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Journal of Human Evolution 123 (2018) 141e147

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Journal of Human Evolution

journal homepage: www.elsevier .com/locate/ jhevol

Macaque remains from the early Pliocene of the Iberian Peninsula

David M. Alba a, *, Eric Delson b, c, d, e, a, Jorge Morales f, Plini Montoya g,Gregorio Romero h, i

a Institut Catal�a de Paleontologia Miquel Crusafont, Universitat Aut�onoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de La UAB,08193 Cerdanyola del Vall�es, Barcelona, Spainb Department of Anthropology, Lehman College of the City University of New York, 250 Bedford Park Boulevard West, Bronx, NY 10468, USAc Department of Vertebrate Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY 10024, USAd PhD Program in Anthropology, The Graduate Center of the City University of New York, 365 Fifth Avenue, New York, NY 10016, USAe New York Consortium in Evolutionary Primatology, New York, NY, USAf Departamento de Paleobiología, Museo Nacional de Ciencias Naturales (CSIC), Jos�e Guti�errez Abascal 2, 28006 Madrid, Spaing Departament de Bot�anica i Geologia, Universitat de Val�encia, Doctor Moliner 50, 46100 Burjassot, Spainh Direcci�on General de Bienes Culturales de la CARM, Casa Díaz Cassou, C/ Santa Teresa 21, 30071 Murcia, Spaini Grupo de Investigaci�on de Geología, Facultad de Química, Universidad de Murcia, 30100 Murcia, Spain

a r t i c l e i n f o

Article history:Received 28 March 2018Accepted 16 July 2018Available online 20 August 2018

Keywords:MacacaMessinianTurolianPuerto de la CadenaMurciaSpain

* Corresponding author.E-mail address: david.alba@icp.cat (D.M. Alba).

https://doi.org/10.1016/j.jhevol.2018.07.0050047-2484/© 2018 Elsevier Ltd. All rights reserved.

a b s t r a c t

Macaques dispersed out of Africa into Eurasia in the framework of a broader intercontinental faunalexchange that coincided in time with the sea level drop associated with the Messinian Salinity Crisis.They are first recorded in Europe (Italy and Spain) by the latest Miocene, being subsequently recorded allover Europe, albeit sparsely, throughout the Pliocene and Pleistocene. These fossil European macaquesare attributed to several (sub)species of the extant Barbary macaque (Macaca sylvanus). In Iberia, fossilmacaques are best documented by Macaca sylvanus florentina from various Early Pleistocene sites,whereas their published Pliocene record is very scarce. Here we report the oldest post-Messinianoccurrence of macaques in the Iberian Peninsula, based on the description and metrical comparisonsof two upper teeth (a male canine and a third molar of two different individuals) from the early Pliocene(MN14, 5.0e4.9 Ma) site of Puerto de la Cadena (Murcia, SE Spain). The male C1 is fully comparable inmorphology with those of extant and fossil M. sylvanus, and larger than those of Mesopithecus. The M3, inturn, displays the typical papionin morphology that characterizes the dentally-conservative genusMacacadthereby discounting an alternate assignment to either the extinct colobine monkey Meso-pithecus or the more dentally-derived papionin Theropithecus. Dental size and proportions of the M3

further support an attribution to an extinct subspecies of M. sylvanus instead of the larger papioninParadolichopithecus. Mostly on biochronologic grounds, the two macaque teeth from Puerto de la Cadenaare here assigned to Macaca sylvanus cf. prisca, albeit tentatively, given the lack of clear-cut criteria todistinguish this subspecies from the younger Macaca sylvanus florentina. The described material repre-sents the oldest well-dated Pliocene record of macaques in Iberia, predating the record of Para-dolichopithecus by almost 1.5 million years.

© 2018 Elsevier Ltd. All rights reserved.

