Ten Years in the Dump: an Updated Review of the Miocene Primate

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Journal of Human Evolution xxx (2016) 1e9

55 Contents lists available at ScienceDirect 56 57 Journal of Human Evolution 58 59 60 journal homepage: www.elsevier.com/locate/jhevol 61 62 63 64 65 1 Ten years in the dump: An updated review of the Miocene 66 2 67 3 primate-bearing localities from Abocador de Can Mata 68 4 69 Q6 (NE Iberian Peninsula) 5 70 6 a, * a b, c a 71 7 Q5 David M. Alba , Isaac Casanovas-Vilar , Miguel Garces , Josep M. Robles 72 8 a Institut Catala de Paleontologia Miquel Crusafont, Universitat Autonoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, 08193, Cerdanyola del 73 9 Valles, Barcelona, Spain 74 10 b Departament de Dinamica de La Terra i de l'Ocea, Facultat de Geologia, Universitat de Barcelona, Martí i Franques s/n, 08028, Barcelona, Spain c 75 11 Institut Geomodels, Grup de Recerca Consolidat de Geodinamica i Analisi de Conques, Universitat de Barcelona, Martí i Franques s/n, 08028, Barcelona, Spain 76 12 77 13 78 14 article info abstract 79 15 80 16 Article history: More than ten years of paleontological fieldwork during the enlargement of the Can Mata Landfill 81 17 Received 2 May 2016 (Abocador de Can Mata [ACM]), in els Hostalets de Pierola (Valles-Pened es Basin, NE Iberian Peninsula) 82 18 Accepted 21 September 2016 led to the recovery of >60,000 Miocene vertebrate remains. The huge sampling effort (due to continuous 83 Available online xxx 19 surveillance of heavy machinery digging activity, coupled with manual excavation and screen-washing of 84 20 sediments) enabled generally rare faunal elements such as pliopithecoid and hominoid primates to be 85 Keywords: 21 found. Thanks to detailed litho-, bio- and magnetostratigraphic controls, accurate dating is possible for 86 Hominoideao 22 all the recovered primate remains from 19 of the 235 localities defined along the 234 m-thick composite Pliopithecoideao 87 stratigraphic sequence of the ACM. Here we report updated estimated (interpolated) ages for these 23 Fossil primateso 88 24 Vales-Pened es Basino paleontological localities and review the timing of the primate succession in this area. Our results 89 indicate that the whole ACM sequence is late Aragonian in age (MN6 and MN7þ8) and includes seven 25 Paleomagnetismo 90 26 Magnetostratigraphyo magnetozones that are correlated to subchrons C5Ar.1r to C5r.2r (ca. 12.6 to 11.4 Ma). Great apes (dryopithecines) are first recorded at 12.4e12.3 Ma, but most of the finds (Anoiapithecus, Pierolapithecus 91 27 and Dryopithecus) cluster between 12.0 and 11.9 Ma, followed by some indeterminate dryopithecine 92 28 remains between 11.7 and 11.6 Ma. Pliopithecoids first appear at 12.1 Ma, being subsequently repre- 93 29 sented by Pliopithecus between 11.9 and 11.7 Ma. The small-bodied hominoid Pliobates is the youngest 94 30 ACM primate, with an estimated age of 11.6 Ma. Although these primates probably overlapped in time, 95 31 their co-occurrence is recorded only twice, at 11.9 Ma (a dryopithecine with Pliopithecus) and at 11.6 Ma 96 32 (a dryopithecine with Pliobates). The rare co-occurrence between great apes and small-bodied catar- 97 33 rhines might be attributable to sampling biases and/or to presumed diverging ecological preferences of 98 34 these groups. In the future, more detailed analyses of the fauna recovered from the long and densely- 99 sampled ACM sequence will hopefully throw new light on this long-standing, unresolved question. 35 100 © 2016 Elsevier Ltd. All rights reserved. 36 101 37 102 38 103 39 104 40 105 41 1. Introduction Josep R. Bataller, together with amateur collector Marius Guerín, 106 42 surveyed the area during the 1920s and first reported on its fossil 107 43 The fossiliferous area of els Hostalets de Pierola (Valles-Pened es fauna (Bataller Calatayud, 1938). Although Guerín collected a 108 44 Basin, NE Iberian Peninsula) is exceptional among the European hominoid M2, it was mistaken for a suid and it was not reassigned 109 45 Miocene in terms of the high diversity of extinct catarrhines to Dryopithecus until much later (van der Made and Ribot, 1999). 110 46 recorded in a densely sampled stratigraphic sequence spanning Subsequent surveys in the area were carried out by paleontologists 111 47 more than one million years (Alba et al., 2011a). Paleontologist Miquel Crusafont and Josep F. de Villalta during the 1940s (e.g., 112 48 Villalta Comella and Crusafont Pairo, 1941a; Crusafont Pairo, 1944; 113 49 Villalta and Crusafont, 1946). In the loosely-defined locality of 114 50 Can Vila, these authors discovered a hominoid mandibular frag- 115 * 51 Corresponding author. e 116 E-mail address: [email protected] (D.M. Alba). ment with M2 M3 (Villalta Comella and Crusafont Pairo, 1941b, 52 117 53 http://dx.doi.org/10.1016/j.jhevol.2016.09.012 118 54 0047-2484/© 2016 Elsevier Ltd. All rights reserved. 119

