Journal of Human Evolution 88 (2015) 1e14

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Journal of Human Evolution

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First record of Mesopithecus (Cercopithecidae, ) from the Miocene of the Iberian Peninsula

* David M. Alba a, , Plini Montoya b, Marta Pina a, Lorenzo Rook c, Juan Abella d, a, Jorge Morales e, Eric Delson f, g, a 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 Area de Paleontologia, Departament de Geologia, Universitat de Valencia, c/ Doctor Moliner 50, 46100 Burjassot, Valencia, Spain c Dipartimento di Scienze della Terra, Universita degli Studi di Firenze, via G. La Pira 4, 50121 Firenze, Italy d INCYT-UPSE, Universidad Estatal Península de Santa Elena, 7047 Santa Elena, Ecuador e Departamento de Paleobiología, Museo Nacional de Ciencias Naturales, CSIC, c/ Jose Gutierrez Abascal 2, 28006 Madrid, Spain f Department of Anthropology, Lehman College, CUNY, Bronx, NY, USA g Department of Vertebrate Paleontology and NYCEP, American Museum of Natural History, 79th Street and Central Park West, New York, NY 10024, USA article info abstract

Article history: We report dental remains of the extinct colobine monkey Mesopithecus from the Turolian (MN13, Late Received 25 March 2015 Miocene, ca. 6.23 Ma) locality of Venta del Moro (Valencia, Spain). They include most of the deciduous Accepted 11 August 2015 dentition and the unerupted germs of the first molars of a single infantile individual, as well as two lower Available online xxx left lateral incisors from two additional individuals. On the basis of morphometric comparisons, mainly based on the M1s, these remains are attributed to the Late Miocene species Mesopithecus pentelicus. They Keywords: represent a significant addition to the knowledge of the deciduous dentition of this taxon, much less Mesopithecus pentelicus well-known than the permanent dentition. Although this genus was widely distributed from the Late Venta del Moro Turolian Miocene through the Pliocene across Europe, southwestern Asia, Pakistan, and China, until now its Late Miocene occurence in the Late Miocene of the Iberian Peninsula had not been documented conclusively. Hence, Spain the reported remains considerably enlarge southwestwards the known geographic distribution of Mes- opithecus. The presence of this genus at Venta del Moro must be understood within the framework of the significant faunal turnover that took place in European faunas during the latest Turolian (the second Messinian mammalian dispersal), which is further documented at this locality by the occurrence of other eastern immigrants. At the same time, the presence of M. pentelicus at this site agrees well with previous paleoenvironmental and sedimentological evidence, indicating a lacustrine depositional environment with strong hydrologic seasonality. © 2015 Elsevier Ltd. All rights reserved.

1. Introduction Europe, southwestern and south Asia, and China (Delson, 1973, 1975; Szalay and Delson, 1979; Jablonski, 2002; Harrison and 1.1. The genus Mesopithecus Delson, 2007; Jablonski et al., 2011, 2014; Alba et al., 2014). Meso- pithecus is mostly known from Turolian and Pliocene sites, with a The extinct genus Mesopithecus Wagner, 1839 (, Cer- single doubtful Vallesian record (Andrews et al., 1996; Table 1). copithecoidea) includes small to medium-sized colobine monkeys Mesopithecus thus slightly postdates (or broadly coincides with) the distributed from the Late Miocene through the Pliocene across divergence date of the crown African and Asian colobine mono- phyletic clades (Wang et al., 2012), which is estimated at about 10.8e8.9 Ma (Sterner et al., 2006; Ting, 2008; Springer et al., 2012). A closer relationship with Asian colobines (tribe Presbytini) was * Corresponding author. E-mail addresses: [email protected] (D.M. Alba), [email protected] (P. Montoya), favored several decades ago on biogeographic grounds (Delson, [email protected] (M. Pina), lorenzo.rook@unifi.it (L. Rook), juan.abella@gmail. 1973, 1994; Szalay and Delson, 1979). More recently, morphology- com (J. Abella), [email protected] (J. Morales), [email protected]. based cladistic analyses (Jablonski, 1998; Byron, 2001) suggested edu (E. Delson). http://dx.doi.org/10.1016/j.jhevol.2015.08.003 0047-2484/© 2015 Elsevier Ltd. All rights reserved. 2 D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 that Mesopithecus is most closely related to the odd-nosed Asian 2002; Alba et al., 2014), we prefer to leave this genus as incertae colobine subclade, in particular to Pygathrix (Jablonski, 1998, 2002), sedis at the (sub)tribe rank. which is further supported by similarities in dental proportions Two species of Mesopithecus have been traditionally distin- (Pan et al., 2004). This was formalized by Groves (2000) with the guished (e.g., Delson, 1973, 1975, 1994; Szalay and Delson, 1979; inclusion of Mesopithecus into the Asian colobine tribe Rhinopi- Rook, 1999; Jablonski, 2002): the type species, Mesopithecus pen- thecini. However, the phylogenetic relationships of Mesopithecus telicus Wagner, 1839 (type locality: Pikermi, Greece), is question- are still unclear, and its unreduced thumb is consistent with this ably recorded in a single Vallesian locality (see note in Table 1) and taxon being a stem colobine preceding the divergence between definitively in numerous Turolian (MN11-MN13) sites from several Asian and African colobines (Jouffroy et al., 1991; Frost et al., 2015). European countries, Iran, and Afghanistan (see Table 1); and Mes- Therefore, following most previous work (Delson, 1973, 1975; opithecus monspessulanus (Gervais, 1849), with Montpellier Szalay and Delson, 1979; Strasser and Delson, 1987; Jablonski, (France) as its type locality is mainly recorded from the Pliocene

Table 1 List of European and southwestern Asian localities where Mesopithecus has been recorded, indicating its estimated age and the taxonomic attribution followed in this paper.

Locality Country Agea Taxonb

Wissberg Germany ?MN9 (11.1e9.7 Ma)c M. pentelicus cf. pentelicus Grebeniki 1 Ukraine MN11 (8.8e7.9 Ma) M. pentelicus cf. pentelicus Nikiti-2 Greece MN11 (8.8e7.9 Ma) M. pentelicus subsp. indet. Ravin des Zouaves-5 (type locality) Greece MN11 (ca. 8.2 Ma) M. pentelicus delsoni Baltavar Hungary MN12 (7.9e7.0 Ma) M. pentelicus cf. pentelicus Chomateres (¼Kisdari) Greece MN12 (7.9e7.0 Ma) M. pentelicus cf. pentelicus Hadjidimovo Bulgaria MN12 (7.9e7.0 Ma) M. pentelicus subsp. indet. Hatvan Hungary MN12 (7.9e7.0 Ma) M. pentelicus cf. pentelicus Kalimantsi Bulgaria MN12 (7.9e7.0 Ma) M. pentelicus subsp. indet. Kromidovo Bulgaria MN12 (7.9e7.0 Ma) M. pentelicus pentelicus Maragheh (middle beds) Iran MN12 (7.9e7.0 Ma) M. pentelicus pentelicus Molayan Afghanistan MN12 (7.9e7.0 Ma) M. pentelicus cf. pentelicus Perivolaki Greece MN12 (ca. 7.3e7.1 Ma) M. pentelicus subsp. indet. Pikermi (type locality) Greece MN12 (ca. 7.1 Ma) M. pentelicus pentelicus Vathylakkos-2 Greece MN12 (ca. 7.3 Ma) M. pentelicus subsp. indet. Vathylakkos-3 Greece MN12 (ca. 7.3 Ma) M. pentelicus subsp. indet. Gorna Sushitsa Bulgaria MN11eMN12 (8.8e7.0 Ma) M. pentelicus cf. pentelicus Ravin X Greece MN11eMN12 (8.8e7.0 Ma) M. pentelicus subsp. indet. Kryopigi Greece MN11eMN13 (7.9e5.1 Ma) Mesopithecus sp. indet. Kumanovo Macedonia (TFYRM) MN12eMN13 (7.9e5.1 Ma) M. pentelicus subsp. indet. Gravitelli Italy MN13 (7.0e5.1 Ma) Mesopithecus sp. indet. Brisighella (Monticino quarry) Italy MN13 (7.0e5.1 Ma) M. pentelicus cf. pentelicus Casino Italy MN13 (7.0e5.1 Ma) M. pentelicus cf. pentelicus Dytiko-1 Greece MN13 (7.0e5.1 Ma) M. pentelicus cf. pentelicus Dytiko-2 Greece MN13 (7.0e5.1 Ma) M. pentelicus cf. pentelicus and M. cf. monspessulanus Dytiko-3 Greece MN13 (7.0e5.1 Ma) M. pentelicus cf. pentelicus Locality indet. (Baccinello V3) Italy MN13 (7.0e5.1 Ma) Mesopithecus sp. indet. Maramena Greece MN13 (7.0e5.1 Ma) M. pentelicus pentelicus Moncucco Italy MN13 (5.4e5.33 Ma) M. pentelicus pentelicus Paganico (Baccinello V3) Italy MN13 (7.0e5.1 Ma) Mesopithecus sp. indet. Podere Firenze (Baccinello V3) Italy MN13 (7.0e5.1 Ma) Mesopithecus sp. indet. Venta del Moro Spain MN13 (ca. 6.23 Ma) M. pentelicus pentelicus Polgardi Hungary MN12eMN13 (7.9e5.1 Ma) M. pentelicus cf. pentelicus Veles Macedonia (TFYRM) MN12eMN13 (7.9e5.1 Ma)d M. pentelicus pentelicus Celleneuve France MN14 (5.1e4.2 Ma) M. monspessulanus Dorkovo Bulgaria MN14 (5.1e4.2 Ma) M. monspessulanus Montpellier (type locality) France MN14 (5.1e4.2 Ma) M. monspessulanus Capeni Romania MN15 (4.2e3.2 Ma) M. monspessulanus Ivanovce Slovakia MN15 (4.2e3.2 Ma) M. monspessulanus Malusteni Romania MN15 (4.2e3.2 Ma) M. monspessulanus Perpignan France MN15 (4.2e3.2 Ma) M. monspessulanus Wolfersheim€ Germany MN15 (4.2e3.2 Ma) M. monspessulanus Hajnacka Slovakia MN16 (3.2e2.6 Ma) M. monspessulanus Villafranca d'Asti (RDB Quarry) Italy MN16 (3.2e2.6 Ma) M. monspessulanus Red Crag England MN17 (ca. 2.3 Ma) M. monspessulanus