1. Introduction

1.1. Puerto de la Cadena

Continental fossil vertebrates from the late Neogene fossili-ferous outcrops of the Puerto de la Cadena area (Murcia, SE Iberian

Peninsula) have been known for many decades, particularly on thebasis of two sites (La Alberca and La Paloma) correlated to the latestMiocene Mammal Neogene (MN) unit MN13 (Montenat andCrusafont, 1970; Mein et al., 1973; Aguirre et al., 1974; de Bruijnet al., 1975; Morales, 1984; L�opez Martínez, 1989; Freudenthalet al., 1998; Manche~no Jim�enez and Fierro Bandera, 2011; P�erez-García et al., 2011; Morales et al., 2013). The location in 2008 of anew site, due to the construction of highway MU-31 in the area ofPuerto de la Cadena, led to the discovery of more than 2000 fossilremains during successive emergency paleontological works in the

D.M. Alba et al. / Journal of Human Evolution 123 (2018) 141e147142

following three years (Manche~no Jim�enez and Fierro Bandera,2011; Pi~nero et al., 2017). This fossil vertebrate assemblage fromPuerto de la Cadena was recently described by Pi~nero et al. (2017).Although preliminary accounts suggested a latest Miocene age forthis site (Manche~no et al., 2013), Pi~nero et al. (2017) conclusivelycorrelated it to the early Pliocene. Among the 26 reported species,Pi~nero et al. (2017) noted the presence of a cercopithecine, whichon the basis of the two available teeth was assigned to Macaca sp.

Figure 1. Location map of Puerto de la Cadena within the Iberian Peninsula (left) andwithin the province of Murcia (inset, right).

1.2. The fossil macaques from the Iberian Peninsula

Macaques are first recorded in Eurasia by the latest Miocene(MN13), based on fossil remains from Almenara-Casablanca M,Spain (K€ohler et al., 2000) and Moncucco Torinese, Italy (Alba et al.,2014), all assigned to cf. Macaca sp. The remains from MoncuccoTorinese are confidently dated to 5.41e5.33 Ma (Colombero et al.,2017), whereas those from Almenara-Casablanca M are roughlycontemporaneous with the Messinian Salinity Crisis (ca. 5.9e5.3Ma; Agustí et al., 2006; Minwer-Barakat et al., 2009; Alba et al.,2014). Coupled with the lack of papionin remains from older lo-calities in Eurasia as a whole, this evidence suggests that macaquesprobably did not disperse out of Africa until the sea level dropassociated with the Messinian Salinity Crisis (Agustí et al., 2006;Alba et al., 2014, 2015; Colombero et al., 2017).

It is uncertainwhethermacaques dispersed into Europe throughsouthern Iberia or followed the Middle East route that was alreadyavailable from pre-Messinian times (Alba et al., 2015). In any case,the dispersal of macaques coincides with a major mammalianturnover that took place ca. 5.5e5.3 Ma and involved multipleintercontinental dispersals between Africa and Europedthe so-called ‘Gerbil Event’ (Agustí et al., 2006) or ‘Third African-IberianDispersal’ (Gibert et al., 2013). Subsequently, during the Plio-Pleistocene, European fossil macaques are represented by severaltaxa that belong to the lineage ofMacaca sylvanus (Linnaeus, 1758),the extant Barbary macaque from northern Africa (Delson, 1974,1980; Szalay and Delson, 1979; Alba et al., 2011, 2014, 2016),which on phylogenetic evidence is considered the basal-mostmember of the genus (e.g., Springer et al., 2012). The Plio-Pleistocene remains of European macaques are customarilyattributed to chronologically successive subspecies of M. sylvanus,which due to the lack of cranial remains are not particularly wellcharacterized (Szalay and Delson, 1979; Delson, 1980; Alba et al.,2008, 2011, 2016; Marig�o et al., 2014). Only the endemic macaquefrom Sardinia, Macaca majori Azzaroli, 1946, is customarilyconsidered distinct enough from extant M. sylvanus to be consid-ered a separate species (Zanaga, 1998; Rook and O'Higgins, 2005;Smith et al., 2014).

Early Pliocene macaques from Europe are customarilyassigned to M. sylvanus prisca Gervais, 1859, which ranges fromthe earliest Ruscinian to the early Villafranchian (MN14eMN16;Szalay and Delson, 1979; Delson, 1980). The oldest record corre-sponds to the site of Montpellier, France (type locality; MN14, ~5Ma), followed by Csarnota 2, Hungary (late MN15, ~3.6e3.2 Ma;Minwer-Barakat et al., 2012a) and several MN16 localities, such asFornace RDB, Italy (early MN16, ~3.2e3.0 Ma; see Rook et al.,2001, and references therein). Although M. s. prisca was consid-ered to be somewhat smaller than more recent and extant sub-species (Szalay and Delson, 1979; Delson, 1980), subsequentworks have failed to substantiate such claims (Delson et al., 2000;Rook et al., 2001). Here we figure and describe in detail the fossilmacaque remains from Puerto de la Cadena, which represent theoldest securely dated record of Macaca from the Iberian Pliocene,and tentatively one of the oldest occurrences of M. s. prisca inEurope.