Please cite this article in press as: Alba, D.M., et al., Ten years in the dump: An updated review of the Miocene primate-bearing localities from Abocador de Can Mata (NE Iberian Peninsula), Journal of Human Evolution (2016), http://dx.doi.org/10.1016/j.jhevol.2016.09.012 YJHEV2264_proof ■ 5 October 2016 ■ 2/9

2 D.M. Alba et al. / Journal of Human Evolution xxx (2016) 1e9

1 1944), first attributed to Dryopithecus fontani but subsequently Vilar et al., 2011a). Although estimated ages have already been 66 2 used to erect Sivapithecus occidentalis (currently considered a published for some of them (e.g., Casanovas-Vilar et al., 2011a, 67 3 nomen dubium; Moya-Sol a et al., 2004; Alba, 2012), as well as one 2016; Alba, 2012), recent refinements in the calibration of the 68 4 or two additional molar crown fragments (Villalta Comella and geomagnetic polarity time scale (Ogg, 2012; Hilgen et al., 2012) and 69 5 Crusafont Pairo, 1944; Crusafont Pairo, 1958; see discussion in; additional field data have resulted in minor readjustments. The 70 6 Golpe Posse, 1982; Alba et al., 2013). Many years later, Crusafont- termination of fieldwork in 2014 offers a good opportunity to 71 7 Pairo and Golpe-Posse (1973) reported a female lower canine report a comprehensive list of updated estimated ages for ACM 72 8 from Can Mata 1, which was formerly attributed to Hispanopithecus localities, thereby including an accurate dating for all of the iden- 73 9 laietanus (e.g., Golpe Posse, 1993), but most recently assigned to tified primate remains that have been recovered from this area. 74 10 Hominidae indet. (Alba, 2012). 75 11 For many years, Can Vila and Can Mata 1 were the only late 2. Geological background 76 12 Aragonian (late middle to early late Miocene) primate-bearing lo- 77 13 calities from the Penedes sector of the Valles-Pened es Basin (Agustí The middle to late Miocene successions in the area of els Hos- 78 14 et al., 1985; Casanovas-Vilar et al., 2011a). This situation drastically talets de Pierola are located in the Penedes sector of the Valles- 79 15 changed in late 2002 thanks to the beginning of paleontological Penedes Basin. The latter is an elongated and NNE/SSW-oriented 80 16 surveillance, sampling and excavation associated with the removal half-graben, situated between the Pre-littoral and the Littoral 81 17 of thousands of tons of Miocene sediments with heavy machinery Catalan Coastal Ranges, which originated due to the rifting of the 82 18 during the building of a new phase of the Can Mata Landfill (Abo- NW Mediterranean during the Neogene (Cabrera et al., 2004; de 83 19 cador de Can Mata [ACM]; Fig. 1a; Alba et al., 2006, 2009, 2011a). Gibert and Casanovas-Vilar, 2011). Except for some marine and 84 20 This paleontological work, carried out almost uninterruptedly (10.5 transitional sequences from the early to middle Miocene, most of 85 21 years of fieldwork in total) until early 2014, led to the recovery of the basin infill consists of alluvial fan sediments that have provided 86 22 more than 60,000 fossil vertebrate remains. This huge sampling a rich record of early to late Miocene continental vertebrates 87 23 effort enabled the recovery of rare taxa such as primates, leading to (Agustí et al., 1985; Casanovas-Vilar et al., 2011b, 2015). The thick 88 24 the description of three new hominoid genera (Moya-Sol a et al., Miocene alluvial sequences of the ACM and surrounding areas 89 25 2004, 2009a; Alba et al., 2015) and a new pliopithecoid species mostly consist of reddish to brown mudstones, coupled with less 90 26 (Alba et al., 2010). Detailed litho-, bio- and magnetostratigraphic abundant sandstones, breccias and conglomerates, which were 91 27 