a Approximate age ranges based on MN units taken from Agustí et al. (2001: Fig. 2), updated based on Morales et al. (2013: Fig. 3) for the Miocene and Minwer-Barakat et al. (2012: Fig. 6) for the Pliocene. Most age estimates taken from MN attributions in Fortelius (2011), except for Greek localities (Koufos, 2009a, b), Grebenkiki 1 (Andrews et al., 1996; Eronen and Rook, 2004), Kumanovo (based on the local geology, see Dumurdzanov et al., 2004 since the locality is unpublished), Casino (Pradella and Rook, 2007), Hatvan (Bernor et al., 2003), Red Crag (Delson, 1974, 1994), Moncucco (Alba et al., 2014), and Venta del Moro (Gibert et al., 2013). b Most significant taxonomic references: Delson (1973, 1974, 1975, 1994), Szalay and Delson (1979), Heintz et al. (1981), de Bonis et al. (1990, 1997), Zapfe (1991), Kullmer (1991), Mottura and Ardito (1992), Kullmer and Doukas (1995), Andrews et al. (1996), Gentili et al. (1998), Rook (1999), Kohler€ et al. (1999), Koufos et al. (2003, 2004), Eronen and Rook (2004), Delson et al. (2005), Koufos (2006a, 2009a, b), Pradella and Rook (2007), Tsoukala and Bartsiokas (2008), Costeur and Malvesy (2010), Fortelius (2011), Rook and Alba (2012), Radovic et al. (2013), and Alba et al. (2014). c Wissberg has yielded a single isolated upper premolar identified as cf. Mesopithecus pentelicus (Delson, 1973 et. seq.; Andrews et al., 1996). The main faunal assemblage from this site is Vallesian (MN 9 or MN10), but there is a possibility that some Turolian fossils were mixed in. Given that it is an “outlier” in terms of the age range of this species, one must be hesitant to accept it without question. d In fact, the material comes from three sites (Brce, Prevalec, and Beluska) in the vicinity of Veles (formerly Titov Veles; Radovic et al., 2013). Although these authors attributed the material to both M. pentelicus and M. delsoni, the remains from these localities are here retained in M. p. pentelicus. D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 3

(MN14eMN17) of Europe (Table 1; see also Pradella and Rook, 2007), although it might have also coexisted with the preceding species during the late Turolian (MN13; see Discussion). A third European species of this genus, Mesopithecus delsoni de Bonis et al., 1990 (type locality: Ravin des Zouaves-5, Greece), has been recognized by several researchers (de Bonis et al., 1990, 1997; Koufos et al., 2003, 2004; Koufos, 2009a, b; Radovic et al., 2013), whereas others have merely considered it a junior subjective syn- onym of M. pentelicus (Zapfe, 1991; Delson, 1994; Andrews et al., 1996; Pradella and Rook, 2007). Following Alba et al. (2014), this taxon is here provisionally recognized at the subspecies rank, i.e., M. pentelicus delsoni de Bonis et al., 1990, for the reasons explained below (see Discussion). Another species, Mesopithecus sivalensis (Lydekker, 1878), is further recognized from the “Dhok Pathan” Figure 1. Location map of Venta del Moro. faunal assemblage (ca. 7.5e5.3 Ma) of the Siwaliks (Harrison and Delson, 2007). Finally, Mesopithecus has also been reported from the latest Miocene of China; it was initially attributed to a new (still (Montoya et al., 2006; see Marín, 2000 and Marín et al., 2002 for undescribed) species (Jablonski et al., 2011), but most recently additional taphonomic details). From a biochronological viewpoint, assigned to Mesopithecus cf. pentelicus (Ji et al., 2013; Jablonski Venta del Moro is correlated to MN13 (late Turolian, or Ventian, et al., 2014; the latter publication listed an unnamed new species sensu Morales et al., 2013). The age of this site was estimated until in the faunal list but not in the text). recently at 5.8e5.5 Ma, close to the Miocene-Pliocene boundary, on Here we describe and illustrate dental remains identified as the basis of biostratigraphic and magnetostratigraphic data Mesopithecus from the Turolian site of Venta del Moro (Valencia, (Opdyke et al., 1997; Montoya et al., 2002, 2006). However, a recent Spain), which has delivered an abundant assemblage of vertebrates, detailed magnetostratigraphic analysis allowed Gibert et al. (2013) invertebrates, and plants (Montoya et al., 2002, 2006). The presence to provide a revised age of 6.23 Ma, clearly preceding the Messinian of a colobine at this locality was first reported by Montoya Salinity Crisis. et al. (2002) without a formal taxonomic assignment. Subsequently, the remains were attributed to Mesopithecus sp. by Montoya et al. 2. Materials and methods (2006) and Marigo et al. (2014), and to M. pentelicus by Alba et al. (2013) in a preliminary report of the teeth described in this pa- Measurements of mesiodistal crown length (MD) and bucco- per. Here we identify the colobine from Venta del Moro as Meso- lingual crown breadth (BL) were employed to compare the pithecus pentelicus pentelicus. described material with other available samples of Mesopithecus.In the case of molars and deciduous premolars, BL measurements 1.2. Chronological and geological background were taken separately across the mesial and distal loph(id)s, although morphometric comparisons were based on maximum The site of Venta del Moro (UTM 642494, 4370967; latitude buccolingual breath. The breadth/length index (BLI), computed as 39.477 N, longitude 1.343 W) is located in an old railway trench BL(maximum)/MD 100, was employed to reflect the proportions close to the town of Venta del Moro (Cabriel Basin, Valencia, E of each tooth. Bivariate dental plots of BL versus MD were employed Iberian Peninsula; Fig. 1). From a geological viewpoint, it is situated to visually assess the size and proportions of the dental remains in the upper part of the Los Isidros Detrital Unit, within the Venta from Venta del Moro with regard to the various comparative del Moro-Villatoya Formation (Mathisen and Morales, 1981; Marín samples. Significant differences for the means of the several et al., 2002; Montoya et al., 2006). The section that outcrops at the measured variables were further tested in the case of the M1 by trench of Venta del Moro is composed of the following types of means of analysis of variance (ANOVA) and post-hoc pairwise deposits (Mathisen and Morales, 1981; Marín et al., 2002; Montoya comparisons (t-tests with p-values adjusted according to the SPSS et al., 2006): (a) lacustrine-palustrine deposits (limestones and implementation of the Bonferroni method to reflect the number of marls); (b) fluviatile deposits (sandstones and conglomerate pale- tests) using the SPSS 15.0 statistical package. Dental measurements ochannels); and (c) alluvial plain deposits, represented by an for the comparative samples were taken on original specimens by alternation of sandstones and red claystones. The lacustrine- the authors of this paper (mostly E.D.), complemented by data from palustrine deposits (Upper Carbonaceous Unit) have an areal the literature (de Bonis et al., 1990, 1997; Zapfe, 1991; Mottura and extension of about 5 1.5 km at Venta del Moro, corresponding to a Ardito, 1992; Kullmer and Doukas, 1995, after; Kullmer, 1991; shallow lacustrine depositional environment that was subject to Koufos et al., 2004; Delson et al., 2005; Koufos, 2006a). Many of fluctuating water table conditions and anoxic episodes (Montoya these data can be accessed online via PRIMO (http://primo.nycep. et al., 2006). Such a facies succession shows several cycles of org), the NYCEP PRImate Morphometrics Online database. inundation-stability-desiccation, and the site is situated in horizons of lignitic marls, marls, and lutitic limestones within the last 3. Systematic paleontology lacustrine cycle recognized in the series (Montoya et al., 2006). Several fossiliferous layers rich in vertebrate remains have been Order Primates Linnaeus, 1758 identified (see Montoya et al. [2006] for further stratigraphic de- Infraorder Catarrhini E. Geoffroy Saint-Hilaire, 1812 tails). The faunal list from Venta del Moro includes ostracods, Superfamily Cercopithecoidea Gray, 1821 mollusks, fishes, reptiles, birds, and 43 mammalian taxa (Aguirre Family Cercopithecidae Gray, 1821 et al., 1973; Morales, 1984; Montoya et al., 2002, 2006, 2009, Subfamily Colobinae Blyth, 1863 2011). The macrovertebrate assemblage is mainly composed of Genus Mesopithecus Wagner, 1839 isolated teeth (such as those described in this paper), implying that Mesopithecus pentelicus Wagner, 1839 the remains were exposed under subaerial conditions for a variable Mesopithecus pentelicus pentelicus Wagner, 1839 period of time until water table fluctuations led to their burial (Fig. 2) 4 D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14