2. Age and geological background

The site of Puerto de la Cadena is located 8 km SW of the city ofMurcia (Fig. 1), close to the towns of La Alberca and El Palmar, onthe northern flank of the Sierra de Carrascoy y el Valle. From ageological viewpoint, the fossiliferous outcrops of Puerto de laCadena (Murcia-Carrascoy Basin) belong to the Cigarr�on Unit,whose sediments were deposited under shallowmarine conditionstowards the bottom and within a continental setting towards thetop (Pi~nero et al., 2017). The lower boundary of this unit consists ofan erosional surface that has been correlated to the end-Messiniandiscontinuity, thus being interpreted as a post-Messinian regressivesequence (Pi~nero et al., 2017). The site of Puerto de la Cadena islocated in the upper portion of the Cigarr�on Unit, being composedof alternating sandstones and mudstones with some conglomeraticlayers, indicative of a fluvial depositional environment with chan-nels and floodplain development (Manche~no Jim�enez and FierroBandera, 2011; Pi~nero et al., 2017).

Magnetostratigraphic analyses (Pi~nero et al., 2017) indicate thatthe short section from Puerto de la Cadena records a reversemagnetozone, which is also recorded in the upper portion of thenearby Barranco del Cigarr�on section, being correlated to eitherC3n.3r (4.997e4.896 Ma; Ogg, 2012) or C3n.2r (4.799e4.631 Ma;Ogg, 2012). Correlation with the former has been favored onbiostratigraphic grounds by Pi~nero et al. (2017:112), who furtherconsidered the fauna from Puerto de la Cadena to be “earliestMN14”. These authors advocated for the placement of the MN13/MN14 boundary within C3n.3r (i.e., about 0.3 million years youngerthan the Miocene/Pliocene boundary at 5.33 Ma), in agreementwith previous proposals by some authors (Opdyke et al., 1997;Agustí et al., 2001). However, such a proposal is at odds with themost common, recently strongly supported view that the MN13/MN14 boundary is somewhat older and might even roughly coin-cide with the Miocene/Pliocene boundary (García-Alix et al., 2008;Minwer-Barakat et al., 2012b; Pi~nero and Agustí, 2017; Pi~nero et al.,2018).

Pi~nero et al. (2017) did not provide positive evidence for theirinterpretation of Puerto de la Cadena as earliest MN14. Theycorrectly indicated that it appears to be ca. 200 kyr younger thanSif�on 413, which occurs just above the local end-Messinian erosionand thus must be within the Pliocene, at ca. 5.3 Ma, but they did notcorrelate Sif�on 413 to a MN unit. Pi~nero and Agustí (2017) revisedthe rodent assemblages from the Sif�on de Librilla section (FortunaBasin) and made a good case that Sif�on 413 yielded several taxaindicative of MN14, especially Occitanomys brailloni, which theysuggested is the best marker for the beginning of MN14 in southernSpain. Therefore, Puerto de la Cadena does not represent the

D.M. Alba et al. / Journal of Human Evolution 123 (2018) 141e147 143

earliest part of MN14, although with an age of 5.0e4.9 Ma, it can beconclusively correlated to both MN14 and the early Pliocene.

3. Material and methods

The macaque fossil remains from Puerto de la Cadena describedhere consist of a left C1 (MAM VLP3-62) and a left M3 (MAM VLP2-715). Although these remains were originally studied by the seniorauthor of this paper in 2013 at the Museo Paleontol�ogico de Elche(MUPE; Elche, Spain), they are currently housed at the MuseoArqueol�ogico de Murcia (MAM; Murcia, Spain). These teeth corre-spond to two different individuals, since they were recovered fromtwo different outcropping slopes of the site (no. 3 in the case of theC1, and no. 2 in the case of theM3). The acronym (VLP) of the catalognumbers stands for ‘Venta La Paloma,’ which is the informal nameoriginally given to the site that is currently known as Puerto de laCadena (Pi~nero et al., 2017).