control further enabled accurate dating of the more than 200 deposited in the distal-to-marginal inter-fan zones of the coa- 92 28 paleontological localities formally defined along the composite lescing alluvial fan systems of Olesa and els Hostalets de Pierola 93 29 stratigraphic sequence of ACM (Moya-Sol a et al., 2009b; Casanovas- (Casanovas-Vilar et al., 2008; Alba et al., 2009, 2011a; Moya-Sol a 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 57 122 58 123 59 124 60 125 61 Figure 1. Abocador de Can Mata (ACM). (a) Map of the main fossiliferous area of els Hostalets de Pierola, indicating the location of the old and new phases of ACM, the nearby 126 62 classical localities of Can Mata, and the magnetostratigraphically sampled sections (in red). (b) Aerial photograph of ACM (courtesy of CESPA Gestion de Residuos, S.A.U.), indicating 127 fi 63 the approximate location of the various sectors in which it was divided during eldwork (in red). (c) Schematic map of the new phase of ACM, indicating the (sub)sector limits. 128 Abbreviations: BCV, Barranc de Can Vila (Can Vila Ravine); BDA, Bassa de Decantacio d'Aigues Pluvials (Settling Pond of Rainwater); BDL, Bassa de Lixiviats (Pond of Leachates); BPA, 64 Base de la Pila d'Acopi (Base of the Stockpile); C1eC9, Cells 1 to 9; CCV; Camí de Can Vila (Can Vila's Trackway); RC, Riera de Claret; VIE, Vial Intern d'Explotacio (Internal Operating 129 65 Road). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.) 130

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1 et al., 2009b). The mudstone-dominated alluvial successions at magnetostratigraphic analysis of six different sections: Barranc de 66 2 ACM show evidence of soil formation at varying intensities, and are Can Vila, Cell 3, Cell 4, and Vial Intern, restricted to the ACM; Camí 67 3 characterized by the widespread presence of early diagenetic car- de Can Vila, further including nearby areas; and Riera de Claret, 68 4 bonates as well as by the alternation of differently colored (pale tan, entirely circumscribed to an area outside the ACM. Moya-Sol a et al. 69 5 redebrown, grayish, or mottled) layers (Alba et al., 2009). These (2009b) depicted only four (instead of six) local polarity magne- 70 6 multicolored sediments are interpreted as the result of alternating tostratigraphic sections because the results for Cell 3, Cell 4 and 71 7 reducing/oxidizing conditions (due to oscillations of the water ta- Camí de Can Vila were fused into a single composite section (see 72 8 ble), with comparatively thin and scarce paleosol horizons being Moya-Sol a et al., 2009b: Fig. 3 and Suppl. Figs. 3e4). In the field, the 73 9 characterized by an intense red coloration. Additional information stratigraphic position of each ACM locality was recorded relative to 74 10 about the geology of the ACM can be found in Alba et al. (2009). its closest lithostratigraphic section(s), being thus correlated on 75 11 lithostratigraphic grounds to the composite magnetostratigraphic 76 12 3. Materials and methods sequence. 77 13 78 14 3.1. Lithostratigraphic correlation 3.3. Definition of the localities 79 15 80 16 A composite lithostratigraphic sequence of ACM was con- Although isolated fossil finds were common during the sur- 81 17 structed on the basis of 38 local sections (47.2 m in average thick- veillance of the excavators' activity, most of the recovered fossils 82 18 ness, but ranging from 7 to 150 m), five of which have associated (>90%) come from formally-defined localities, which were desig- 83 19 paleomagnetic data. Although a simplified correlation of the then nated with a particular name for practical reasons. These formally- 84 20 available lithostratigraphic sections was provided by Alba et al. defined localities are designated by the ACM acronym, separated by 85 21 (2009), the complete lithostratigraphic panel (including correla- a slash from the (sub)sector acronym (Fig. 1b,c), and followed by 86 22 tion of all the local sections) remained unpublished. All the docu- one or two characters denoting the locality (see SOM Table S2 for 87 23 mented lithostratigraphic sections largely overlap with adjacent further details). These localities are restricted areas within a single 88 24 ones and were correlated with one another based on 42 correlation stratigraphic horizon (usually between 0.2 and 1 m in thickness) 89 25 levels (See Supplementary Online Material [SOM] Table S1 and from which fossil remains were collected, being quite heteroge- 90 26 Fig. S1). The same method (although not explicitly described) was neous in lateral extension and number of recovered fossil remains. 