3.1. Described specimens and measurements

The dental fossil material described (Table 2) is currently housed at the Museu de Geologia of the Universitat de Valencia (MGUV) in Spain. Metrical data on the material are given in Table 3.

3.2. Preservation

After the initial discovery of an isolated deciduous premolar during manual excavation of the site, additional dental remains were recovered by screen-washing sediments, mainly from the same stratigraphic level. On the basis of the degree of wear and the lack of repeated elements, most of the dental remains are attrib- utable to a single individual. These dental elements in most cases only preserve the crowns and, in a few instances, part of the roots. The upper and lower first molar crowns correspond to unerupted germs that had almost completed formation, indicating that this individual was an infant, as is also suggested by the light wear on the deciduous premolars. Two lower left I2s (MGUV-19622 and MGUV-24140), preserving only slightly worn crowns and partial roots, were also recovered. Compared to other catarrhines, colobines display a tendency to- wards early molar eruption relative to the anterior teeth (Harvati, 2000; Harvati and Frost, 2005, 2007), with the M1 and even the M2 erupting before the I2 in M. pentelicus pentelicus (Harvati and Frost, 2005, 2007). Given that unworn dI2s are present in the very young individual from Venta del Moro, and that its M1swere unerupted and not completely formed, the attribution of either of Figure 2. Dental remains of Mesopithecus from Venta del Moro. A) Left dI1 MGUV- the two available I2s (which show some apical wear and well- 15778 in occlusal (A1), mesial (A2), labial (A3), distal (A4), and lingual (A5) views; B) Left dC1 MGUV-12758 in occlusal (B1), mesial (B2), buccal (B3), distal (B4), and formed partial roots) to the same infantile individual is not lingual (B5) views; C) Right dP3 MGUV-12755 in occlusal (C1), mesial (C2), buccal reasonable. It is therefore concluded that the two left I2s corre- (C3), distal (C4), and lingual (C5) views; D) Right dP4 MGUV-12756 in occlusal (D1), spond to two different, (sub)adult individuals, resulting in a mini- 1 mesial (D2), buccal (D3), distal (D4), and lingual (D5) views; E) Right M germ mum number of three individuals represented within the Venta del MGUV-12757 in occlusal (E1), mesial (E2), buccal (E3), distal (E4), and lingual (E5) Moro assemblage. views; F) Labial crown fragment of left dI1 MGUV-12751 in labial view; G) Labial crown fragment of right dI2 MGUV-12751 in labial view; H) Left dI2 MGUV-12759 in occlusal (H1), mesial (H2), labial (H3), distal (H4), and lingual (H5) views; I) Left dC1 3.3. Description MGUV12760 in occlusal (I1), mesial (I2), buccal (I3), distal (I4), and lingual (I5) views; J) Left dP MGUV-12752 in occlusal (J1), mesial (J2), buccal (J3), distal (J4), and 3 The dental morphology is typical of that seen in most colo- lingual (J5) views; K) Left dP4 MGUV-12753 in occlusal (K1), mesial (K2), buccal (K3), distal (K4), and lingual (K5) views; L) Right dP4 MGUV-12761 in occlusal (L1), mesial bines, extant or extinct, as described by Delson (1973) and Szalay (L2), buccal (L3), distal (L4), and lingual (L5) views; M) Mesial fragment of right M1 and Delson (1979). However, details of colobine deciduous denti- germ MGUV-14811 in occlusal (M1), mesial (M2), buccal (M3), distal (M4), and tion, much less that of Mesopithecus, are not well known, and thus lingual (M5) views; N) Left M MGUV-12754 in occlusal (N1), mesial (N2), buccal 1 we provide more detail here than might otherwise be expected. (N3), distal (N4), and lingual (N5) views; O) Left I2 MGUV-19622 in occlusal (O1), mesial (O2), labial (O3), distal (O4), and lingual (O5) views; P) Left I2 MGUV-24140 in The permanent teeth require less descriptive detail. The diagnostic occlusal (P1), mesial (P2), labial (P3), distal (P4), and lingual (P5) views. In occlusal features of Mesopithecus do not relate to its dentition, but rather to views, mesial is depicted to the left. its cranium and postcranium. Its dentition is typically colobine,

Table 2 List of Mesopithecus pentelicus pentelicus specimens from Venta del Moro.

Collection No.a Anatomical description Figure

MGUV-12751 Dental fragments, including buccal crown fragments of L dI1 and R dI2 Fig. 2FeG MGUV-12752 L dP3 crown Fig. 2J MGUV-12753 L dP4 crown Fig. 2K MGUV-12754 L M1 germ Fig. 2N MGUV-12755 R dP3 crown Fig. 2C MGUV-12756 R dP4 crown Fig. 2D MGUV-12757 R M1 germ Fig. 2E MGUV-12758 L dC1 crown with basalmost portion of the root Fig. 2B

MGUV-12759 L dI2 crown Fig. 2H MGUV-12760 L dC1 crown Fig. 2I MGUV-12761 R dP4 crown with partial roots Fig. 2L MGUV-14811 R M1 germ mesial fragment Fig. 2M MGUV-15778 L dI1 crown and partial root Fig. 2A

MGUV-19622 L I2 crown with root Fig. 2O MGUV-24140 L I2 crown with partial root Fig. 2P a MGUV ¼ Museum of Geology, Universitat de Valencia. D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 5

Table 3 midcrown length. Two long and sharp crests descend from the apex Dental measurements (in mm) and ratios of Mesopithecus pentelicus pentelicus from of the main cusp, the distal one being longer than the mesial one. At Venta del Moro.a the mesial and distal ends of the crown, these crests merge with the Collection No.b Tooth MD BLm BLd BLI distinct and somewhat irregular lingual cingulum, which is situated