Measurements of mesiodistal crown length (MD) and bucco-lingual crown breadths (BL), taken to the nearest 0.1 mm with adigital caliper, were used to compare the Puerto de la Cadenaspecimens with other available samples of extantM. sylvanus, fossilMacaca and Paradolichopithecus (and for canines, Mesopithecusspecies and Dolichopithecus ruscinensis) from Europe. In the case ofupper canines, MD was defined as the maximum crown diameter,and BL as the maximum crown diameter perpendicular to MD.Bivariate dental plots of BL vs. MDwere employed to visually assessthe size and proportions of the dental remains from Puerto de laCadena with regard to the comparative sample. Given that caninesare generally not diagnostic to distinguish between cercopithecidsubfamilies (e.g., Delson, 1973; Hill and Gundling, 1999; Delsonet al., 2005), and the fact that Macaca is known to co-occur withthe colobinesMesopithecus and/orDolichopithecus in some Pliocenelocalities (Delson, 1974; Szalay and Delson, 1979; Delson et al.,2005), upper canine measurements for the latter genera werealso included in the comparisons. Dental measurements for thePuerto de la Cadena specimens and the comparative sample arereported in Supplementary Online Material (SOM) Table S1. Theywere taken from NYCEP's PRIMO (PRImate Morphometrics Online)database (http://primo.nycep.org) or the literature (Alba et al.,2011; Bona et al., 2016), or measured by one of the authors of thispaper (D.M.A.) at the American Museum of Natural HistoryDepartment of Mammalogy (AMNH-M) in New York, USA.

4. Systematic paleontology

Order Primates Linnaeus, 1758Infraorder Catarrhini �E. Geoffroy Saint-Hilaire, 1812Superfamily Cercopithecoidea Gray, 1821Family Cercopithecidae Gray, 1821Subfamily Cercopithecinae Gray, 1821Tribe Papionini Burnett, 1828Subtribe Macacina Owen, 1843Genus Macaca Lac�ep�ede, 1799Macaca sylvanus (Linnaeus, 1758)Macaca sylvanus cf. prisca Gervais, 1859(Fig. 2)

4.1. Description

Upper canine This specimen (Fig. 2fej) preserves the root (buccalheight ¼ 20.6 mm) and the basal-most portion of the crown(preserved buccal crown height ¼ 11.9 mm), thus enablingreliable measurements of the relevant dimensions(MD ¼ 11.2 mm, BL ¼ 8.6 mm; see also SOM Table S1). This tooth

displays some degree of rounding and erosion overall (mostevident at the crown and root preserved apices), which suggestssome transport before burial. It also shows marked abrasionthroughout the surface of the root and the crowndnot only aresome superficial flakes of enamel and cement missing, but theremaining portions of the root and crown are abraded, probablyindicating soil acid corrosion.

On the basis of its large dimensions, extension of the mesio-lingual groove onto the root and overall dagger-like morphology,this canine can be attributed to a male individual. Both the root andthe preserved portion of the crown display a labiolingually com-pressed (crown breadth/length index 77%), subtriangular cross-section. The root is somewhat tilted distally, displaying in buccaland lingual views a rather straight distal contour, but a moremarkedly convex mesial profile. There is a broad and markedmesiolingual sulcus that runs along the root and the crown throughthe cervix, as well as a shallower and more diffuse lingual sulcus.These sulci result in biconvex lingual and, especially, mesial pro-files, which contrast with the flat or even slightly convex buccalsides of the root and the crown, respectively. The crown is brokenaway obliquely to a large extent, so that only its basal portion ispreserved (especially on the mesial and buccal sides), therebyexposing the pulp cavity. The cementoenamel junction on thebuccal side shapes a shallow inverted ‘V’. Distally, the basal-mostportion of the honing facet against the P3, which extends basallybeyond the cervix, is preserved.

Upper molar Only the crown of this tooth is preserved (Fig. 2aee).Although small portions of enamel are missing at some points closeto the cervix, the overall dimensions of the crown can be reliablytaken (MD ¼ 10.2 mm, mesial BL ¼ 10.2 mm, distal BL ¼ 8.5 mm;see also SOM Table S1). This tooth displays only a slight degree ofwear, with very minimal dentine exposure at the apices of allmain cusps except the paracone. There is a small contact facet onthe mesial crown wall, but no distal contact facet; in combinationwith the marked distal tapering and the presence of an accessorycusp on the distal margin (frequent in papionin last molars,although usually being situated more buccally; Delson, 1973),these features point to the tooth being an M3.

