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

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First Record of Mesopithecus (Cercopithecidae, Colobinae) from the Miocene of the Iberian Peninsula Journal of Human Evolution 88 (2015) 1e14 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol First record of Mesopithecus (Cercopithecidae, Colobinae) 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 (Primates, 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.
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