MGUV-12572 dP3 6.3 3.2 3.8 60.32 at about one-third of crown height above the cementoenamel MGUV-12753 dP4 6.6 4.5 5.1 77.27 junction. At the distal crown end, where it joins the distal crest, MGUV-12754 M1 7.4 (5.4) (5.8) (78.38) the cingulum terminates in a tubercule-like small swelling. A 3 MGUV-12755 dP 5.6 4.6 4.8 85.71 moderately-developed lingual swelling of the crown base is pre- MGUV-12756 dP4 6.5 6.0 6.1 93.85 MGUV-12757 M1 7.4 (6.3) (5.9) (85.14) sent at about midcrown length, where the cingulum displays a V- MGUV-12758 dC1 5.1 3.7 72.55 shape. From this point, a poorly-defined, short, and vertical crest MGUV-12759 dI2 3.8 2.8 73.68 originates, thus separating two concave lingual portions of the MGUV-12760 dC1 4.8 3.3 68.75 crown, although it does not reach the crown apex. MGUV-12761 dP 6.6 4.6 5.2 78.79 4 The dP3 (MGUV-12755, Fig. 2C) crown only shows a slight de- MGUV-14811 M1 e (5.5) e MGUV-15778 dI1 4.4 2.9 65.91 gree of wear on the tips of the four main cusps, so that its occlusal