This molar is as broad as long (breadth/length index 100%) anddisplays a suboval contour that markedly tapers distally, since themesial lobe is much broader than the distal one. The crown isslightly longer buccally than lingually, due to the more mesial po-sition of the paracone compared to the protocone. The crown ismoderately waisted, with biconvex lingual and buccal occlusalprofiles. The mesial and distal lobes are separated by a moderately-developed median lingual cleft, which is located well above thecervix and displays no interconule, as well as a groove-like medianbuccal cleft that does not reach the crown base either. The mesiallingual and buccal clefts are narrow and groove-like, whereas nodistinct distal clefts can be discerned. The shallow median buccaland lingual notches do not surpass one-third of crown height. Theocclusal surface displays the typical cercopithecid bilophodontpattern, with four main cusps that are not particularly compressedbuccolingually. The paracone is slightly larger and higher than theprotocone, and the distal cusps are shorter, less extensive andmoreclosely packed than the mesial ones. The degree of lingual flare isslightly more marked than the buccal. The mesial fovea is crescent-shaped and extends to some extent above the mesial shelf. Thedistal fovea is intermediate in size between the mesial and thecentral ones. The hypocone has a twinned appearance, due to thedevelopment of the posthypocrista into an accessory cuspule(distoconule), which is separated from the hypocone by subtlebuccal and lingual grooves and is smaller than the four main cusps.There are no discernible cingula.

Figure 2. Dental remains ofMacaca sylvanus cf. prisca from Puerto de la Cadena. aee) Left M3 (MAM VLP2-715), in occlusal (a), mesial (b), lingual (c), distal (d), and buccal (e) views.fej) Left C1 (MAM VLP3-62), in occlusal (f), mesial (g), lingual (h), distal (i) and buccal (j) views.

D.M. Alba et al. / Journal of Human Evolution 123 (2018) 141e147144

4.2. Morphometric comparisons

The dental size and proportions of the two macaque teeth fromPuerto de la Cadena (SOM Table S1) are compared with those ofother fossil macaques in Figure 3. The male C1 (Fig. 3a), like those ofPlio-Pleistocene European subspecies of M. sylvanus for which thistooth is known, overlaps in size with those of the extant Barbarymacaque, fromwhich onlyM. majorimarkedly differs by its smallerocclusal dimensions. Given the variability of the extant subspecies,and the very limited sample available for extinct M. sylvanus sub-species, no taxonomic conclusions can be drawn from this tooth,other than concluding that its dimensions are compatible with anassignment to this species. An attribution to a colobine can bediscounted on size grounds in the case of Mesopithecusdsince themale canines of this genus are much smaller (intermediate in size

Figure 3. Bivariate dental plots of mesial buccolingual (BL) breadth vs. mesiodistal (MD) lecompared to those of other fossil European and African macaques, Paradolichopithecus, Mes(C1_). b) Upper first molar (M1). The measurements for the comparative sample are report

between those of M. majori and those of extant M. sylvanus;Fig. 3a)dbut not so confidently in the case of Dolichopithecus.

More taxonomically-informative comparisons can be maderegarding theM3 from Puerto de la Cadena due to the larger sampleavailable for fossil macaques (Fig. 3b). A referral to the largerpapionin Paradolichopithecus, which is recorded in the late Plioceneof Iberia (Delson et al., 2014; Marig�o et al., 2014), can be confidentlydiscounted based on its much larger occlusal dimensions (Fig. 3b).In contrast, the dimensions of the M3 from Puerto de la Cadena(Fig. 4a) only minimally exceed those of M. sylvanus sylvanus(Fig. 4feh), and fit well with those of some extinct subspecies(Figs. 3b and 4bee; see also Alba et al., 2011), including a malespecimen of M. s. prisca from Fornace RDB (Fig. 4c; see also Rooket al., 2001:Fig. 3). Male and female specimens of the extant Bar-barymacaque largely overlap, althoughmales tend to display larger

ngth (in mm) for the two teeth of Macaca sylvanus cf. prisca from Puerto de la Cadena,opithecus and Dolichopithecus, as well as extant Macaca sylvanus. a) Male upper canineed in SOM Table S1.

