-Pliocene of other Balkan countries, but also stands in stark contrast to the long history of research on Neogene vertebrates in Serbia (see e.g. Markovi
c and Milivojevi
c, 2010; Milivojevi
c, 2016). Considering the extraordinary richness of Ridjake fossil assemblage, it is highly likely that additional primate specimens will be recovered from this site. Moreover, the discovery of fossil monkeys at Ridjake demonstrates that future systematic research in Serbia certainly has the potential for locating other primate-bearing Neogene strata. Acknowledgements We express our gratitude primarily to senior geological conservator Milos Milivojevi
c (Natural History Museum in Belgrade), a valuable member of the team conducting the field and lab research. We also thank the Maksimovi
c family, the landowners of the fossiliferous locality at Ridjake, Petar Steji
c (Geological Survey of Serbia) for providing access to the unpublished geological data on the site, and SrCan Vulovi
c (National Museum Kraljevo) for photographing the fossils. Some data for this project were downloaded from PRIMO, the NYCEP Primate Morphology Online database (http://primo.nycep.org). We thank Dr. Eric Delson and colleagues for access to these data. Thanks to Michael Plavcan, Eric Delson and one anonymous reviewer for their input on the initial draft. This work was supported by Natural Sciences and Engineering Research Council of Canada (grants no RGPIN-2019-04113 and RGPAS-2019-00039). Supplementary data Supplementary data to this article can be found online at https://doi.org/10.1016/j.jhevol.2019.102681. References Aguirre, E., Soto, E., 1978. Paradolichopithecus in La Puebla de Valverde, Spain: Cercopithecoidea in European Neogene stratigraphy. Journal of Human Evolution 7, 559e565. -Sol Alba, D.M., Moya a, S., Madurell, J., Aurell, J., 2008. Dentognathic remains of Macaca (Primates, Cercopithecidae) from the late early Pleistocene of Terrassa (Catalonia, Spain). Journal of Human Evolution 55, 1160e1163.
Montoya, P., Morales, J., Rook, L., ~ o, M.A., Alba, D.M., Carlos Calero, J.A., Manchen 2011. Fossil remains of Macaca sylvanus florentina (Cocchi, 1872) (Primates, Cercopithecidae) from the Early Pleistocene of Quibas (Murcia, Spain). Journal of Human Evolution 61, 703e718. Alba, D.M., Delson, E., Carnevale, G., Colombero, S., Delfino, M., Giuntelli, P., Pavia, M., Pavia, G., 2014. First joint record of Mesopithecus and cf. Macaca in the Miocene of Europe. Journal of Human Evolution 67, 1e18. Benefit, B.R., 1993. The permanent dentition and phylogenetic position of Victoriapithecus from Maboko Island, Kenya. Journal of Human Evolution 25, 83e172. Bengtson, P., 1988. Open nomenclature. Palaeontology 31, 223e227. Burnett, C.T., 1828. Illustrations of the Manupeda or apes and their allies: being the arrangement of the Quadrumana or Anthropomorphous beasts indicated in the outline. The Quarterly Journal of Science, Literature and Art 26, 300e307. Delson, E., 1973. Fossil colobine monkeys of the circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia). Ph.D. Dissertation, Columbia University. Delson, E., 1974. Preliminary review of cercopithecid distribution in the circum
moires du Bureau de Recherches Ge
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