MGUV-19622 I2 3.6 3.0 83.33 anatomy may be described in detail. It is an elongated tooth (much MGUV-24140 I2 3.3 3.1 93.94 longer than wide), with a markedly asymmetrical, trapezoidal a BLd ¼ buccolingual breadth on the distal lobe; BLm ¼ buccolingual (or labio- occlusal outline. Thus, the crown is longer on the buccal than on the lingual) breadth, either maximum or, in deciduous premolars and permanent mo- lingual side, with a markedly inclined mesial marginal ridge, and ¼ ¼ lars on the mesial lobe; MD mesiodistal length; BLI BL(maximum)/MD 100. the distal lobe is much wider than the mesial one. The buccal cusps Values within parentheses denote that measurements are estimates due to incomplete preservation. are much more peripheral than the corresponding lingual cusps, b MGUV ¼ Museum of Geology, Universitat de Valencia. and the mesial ones are taller than the distal ones. The lingual surface of the tooth flares extensively, so that the lingual occlusal margin between protocone and hypocone is just slightly lingual to the buccolingual midline of the crown, as is typical of cercopithecid with no clearly diagnostic aspect. Mesopithecus is the only colo- dP3s. All the cusps are quite buccolingually compressed. The bine of this size known in Europe (Dolichopithecus is significantly paraloph is thin and slightly curved, but quite transversely aligned, larger but also has no generically diagnostic dental features), so whereas the metaloph is more obliquely oriented. The paraloph, that identification of the Venta del Moro sample as belonging to together with the mesially-directed preprotocrista, the longer but Mesopithecus is reasonable and parsimonious, but not definitive. similarly-oriented preparacrista, and the obliquely-oriented mesial Conceivably, these teeth could belong to an as yet unknown small marginal ridge, delimit a well-developed, asymmetrical, mesial species of Dolichopithecus or even to Libypithecus, known from fovea, which is slightly broader than long. The central fovea (trigon contemporaneous sites in North Africa. One option would be to basin) is more extensive and deeper than the mesial fovea, further indicate the small degree of hesitancy we have with regard to the displaying a rectangular outline (somewhat longer than broad). identification of this sample by formally referring to it as cf. There is a secondary crest that runs from the apex of the metacone M. pentelicus pentelicus. This hesitancy would thus need to be towards the center of the central fovea, which shows a moderate extended to all western European dental-only samples, such as development of secondary enamel crenulations. The more those in the Miocene of Italy, Hungary, and Greece. Rather than restricted and shallow distal fovea (talon basin), which is clearly revising the nomenclature for all these populations (and by broader than long, is mostly restricted to the buccal moiety of the extension for all fragmentary samples of Cenozoic teeth), crown, being distally closed by a moderately-developed and curved we think it is simpler to accept the most likely identification as distal marginal ridge. There is no distinct buccal cingulum, whereas Mesopithecus and then determine the infrageneric allocation on a poorly-developed cingulum is present on the mesiolingual the basis of size and morphology, as we do below. portion of the crown wall. There is a moderately-developed lingual 3.3.1. Upper deciduous teeth All the deciduous dentition is rep- cleft just above the cingulum. resented in the sample, except for the dI2. The dI1 (MGUV-15778, Like the preceding tooth, the dP4 (MGUV-12756, Fig. 2D) only Fig. 2A), which merely shows a slight degree of wear along the shows a minimal degree of wear (only at the tip of the two mesial apical crown margin, is a slightly spatulate, waisted, and cusps), although a large flake of enamel is missing from the lingual relatively short-crowned tooth. In occlusal view, the crown surface of the protocone. This tooth similarly displays a bilopho- displays an elliptical profile, being mesiodistally longer than dont occlusal pattern with quite buccolingually compressed cusps, labiolingually wide. In mesial/distal views, the labial profile is the buccal ones being much more peripheral than the lingual cusps. straight to slightly convex, whereas the lingual one displays a Nevertheless, the dP4 displays numerous differences compared to more convex contour. In labial/lingual views, the crown is the dP3: the former is larger and relatively broader (although still somewhat asymmetrical, with a straighter and taller mesial being longer than broad) and much more symmetrical (the mesial contour and a somewhat shorter, more convex and obliquely lobe being as broad as the distal one), the lingual flare is not as oriented distal profile. There is a narrow but marked lingual extensive, the preparacrista is not longer than the preprotocrista cingulum, situated at about one-third of crown height above the and the mesial marginal ridge is not obliquely oriented (resulting in cementoenamel junction. This cingulum displays a moderately- a subrectangular mesial fovea, clearly broader than long), the distal developed, triangular swelling on the medial portion of the cusps are almost as tall as the mesial ones, the central fovea shows crown that approximately reaches crown midheight but which is no development of secondary enamel crenulations, the distal fovea not further continued apically along the lingual crown wall. is more extensive and centrally situated, there is a buccal cleft The dC1 (MGUV-12758, Fig. 2B) is a moderately high and (although short and poorly-developed), and the lingual cleft is pointed tooth that displays a single main cusp. The crown is buc- more well-developed; and finally, the lingual cingulum is broader colingually compressed, with a suboval occlusal out- and more marked, although being similarly restricted to the linedmesiodistally longer than buccolingually broad, and mesiolingual portion of the crown. somewhat broader at the distal than at the mesial portion of the 3.3.2. Lower deciduous teeth The dI1 is only represented by the crown. The main cusp is situated slightly mesially relative to labial crown portion (MGUV-12751, Fig. 2F), so that few details of its 6 D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 morphology can be ascertained. It is a relatively low and hypoconid and entoconid, respectively situated at the distobuccal mesiodistally short, spatulate, and quite symmetrical tooth, with and distolingual corners of the crown. Two thin cristids (pre- its apical crown margin being slightly inclined downwards from hypocristid and preentocristid) run mesially from these cuspids to mesial to distal and displaying several moderately-distinct the bases of the protoconid and metaconid, respectively. The two mamelons. The dI2 is represented by the complete crown of the distal cuspids are linked to each other only by the marked and left side (MGUV-12759, Fig. 2H) and also by the labial portion of curved distal marginal ridge, which distally closes the central fovea the right (MGUV-12751, Fig. 2G). The dI2 differs from the dI1 by (talonid basin). The latter is similar in size to the mesial fovea, but being pointed and more asymmetric. Its crown displays a suboval much deeper and subquadrangular (being about as broad as long). contour, being mesiodistally longer than labiolingually broad, and There is no transverse cristid homologous to the hypolophid con- broader at the distal than at the mesial portion of the crown. In necting the two distal cuspids, so that no distal fovea can be mesial/distal views, the dI2 crown displays a uniformly and distinguished from the central one. At most, a short, transverse moderately convex profile, whereas the lingual crown wall is hypoentocristid might have been originally present but subse- markedly convex except at the base, which is inflated and quently eroded by wear. On the buccal crown wall, there is a poorly- protrudes lingually. In labial/lingual views, this tooth is much developed vertical groove that does not reach the crown base, higher mesially rather than distally, with the point of maximal between the paraconid and the protoconid, as well as a much wider protrusion being slightly mesially displaced relative to crown and deeper buccal cleft between the protoconid and hypoconid that midlength. This point cannot be considered a cuspid in a strict also does not reach the crown base but terminates in a pocket. sense, but rather the most protruding of the three mamelons that Lingually, the medial lingual notch between metaconid and ento- can be distinguished along the course of the mesial portion of the conid is fairly deep, as is typical of colobine lower molariform teeth. apical ridge, which is moderately convex and obliquely inclined. Both the right (MGUV-12761, Fig. 2L) and the left (MGUV-12753, The distal portion of the apical margin is on the contrary much Fig. 2K) dP4 are preserved, the former further preserving part of the more steeply inclined, and on the distal crown end it further roots socketed within a portion of the mandibular bone. These shows a distinct cuspulid, which reaches approximately teeth only show a slight degree of wear at the tips of the two mesial midcrown height. This “distocuspulid” is separated from the cuspids and at the mesial marginal ridge. The dP4 is somewhat remaining portion of the lingual crown aspect by a distinct but longer and much broader than the dP3, further displaying a sub- short vertical groove. Both the mesial marginal ridge and the rectangular instead of subtriangular outline, although similarly somewhat shorter but more markedly convex distal marginal being wider across the distal than the mesial lobe. Unlike the ridge are rather blunt and indistinct, just like the curved lingual preceding deciduous premolar, the dP4 displays a complete bilo- cingulid that runs close to the crown base from the distal end of phodont occlusal pattern, with four main cuspids (the lingual ones the latter up until about one-third of crown length before more peripheral than the buccal) transversely arranged and united reaching the mesial end. by well-developed lophids, the hypolophid being somewhat longer The dC1 is represented by a single and partially broken crown than the protolophid because the two mesial cuspids are closer to (MGUV-12760, Fig. 2I), although most of its morphology can be one another. The marked and sharp preprotocristid and pre- readily ascertained. This tooth somewhat resembles the dI2, but the metacristid curve in a mesial direction until merging with the crown is higher and more buccolingually compressed, further dis- similarly-developed mesial marginal ridge. Together with the playing a distinct main cuspulid clearly situated on the mesial protolophid, these cristids delimit an extensive and deep, semi- moiety of the crown, a much lower secondary cuspulid situated on circular mesial fovea, which is approximately as broad as long. This the distal end of the crown, and a lingual cingulid that (even though fovea is partly occupied by a third, supplementary cristid that runs blunt and poorly-developed) extends until the mesial end of the from the apex of the metaconid and markedly curves until it ends at crown. There, the cingulid merges with the straight and moderately the center of the fovea. The central fovea is even deeper and more sharp mesial cristid that descends from the main cuspid until about extensive than the mesial one, further being subrectangular in crown midheight. Another blunter and more poorly-defined distal shape (slightly longer than broad), although somewhat asymmet- cristid runs in the opposite direction, until it reaches the distal rical due to the oblique direction of the prehypocristid compared to cuspulid, which is apically worn against the dC1. The buccal crown the mesiodistal crown axis. The postprotocristid and post- wall is markedly convex, whereas the lingual one is somewhat metacristid are shorter than the corresponding prehypocristid and concave, particularly along its distal two-thirds. preentocristid. The distal cuspids are only slightly less tall than the The dP3 (MGUV-12572, Fig. 2J) shows a slight degree of wear, mesial ones, and besides being united by the marked and convex with some dentine exposure at the apices of most cuspids. The distal marginal ridge, they are connected by a distinct hypolophid. crown is relatively narrow (much longer than broad) and displays a The latter separates the central fovea from the shallower and more subtriangular oclusal outline, with the distal lobe being somewhat restricted distal fovea, which is broader than long. As in the dP3, broader than the mesial one. The trigonid is much higher than the there is a shallow vertical groove on the buccal crown wall close to talonid, with the very buccolingually-compressed protoconid being its mesial end, although in the dP4 it is less well-developed and the highest cusp of the former. A long preprotocristid runs in a situated farther away from the crown base. On the contrary, the mesial direction until it merges with the rather rudimentary par- buccal cleft between the protoconid and hypoconid is broader and aconid, situated at the mesial end of the crown. From the distal deeper than in the dP3, although no ectostylid is present. The me- aspect of the protoconid, two crests originate: a distally-directed dian lingual notch is again deep. postprotocristid (shorter than the preprotocristid) and a 3.3.3. Permanent dentition Only five permanent colobine teeth transversely-aligned protolophid. The latter connects the proto- are known from Venta del Moro: the two lower second incisors conid to the smaller, less protruding, and less buccolingually- mentioned above and the germs of upper and lower first molars. compressed metaconid that gives rise to weak pre- and post- The right M1 unerupted germ (MGUV-12757, Fig. 2E) preserves its metacristids that form the lingual marginal ridge. The pre- occlusal morphology, except that the basalmost portion of the protocristid, protolophid, and premetacristid delimit a sub- crown had not yet been formed, so that breadth measurements triangular mesial fovea (trigonid basin), which is about twice as on this particular specimen should be taken only as minimum long as it is broad. The talonid is not only lower, but also shorter estimates. Although larger, the crown displays an occlusal outline than the trigonid; it displays two distinct low cuspids, the and a general occlusal morphology reminiscent of the dP4, D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 7 including the arrangement and development of the cusps (the deciduous premolars and the permanent molars, by the deeply mesial ones being taller than the distal and the paracone being excavated median lingual notch of lower molariform teeth, by the the largest one), although they are more buccolingually reduced trigonid and the markedly greater breadth of the distal 4 compressed than in the dP . These two teeth also resemble each portion of the crown compared to the mesial one in the M1 (and other in the development of the foveae. However, the lingual dP4), by the lack of a distinct distal fovea on the dP3, and by the faces of the protocone and hypocone show very little flare, so distal “prong” of the dI2 (also characteristic of the colobine I2; that the basins are far more extensive lingually than in either Delson, 1973). deciduous tooth; this pattern is typical of all cercopithecid upper Although the dental morphology of the Venta del Moro sample molars. The presence of cingula and the extent of the clefts fits well with previously recovered remains of Mesopithecus from cannot be adequately ascertained due to incomplete crown several sites (Table 1), extensive comparisons are hampered by the formation, although a wide and deep lingual cleft, at least as lack of deciduous teeth in most of these samples. Thus, the genus developed as in the dP4, was present. Mesopithecus is best known on the basis of the sample of fossils The M1 is represented by a left complete unerupted germ from the type locality of M. pentelicus, where as many as 60 in- (MGUV-12754, Fig. 2N) and a right mesial germ fragment (MGUV- dividuals including 22 crania have been recovered (Zapfe, 1991; 14811, Fig. 2M). In both instances, the crown is not completely Costeur and Malvesy, 2010). Not all the deciduous tooth loci are formed, so that breadth measurements again must be considered represented among the Pikermi sample (Delson, 1973; Zapfe, 1991). minimum estimates. The basic occlusal morphology resembles that In fact, the deciduous dentition of M. pentelicus is best known from of the dP4. There are, however, some noticeable differences; thus, in the Greek locality of Maramena (Kullmer, 1991; Kullmer and the M1 the trigonid is markedly higher than the talonid and at the Doukas, 1995). Additional deciduous material is available from same time more clearly reduced in area, with a less extensive Vathylakkos 2 (Koufos et al., 2004), Baccinello V3 (Delson, 1973; mesial fovea that is clearly broader than long. Moreover, the met- Rook, 1999), and Gorna Sushitsa (E.D., pers. obs.: subadult maxilla aconid is more mesially situated relative to the protoconid FM1530, erroneously described as from Kromidovo by Koufos et al. (resulting in an obliquely inclined protolophid) and the crown is [2003] is labelled to be from Gorna Sushitsa, which was confirmed relatively broader than that of the dP4. Whereas the short and by N. Spassov, pers. comm. to E.D.). Plots of dental proportions in curved premetacristid directly merges with the mesial marginal both the deciduous and permanent teeth from Venta del Moro ridge, the preprotocristid is separated from the latter by a shallow compared to most other available Mesopithecus teeth are reported and short vertical groove and a concomitant stylid (better-defined in Figures 3e5 (summarized in Table 4). than in the dP4) that curves along the mesiobuccal crown wall from Both the lower (Fig. 3) and upper (Fig. 4) deciduous teeth fit the mesiobuccal end of the crown toward its base. There is a deep within the range of dimensions of M. pentelicus sensu stricto (s.s.) and broad buccal cleft, the extent of which cannot be completely from Pikermi and Maramena, but detailed statistical comparisons ascertained due to incomplete crown formation, and a deep median are precluded by both small sample sizes and the lack of ho- lingual notch. mologous material attributable to M. monspessulanus.Withre- All of the teeth described above were probably derived from a gard to the permanent dentition (Fig. 5), the two available I2 from single young juvenile or infant individual, while the two remaining Venta del Moro, similar to those available for M. monspessulanus, lower teeth appear to be the sole representatives of two different tend to be narrower than most of those known for M. pentelicus (sub)adults. The left I2 is represented by two nearly identical (although some specimens from Pikermi display similar di- specimens (MGUV-19622 and MGUV-24140, Fig. 2OeP), both of mensions and proportions). With regard to M1, the unerupted which are only slightly worn, with minimal dentine exposure on germ from Venta del Moro falls within the scatter of points of the apical margin. Both teeth are spatulate and non-waisted, with a M. pentelicus. Although the completed crown might have been slightly asymmetrical crown that is broadest at the base and slightly broader, thus falling well within the variability of somewhat tilted to mesial and a markedly convex apical ridge. M. pentelicus, the recorded measurements already indicate much There is enamel on both the labial and lingual sides of the crown. In larger occlusal dimensions than the single available specimen of mesial view, the cementoenamel junction displays an inverted-V M. monspessulanus, thus suggesting that an attribution to the shape, whereas on the distal side it is situated in a lower position latter species would not be warranted. This is further confirmed towards the root apex. The labial crown wall displays a convex by the dimensions of the M1 from Venta del Moro that are clearly contour, whereas the lingual one is slightly concave to flat. The larger than those of M. monspessulanus but fall well within the latter displays a smooth surface, except for a vertical pillar that is variability of M. pentelicus (see below). situated on the basal portion of the crown. In this respect, the two The M1 sample available for M. monspessulanus is large enough available specimens show some variation: in MGUV-19622, the to permit statistical comparisons between the two species; the crown base is more swollen lingually and the pillar is broader and following groups were distinguished (see descriptive statistics shorter, being separated from the marked distal marginal ridge by a and further details in Table 4): M. p. pentelicus from Pikermi, M. p. narrow vertical groove; in MGUV-24140, on the contrary, the basal pentelicus from Maramena, M. p. pentelicus from other localities, portion of the pillar is subdivided into three distinct folds of M. pentelicus subsp. indet., and M. monspessulanus (including M. cf. enamel, one of which is longer and continues until about midcrown monspessulanus). ANOVA results indicate that there are significant height, terminating close to the distal marginal ridge. Both speci- differences for the three investigated variables at p < 0.001 mens display a similarly marked mesial marginal ridge but no sign (F ¼ 9.547 for length, F ¼ 15.156 for maximum breadth, and of a lingual cingulid. F ¼ 6.082 for breadth/length index). Post-hoc pairwise compari- sons (see Methods) indicate that M. monspessulanus significantly 3.4. Morphological and metrical comparisons differs (at least at p < 0.05) from all the samples attributed to M. pentelicus in both length and width of the M1,withthesingle The general morphology of the dental sample from Venta del exception of length of M. monspessulanus compared to Moro corresponds to the typically colobine pattern displayed by M. pentelicus subsp. indet. It can thus be concluded that Mesopithecus (Delson, 1973; Szalay and Delson, 1979; Zapfe, 1991). M. monspessulanus generally differs from M. pentelicus by dis- This is indicated, among other features, by the marked crown relief playing an absolutely shorter and narrower M1.Significant dif- with somewhat buccolingually compressed cusps in both the ferences (at p < 0.01) are found in M1 proportions between 8 D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14