Figure 4. Macaca sylvanus cf. prisca from Puerto de la Cadena (a) compared to extinct Macaca sylvanus prisca from Montpellier (b) and Fornace RDB (c), Macaca sylvanus florentinafrom Quibas (dee), and extant Macaca sylvanus sylvanus from North Africa (feh). All specimens are depicted in occlusal view. a) Left M3 (MAM VLP2-715). b) Left M3 (FSL 40136). c)Left M3 (NMB VJ 88). d) Left M3 (UM Q03-E-155b). e) Right (reversed) M3 (UM Q05-Ec-87). f) Right (reversed) M3 (AMNH-M 19014, female). g) Left M3 (AMNH-M 2060/5484,female). h) Left M3 (AMNH-M 185277, female). Specimens identified as UM were formerly housed at the Universidad de Murcia (Spain; Alba et al., 2011), but currently they arehoused in the MAM collections. The specimens labeled as FSL and NMB are respectively housed in Lyon and Basel (for further details on institutional acronyms, see SOM Table S1).

D.M. Alba et al. / Journal of Human Evolution 123 (2018) 141e147 145

dimensions on average. The extinct taxa also overlap with theextant subspecies to a large degree, although M. majori mostlyoverlaps with female specimens, whereas both M. libyca (Stromer,1920) from the late Miocene of north Africa and extinct subspe-cies of M. sylvanus mostly overlap with male specimens of theBarbary macaque, and show in all instances larger dimensions thanM. majori. Unlike Macaca sylvanus pliocena Owen, 1846 and thesingle specimen of M. libyca, the M3 from Puerto de la Cadena re-sembles some specimens of the early Pliocene M. s. prisca and theyounger Macaca sylvanus florentina (Cocchi, 1872) in having largerdimensions than the extant subspecies. As such, the size of thedescribed M3 is compatible with an attribution of the Puerto de laCadena macaque to either of these two extinct subspecies ofM. sylvanus.

5. Discussion and conclusions

The male C1 from Puerto de la Cadena is fully comparable inmorphology with those of both extant and fossil M. sylvanus (e.g.,see Alba et al., 2011). Although it should be taken into account thatthis tooth is less informative from a taxonomic viewpoint thancheek teeth (Delson, 1973), an alternate assignment to the colobineMesopithecus can be discounted on size grounds. In turn, the uppermolar from Puerto de la Cadena is identified as an M3 and displaysthe typical papionin morphology that characterizes the dentally-conservative genus Macaca (Fig. 4). This fact allows us to rule outan attribution to the colobine monkeys Mesopithecus or Dolichopi-thecus, which would be characterized by a more marked occlusalrelief, deeper median notches, a deeper lingual cleft, more bucco-lingually compressed cusps, a larger mesial fovea, a more waisted

crown (more markedly biconvex buccal and lingual contours), anasymmetrical distal margin, and a less developed buccal flare of thecrown (Delson, 1973, 1975; Szalay and Delson, 1979), as well assmaller size. Several of these features further discount an attribu-tion to the papionin Theropithecus, which is characterized by highercusps, more deeply excavated foveae, and better-developednotches, among other features (Delson, 1973, 1975; Szalay andDelson, 1979). Finally, an attribution to the dentally-generalizedpapionin Paradolichopithecus can be discounted on the basis of size.

An attribution of the material from Puerto de la Cadena to thegenusMacaca, and in particular toM. sylvanus, is further supportedby the dental size and proportions of the described remains. In fact,the teeth are compatible with an assignment to M. s. prisca, whichwould seem logical on biochronologic (and paleobiogeographic)grounds. The teeth of specimens assigned to the various extinctsubspecies of M. sylvanus overlap to a large extent with those ofextantM. sylvanus (e.g., Alba et al., 2011). Thus, althoughM. s. priscawas considered to be smaller than the Barbary macaque and otherextinct subspecies (Szalay and Delson, 1979; Delson, 1980;Jablonski, 2002), in fact there is a considerable overlap (Rooket al., 2001). Our comparisons indicate that the M3 from Puertode la Cadena is somewhat larger than those of the extant Barbarymacaque, as is also the case for some specimens of both M. s. priscaand M. s. florentina (see Alba et al., 2011). Given the lack of cleardental metric criteria to distinguish between these two subspeciesof M. sylvanus, our attribution to the former largely relies on bio-chronological criteria and, as such, it must be considered tentative(M. sylvanus cf. prisca), at least until more complete macaque re-mains from similarly-aged European localities enable a morecomplete taxonomic assessment.