Figure 3. Bivariate dental plots of maximum breadth (BL) versus length (MD), in mm, for the lower deciduous teeth of Mesopithecus pentelicus pentelicus from Venta del Moro compared to Mesopithecus spp. from other European and southwestern Asian localities.

M. monspessulanus and the topotypic sample of M. p. pentelicus 4. Discussion from Pikermi, with the former displaying on average a relatively narrower M1, although their ranges largely overlap and such dif- 4.1. Taxonomic attribution ferences are not substantiated by the remaining samples of M. pentelicus. The sample from Maramena further differs at The taxonomic distinction among the various purported Euro- p < 0.001 from the Pikermi sample regarding mesiodistal length, pean species of Mesopithecus is unclear. Resolving the alpha- although differing from the latter in the opposite direction than taxonomy of Mesopithecus is outside the scope of this paper, but M. monspessulanus (i.e., displaying longer instead of shorter M1), some considerations are required regarding the validity of the so that an attribution to the younger species is not warranted on various taxa. Koufos et al. (2003; see also Koufos, 2006a) adopted the basis of the performed statistical tests. Compared to these the view that the various recorded populations represent a single samples (Table 4), the Venta del Moro unerupted germ of M1 lineage that mostly varies in size, and they favored recognizing overlaps in size with the maximumeminimum ranges of most of three distinct species with intermediates between them, while the M. pentelicus samplesdthe only exception being the Mar- accepting that other solutions (a single variable species, amena remains that display a somewhat longer M1, although this M. pentelicus, or three distinct subspecies within the latter) might might be merely attributable to small sample size. On the con- also be acceptable from a taxonomic viewpoint. The distinction of trary, the M1 germ from Venta del Moro shows larger dimensions chronospecies in the fossil record is of course warranted. However, in both length and breadth than the maximum values recorded for the existence of numerous intermediate forms between M. monspessulanus. On the basis of dental size, we conclude that if M. pentelicus s.s. and the purported distinct species M. delsoni (such it is Mesopithecus, the Venta del Moro sample is best attributed to as those from Vathylakkos 2; Koufos et al., 2004), as well as the M. p. pentelicus. coexistence in the same locality of typical M. pentelicus with D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 9

Figure 4. Bivariate dental plots of maximum breadth (BL) versus length (MD), in mm, for the upper deciduous teeth of Mesopithecus pentelicus pentelicus from Venta del Moro compared to Mesopithecus spp. from other European and southwestern Asian localities. Mesopithecus p. pentelicus (other localities) also includes some material attributed to M. p. cf. pentelicus (see Table 1). intermediate forms (e.g., Kalimantsi; Koufos et al., 2003), makes it chronospecies (Delson et al., 2005). On this basis, together with the preferable, in our opinion, to provisionally distinguish these two more clear-cut morphological criteria previously considered to taxa only at the subspecies level (i.e., M. p. pentelicus and distinguish the two species (see below), M. monspessulanus is here M. pentelicus delsoni), at least until a thorough revision of the genus considered a different species from M. pentelicus. The possible is undertaken. The subspecies rank is useful to distinguish different coexistence of these two species during MN13, however, compli- morphs that can be related to differences in geography and/or cates the taxonomic attribution of the Venta del Moro remains, chronology, but for which intermediate forms are expected to exist. given the limited number of permanent molars available and the Such intermediate forms, previously referred to as M. pentelicus/ fact that the deciduous dentition of M. monspessulanus is unknown. delsoni by Koufos (2009a,b), together with others previously Previous research has suggested that differences between labelled as M. aff. delsoni and M. cf. delsoni, are here merely referred M. pentelicus sensu lato (s.l.) and M. monspessulanus are less to M. pentelicus subsp. indet. On the contrary, the possibility that ambiguous than those claimed to exist between M. p. pentelicus and both M. pentelicus and M. monspessulanus are recorded at some M. p. delsoni (contra Koufos, 2006a). M. pentelicus can be distin- localitiesdespecialy Dytiko-2 in Greece (de Bonis et al., 1990, 1997; guished from M. monspessulanus by its larger size (best consensus Andrews et al., 1996; Koufos et al., 2004; Delson et al., 2005) and estimated body masses of 13 kg for males and 9 kg for females of perhaps also in Baccinello V3 localities (see Rook, 1999)dsuggests M. pentelicus, compared to 10 kg for males and 7 kg for females of that the latter species might have coexisted for a short time with its M. monspessulanus; Delson et al., 2000), the somewhat larger and purported ancestor, M. pentelicus. According to this view, relatively broader lower molars, the more marked molar flare, and M. monspessulanus might have originated near the end of the the greater retroflexion of the humeral medial epicondyle (indi- Miocene from one or several small populations of M. pentelicus that cating a greater degree of terrestriality in the former; Delson, 1973, became adapted to a more arboreal niche (Delson, 1973; Andrews 1975; Szalay and Delson, 1979; de Bonis et al., 1990; Ciochon, 1993; et al., 1996), coexisting for some time with its ancestral species Andrews et al., 1996). M. p. delsoni purportedly differs from M. p. and thus being best regarded as biospecies instead of merely as pentelicus by the larger size of the former, together with 10 D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14

Figure 5. Bivariate dental plots of maximum breadth (BL) versus length (MD), in mm, for the permanent teeth of Mesopithecus pentelicus pentelicus from Venta del Moro compared to Mesopithecus spp. from other European and southwestern Asian localities. Mesopithecus p. pentelicus (other localities) also includes some material attributed to M. p. cf. pentelicus (see Table 1).

mandibular shape differences, the large P3 honing facet, and the MN13/MN14 boundary), so that an attribution to weak and rounded metalophid and well-developed talonid with a M. monspessulanus could not be clearly discounted on chronological large hypoconulid of the M3 (de Bonis et al., 1990; Koufos et al., grounds. There are further taxonomic uncertainties regarding the 2003, 2004; but Andrews et al., 1996 rejected some of these sample from Vathylakkos 2, previously attributed to M. pentelicus claims). None of the features mentioned above refer to the decid- by some (e.g., Pradella and Rook, 2007), but later reassigned by uous dentition, so that the taxonomic assignment followed in this Koufos and colleagues (e.g., Koufos, 2009a, b) to a form interme- paper mostly relies on metrical comparisons of the first molars. diate between M. pentelicus and the purported different species The large sample of teeth available from Pikermi (Zapfe, 1991), M. delsoni: M. cf. pentelicus according to de Bonis et al. (1997), M. aff. the type locality of M. pentelicus s.s., allows us to adequately eval- pentelicus according to Koufos et al. (2004),orM. delsoni/pentelicus uate whether other samples fit well with this species. This is in Koufos (2009a, b); here it is simply termed M. pentelicus subsp. however only true for the permanent dentition, because not all the indet. The same putatively intermediate morph could be present in deciduous dentition is known from Pikermi (i.e., the anterior de- the localities of Kalimantsi, Hadjidimovo, and Perivolaki (Koufos, ciduous teeth are missing; see Zapfe, 1991) and sample sizes for the 2009a), having been previously attributed to M. aff. delsoni by available deciduous teeth are much smaller. Unfortunately, the Koufos et al. (2003) or M. cf. delsoni by Koufos (2006a). Finally, the attribution at the species level of the more complete sample of single deciduous premolar from Baccinello V3 was attributed to deciduous teeth from Maramena is open to different in- Mesopithecus sp. by Rook (1999), who nevertheless noted that this terpretations, having been previously attributed to either specimen did not differ from the Maramena sample. Most recently, M. pentelicus (Kullmer and Doukas, 1995; Delson et al., 2005)or M. p. pentelicus has been reported from the latest Miocene of Mesopithecus sp. (Koufos, 2009a). This is further complicated by Moncucco Torinese in Italy (Alba et al., 2014), but the sample from some degree of uncertainty in the age of Maramena. Biostrati- this locality includes no teeth homologous with those from Venta graphic considerations based on small merely suggest an del Moro. age close to the Turolian/Ruscinian boundary (Schmidt-Kittler et al., Our statistical comparisons of M1 confirm the previous results of 1995), but an age closer to the latest Turolian is favored on the basis Andrews et al. (1996), according to which this tooth is significantly of both macromammals and the lack of Promimomys (Koufos, smaller in M. monspessulanus compared to M. pentelicus s.s. Our 2006b, 2009a). Overall, we concur with the latter author that results further indicate that, at least on the basis of M1 crown di- Maramena is likely latest Turolian (even latest MN13, close to the mensions, both the sample from Maramena and that from Venta D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 11

Table 4 the Greco-Iranian Paleobioprovince. These suggest that Measurements (in mm) and ratios of the M1 of M. pentelicus pentelicus from Venta M. pentelicus would have generally been an inhabitant of relatively del Moro compared to the descriptive statistics for the same variables in several open, mosaic habitats, consisting of a patchily forested open samples of Mesopithecus spp.a savanna with bushy areas and gallery forests (Delson, 1994; MD N Mean SD 95% CI Range Youlatos, 2003; Koufos et al., 2003; Koufos, 2009b), although evi- M. p. pentelicus (Venta del Moro) 1 7.40 dence from the Bulgarian sites suggests that M. pentelicus might M. p. pentelicus (Pikermi) 40 6.90 0.33 6.80 7.01 6.3 7.8 have inhabited slightly wooded, homogeneous landscapes with a M. p. pentelicus (Maramena) 5 7.72 0.09 7.61 7.83 7.6 7.8 well-developed grassy herbaceous layer (Clavel et al., 2012). Mes- M. p. pentelicus (other localities)b 12 7.25 0.52 6.92 7.58 6.5 8.2 M. pentelicus subsp. indet.c 19 6.88 0.58 6.60 7.16 5.6 7.5 opithecus monspessulanus, on the other hand, would probably have M. monspessulanusd 9 6.43 0.28 6.22 6.65 6.0 6.9 inhabited more wooded and humid environments, in agreement

BL N Mean SD 95% CI Range with inferences of its more arboreal behavior and relevant paleo- environmental reconstructions (Delson, 1973, 1975, 1994; Szalay M. p. pentelicus (Venta del Moro) 1 5.80 and Delson, 1979; Ciochon, 1993). M. p. pentelicus (Pikermi) 40 5.97 0.36 5.85 6.08 5.1 6.9 M. p. pentelicus (Maramena) 5 5.91 0.17 5.70 6.12 5.7 6.2 Sedimentological and geological data from Venta del Moro are M. p. pentelicus (other localities)b 12 6.15 0.55 5.80 6.50 5.5 7.0 indicative of deposition at the shores of a paleolacustrine envi- M. pentelicus subsp. indet.c 19 5.77 0.33 5.62 5.93 5.3 6.5 ronment characterized by strong hydrologic seasonality (Montoya d M. monspessulanus 9 5.01 0.23 4.83 5.19 4.6 5.3 et al., 2002). A detailed palynological study of the stratigraphic BLI N Mean SD 95% CI Range sequence of Venta del Moro (Casas-Gallego et al., 2015) has shown M. p. pentelicus (Venta del Moro) 1 78.38 the existence of a steppe-like landscape under a dry climate, with M. p. pentelicus (Pikermi) 40 86.57 5.64 84.77 88.38 74.7 100.0 hygrophytic elements further indicating the presence of swamp M. p. pentelicus (Maramena) 5 76.62 1.83 74.34 78.89 73.9 79.0 environments. This study has also inferred the persistence and b M. p. pentelicus (other localities) 12 84.92 5.85 81.20 88.63 76.3 93.3 expansion of open ecosystems, which probably began to develop M. pentelicus subsp. indet.c 19 84.47 8.17 80.53 88.41 74.2 102.3 M. monspessulanusd 9 77.91 2.23 76.20 79.63 75.4 81.5 during the Early Miocene and were important in the central sector of the Iberian Peninsula during the Middle Miocene (Casas-Gallego a MD ¼ mesiodistal length; BL ¼ maximum buccolingual breadth; BLI ¼ breadth/ length index; N ¼ sample size; SD ¼ standard deviation; CI ¼ confidence interval. et al., 2015). Overall, the presence of M. pentelicus instead of b This material comes from the localities of Veles, Kromidovo, Maragheh (middle M. monspessulanus at Venta del Moro fits well with the relatively beds), Pikermi II (Chomateres), Molayan, Baltavar, and Casino; the last four localities open and mosaic habitats generally inhabited by this taxon in other are attributed to M. pentelicus cf. pentelicus. localities. c This material comes from the localities of Vathylakkos-2, Vathylakkos-3, Ravin X, Perivolaki, Kalimantsi, and Kumanovo. d This material comes from the localities of Villafranca d'Asti, Montpellier, 4.3. Biochronology and paleobiogeography Wolfersheim,€ and Hajnacka. The dental remains reported in this paper, attributed to M. p. pentelicus, show that Mesopithecus was likely present in the Iberian del Moro can be attributed to M. p. pentelicus. The newly reported Peninsula during the Late Miocene. Besides the previous (pre- deciduous teeth from Spain therefore represent a significant liminary) reports of Mesopithecus sp. at Venta del Moro (Montoya addition to the available sample of deciduous teeth of the latter et al., 2006; Marigo et al., 2014), the presence of this genus in taxon, previously mainly known on the basis of the sample from Iberia had been previously reported by Guillen Castejon (2010) on Maramena and, to a lesser extent, Pikermi. the basis of dental remains from the karstic locality of Canal Negre 1 (Barcelona, Spain). In particular, an M1-M2 series was attributed to 4.2. Paleoenvironmental inferences M. pentelicus (see Guillen Castejon, 2010: Pl. 9 Fig. 9), whereas an isolated molar, initially identified as an M1 of Mesopithecus sp., was The wide geographical distribution attained by Mesopithecus later reassigned by the same author to an M1 of M. monspessulanus during the Turolian, including sites with low humidity conditions (see Guillen Castejon, 2013: Figs. AeD). By size and morphology, (Eronen and Rook, 2004), is likely to be related to the ecology and the latter specimen rather seems to be a dP4 of a papionin and, in biology of this taxon, characterized by an opportunistic diet any case, Canal Negre 1 represents a mixed assemblage of Late including hard seeds and a semi-terrestrial locomotor repertoire Miocene to Pleistocene faunas (Guillen Castejon, 2010). Therefore, (see below). Thus, the diet of Mesopithecus has been inferred to be even if the presence of Mesopithecus was confirmed there, its re- more similar to that of extant odd-nosed langurs than to that of mains would be uncontextualized from a biochronological view- leaf-eating langurs (Pan et al., 2004). This is supported by 2D point. On the contrary, the remains from Venta del Moro can be microwear studies, indicating a mixed frugivorous-folivorous diet accurately dated to 6.2 Ma (Gibert et al., 2013), thus conclusively with hard fruits and/or seeds for Mesopithecus (Reitz and Benefit, indicating that Mesopithecus dispersed into westernmost Europe by 2001; Reitz, 2002; Merceron et al., 2009a; Solounias et al., 2010), at least the latest Miocene (MN13). as well as by more refined dental microwear texture analyses, The record of Mesopithecus at Venta del Moro represents the suggesting that Mesopithecus was a widespread, opportunistic reappearance of primates in the fossil record of the Iberian Penin- feeder that often consumed hard seeds (Merceron et al., 2009b). sula, nearly three million years after the disappearance of both With regard to positional behavior, M. p. pentelicus shows a mosaic hominoids and pliopithecoids during the late Vallesian (Moya-Sol a of arboreal and terrestrial-related features, indicating that this et al., 2001; Marigo et al., 2014; Fig. 6). The last Iberian hominoid taxon was adaptated to a semi-terrestrial, -like lifestyle, record, represented by Hispanopithecus, occurs at the site of La probably being at least as terrestrial as the extant langur Semno- Tarumba 1 (Casanovas-Vilar et al., 2011; Alba, 2012) at ca. 9.5 Ma, pithecus entellus (Delson, 1973, 1975, 1994; Szalay and Delson, 1979; whereas the last pliopithecoid record belongs to Egarapithecus, Zapfe, 1991; Youlatos, 1999, 2003; Youlatos and Koufos, 2010; from the latest Vallesian (ca. 9 Ma) of Torrent de Febulines (Moya- Youlatos et al., 2012; but see Escarguel, 2005, for an alternative Sola et al., 2001). The disappearance of these groups from the interpretation). Both the paleodietary and the locomotor inferences Iberian record shortly after the Vallesian Crisis has been related to about Mesopithecus agree with paleoenvironmental re- the gradual replacement of tropical and subtropical plants by de- constructions drawn for Pikermi and other Turolian localities from ciduous elements due to climatic changes involving cooling and 12 D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14