D.M. Alba et al. / Journal of Human Evolution 123 (2018) 141e147146

Before going extinct during the Late Pleistocene, Europeanmacaques attained awide geographical distribution throughout thePlio-Pleistocenedeven if their sparse record suggests that, due tosuboptimal ecological conditions, they were not a common faunalelement (Meloro and Elton, 2012; Elton and O'Reagan, 2014). Mostof the macaque records from the Iberian Peninsula are based onfragmentary remains that have not been assigned to (sub)speciesand/or described in detail (Marig�o et al., 2014, and referencestherein; see also Elton and O'Reagan, 2014:Appendix A). There are,however, some exceptions, including Late Pleistocene remainsassigned to M. s. pliocena (Casta~nos et al., 2011) and, especially, themore abundant Early Pleistocene material of M. s. (cf.) florentinafrom sites such as Incarcal (ca. 1.5e1.2 Ma; Alba et al., 2016), Quibas(ca. 1.3e1.0 Ma; Alba et al., 2011), and Vallparadís and Cal Guardiola(ca. 1.2e0.8 Ma; Alba et al., 2008).

The published Pliocene record of Iberian macaques is, incontrast, very scarce, being restricted to the late Pliocene sites ofOrrios 7 (MN15; Mein et al., 1990) and Cova Bonica (MN16, 3.2e2.6Ma; Delson, 1971; Crusafont-Pair�o and Golpe-Posse, 1984), attrib-uted to Macaca sp. by Marig�o et al. (2014) and M. s. prisca byCrusafont-Pair�o and Golpe-Posse (1984). Guill�en Castej�on (2010)further reported the presence of both M. cf. sylvanus prisca andM. s. florentina from the karstic infills of Canal Negre 1. However,given dating uncertainties (Miocene-Pleistocene), and pending amore detailed study, Marig�o et al. (2014) left these remains unas-signed to subspecies rank. Additional remains of M. s. prisca mightbe available from an unpublished collection of papionin fossils fromCova Bonica, where macaques coexisted with a small-sized Para-dolichopithecus species (Delson,1971;Moy�a Sol�a et al., 1990; Delsonet al., 2014; Marig�o et al., 2014). The ongoing study of these re-mains, together with those from the slightly older site of Moreda 1a(late MN15, 3.5e3.2 Ma), should clarify whether Para-dolichopithecus sp. further coexisted with Macaca in the latter lo-cality (Marig�o et al., 2014). In any case, with a much older (earlyMN14) age of ca. 5.0e4.9 Ma, the two macaque teeth from Puertode la Cadena, attributed here to M. s. cf. prisca, clearly attest thepresence of macaques in the earliest Pliocene of the IberianPeninsula, thereby predating by almost 1.5 million years the recordof the larger terrestrial papionin Paradolichopithecus.

Acknowledgments

This work has been funded by the Generalitat de Catalunya(CERCA Program), and the Agencia Estatal de Investigaci�on(Ministerio de Economía, Industria y Competitividad)eEuropeanRegional Development Fund of the European Union (CGL2015-68333-P, MINECO/FEDER-UE; and CGL2016-76431-P and CGL2017-82654-P, AEI/FEDER-UE). Fieldwork at Puerto de la Cadena wascarried out with funding from the Fundaci�on S�enecaeAgencia deCiencia y Tecnología de la Regi�on deMurcia (11891/PHCS/09), and itwas possible thanks to the collaboration of the Demarcaci�on deCarreteras del Estado in Murcia (Ministerio de Fomento) and theDirecci�on General de Bienes Culturales of the ComunidadAut�onoma de la Regi�on de Murcia, and the Asociaci�on CulturalPaleontol�ogica Murciana. D.M.A. further thanks the MuseoArqueol�ogico de Murcia (Murcia, Spain) for permission to study thematerial, the Museo Paleontol�ogico de Elche (Elche, Spain) for thefacilities given, Eileen Westwig (Mammalogy Department, Amer-ican Museum of Natural History) for access to extant primatecomparative material, and Raef Minwer-Barakat for discussion onMessinian biochronology. Finally, we thank Mike Plavcan (Editor),Sarah Elton, Chris Gilbert, and an additional anonymous reviewerfor helpful comments and suggestions that improved a previousversion of this paper.

Supplementary Online Material

Supplementary online material related to this article can befound at https://doi.org/10.1016/j.jhevol.2018.07.005.

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