Figure 6. Biochronologic scheme depicting the dispersals that took place in the Iberian Peninsula during the Late Miocene compared to the approximate stratigraphic ranges of Mesopithecus records (Mesopithecus sp. indet. excluded) elsewhere in Europe (see Table 1 for further details). Question mark denotes uncertainty about the Vallesian age of the Wissberg tooth. increased seasonality that took place during the Late Miocene Event) by Agustí et al. (2006) and corresponds to the so-called Third (Agustí et al., 2003; Alba, 2012; Marmi et al., 2012). In Eastern African-Iberian Dispersal of Gibert et al. (2013). We would prefer to Europe, the highly seasonal new climatic conditions allowed for a term these intervals of range extension just “mammalian turn- rapid dispersal of the colobine monkey Mesopithecus, which was overs,” but we do not wish to add yet another formal descriptor to already present in Greece by MN11 (early Turolian, ca. 8.5 Ma; the already crowded list. The presence of this papinion of African Koufos, 2009a, b; Table 1 and Fig. 6). In contrast, the presence of this origin in the roughly coeval (5.4e5.3 Ma) locality of Moncucco taxon in Western Europe cannot be reasonably ascertained until the Torinese in Italy (Alba et al., 2014), together with its absence from late Turolian (MN13), when Mesopithecus is recorded from several Venta del Moro and other Iberian sites, supports the contention Italian sites (Gentili et al., 1998; Rook, 1999, 2009; Alba et al., 2014; that dispersed into Europe at the same time as the sea Table 1), as well as from Venta del Moro in Spain. Neither Meso- level drop associated with the Messinian Salinity Crisis. Also about pithecus nor other primates have been recorded from MN11 and this time, we see the dispersal into Iberia of both Asian and African MN12 of the Iberian Peninsula (Marigo et al., 2014; Fig. 6), in spite taxa with semiarid adaptations, such as gerbils (Agustí, 1990; of the rich faunal assemblages from the Teruel Basin and other sites Minwer-Barakat et al., 2009). It is not yet clear if these migrations such as Crevillente (e.g., Alcala, 1994; Montoya, 1994). It is therefore crossed the Gibraltar region (as suggested by Gibert et al., 2013 unlikely that the absence of Mesopithecus from the Iberian record among others) or if they involved, as it seems most likely in the before MN13 might be due to a preservation bias, but instead it is case of Mesopithecus, the longer route through the Middle East that probably a real absence. was already available from pre-Messinian times (Gilbert et al., The appearance of Mesopithecus in the Iberian Peninsula at ca. 2014). 6.2 Ma, during the late Turolian (MN13) or Ventian (sensu Morales et al., 2013), is linked to an important faunal renewal that affected Acknowledgments more than 50% of the Turolian macromammals. Thus, the verte- brate fossil assemblage of Venta del Moro consists of a mixture of This work has been funded by the Spanish Ministerio de Econ- native taxa and immigrants from Southeast Europe, Asia, and Africa omía y Competitividad (CGL0211-28681, CGL2011-25754, CGL2011- (Montoya et al., 2006; van der Made et al., 2006; Gibert et al., 2013; 27343, CGL2014-54373-P, and RYC-2009-04533 to D.M.A.), the Morales et al., 2013; Colombero et al., 2014). This faunal turnover Spanish Ministerio de Educacion (AP2010-4579 to M.P.), and the (documented at Venta del Moro, prior to the start of the Messinian Generalitat de Catalunya (2014 SGR 416 GRC). Fieldwork at Venta Salinity Crisis) has been named the Second Messinian Mammalian del Moro from 1995 onwards was approved and supported by the Event (or Paraethomys Event) by Agustí et al. (2006) and corre- Conselleria de Cultura of the Generalitat Valenciana, with the sponds to the so-called Second African-Iberian Dispersal of Gibert permission of the Ministerio de Fomento and the company ADIF. et al. (2013). The next record of cercopithecids in the Iberian J.A. thanks the “Proyecto Prometeo” of the “Secretaría de Educacion Peninsula is Macaca from the karstic site of Almenara-Casablanca M Superior, Ciencia, Tecnología e Innovacion, ” Republic of Ecuador. € (Kohler et al., 2000), coinciding with another faunal turnover dated D.M.A. drafted this paper mostly during a short stay at the Amer- between 5.5 and 5.3 Ma and therefore contemporary with the ican Museum of Natural History (AMNH) with the New York Con- Messinian Salinity Crisis (Gibert et al., 2013). This faunal exchange sortium in Evolutionary Primatology (NYCEP) in 2011, and he has been termed the Third Messinian Mammalian Event (or Gerbil thanks all their staff, faculty, and students for their hospitality and D.M. Alba et al. / Journal of Human Evolution 88 (2015) 1e14 13 collaboration. Finally, we thank the Editor (Sarah Elton), Associate Delson, E., Terranova, C.J., Jungers, W.L., Sargis, E.J., Jablonski, N.G., Dechow, P.C., 2000. Editor, and three anonymous reviewers for their constructive Body mass in Cercopithecidae (Primates, Mammalia): estimation and scaling in extinct and extant taxa. Anthropol. Papers Am. Mus. Nat. Hist. 83, 1e159. comments and suggestions. Delson, E., Thomas, H., Spassov, N., 2005. Fossil Old World monkeys (Primates, Cercopithecoidae) from the Pliocene of Dorkovo, Bulgaria. Geodiversitas 27, 159e166. Dumurdzanov, N., Serafimovski, T., Burchfiel, B.C., 2004. Evolution of the Neogene- Pleistocene Basins of Macedonia. Geol. Soc. Am. Dig. 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