91 27 applied by Moya-Sol a et al. (2009b) to the then available sections. Some localities (e.g., ACM/C1-E*) yielded only a single or a few fossil 92 28 The levels used to correlate the local sections with one another remains, but were nonetheless formally defined given the signifi- 93 29 are composed of fine- to very fine-grained sediments (claystones cance of the finds. Other localities are medium to large-sized ac- 94 30 and siltstones), with some sporadic presence of quartzite clasts and cumulations of micro- and/or macrovertebrate remains. Localities 95 31 frequently also with pedogenic carbonate nodules. These levels do that exclusively yielded microvertebrate remains are usually 96 32 not differ noticeably in lithology from other argillaceous and silty restricted to a few square meters, but the number of recovered 97 33 layers from the area, but can be readily distinguished from them remains may range from a few tens to several thousands of speci- 98 34 because of their intense red coloration, which is distinctly darker mens. Localities with large mammal remains are also heteroge- 99 35 than that of most other fine-grained sediments from the ACM neous in extension, ranging from small accumulations of a few 100 36 sequence. These correlation levels are interpreted as paleosols, square meters and less than a hundred recovered remains, to very 101 37 with their intense red coloration denoting well aerated (oxidizing) rich and extensive accumulations of >100 m2 and >5000 macro- 102 38 and lowered water table conditions (Alba et al., 2009). Among the mammal (and often also micromammal) remains. 103 39 correlation levels used by us, only level a stands out due to its 104 40 bluishegrayish coloration, which uniquely along the ACM section 3.4. Estimation of locality age by interpolation 105 41 indicates exceptionally marked reducing and elevated water table 106 42 conditions (which were otherwise generally absent from this An estimated age was calculated for each locality using linear 107 43 depositional environment). Stratigraphic layers consisting of interpolation from subchron boundaries (e.g., Barry et al., 2002; see 108 44 sandstones or conglomerates were discarded as correlation levels also Casanovas-Vilar et al., 2014: Fig. 3), i.e., by relying on the 109 45 at ACM because of their restricted (decametric) lateral continuity locality's correlation with a particular subchron, its stratigraphic 110 46 compared to the above-mentioned claystone and siltsone correla- position within that subchron, and the sedimentation rate 111 47 tion levels (which display a lateral continuity in the order of hun- computed for that subchron, with sedimentation rate being 112 48 dreds of meters within the excavated area). This is exemplified by calculated on the basis of the temporal duration and stratigraphic 113 49 the above-mentioned level a, which is documented in twelve local thickness of each subchron in the composite magnetostratigraphic 114 50 sections and displays a minimum lateral extension of >600 m sequence. For the top subchron at ACM, sedimentation rate was 115 51 within the ACM area. This and other correlation levels were computed based on its actual thickness (taking into account nearby 116 52 repeatedly documented in the successive cells and other structures outcrops outside the ACM), whereas for the bottom one, sedi- 117 53 of the dump as they were consecutively excavated (SOM Table S1), mentation rate was approximated by that of the adjacent subchron. 118 54 thanks to the extensive outcrops generated by the activity of the Due to the intermittent nature of sediment deposition in fluvial/ 119 55 heavy machinery (Fig. 2). alluvial environments, characterized by short episodes of sedi- 120 56 mentation separated by longer hiatuses, computed sedimentation 121 57 3.2. Magnetostratigraphy rates vary depending on the time-scale of observation (Sadler, 1981, 122 58 1999) and interpolated ages are prone to some error due to short- 123 59 The magnetostratigraphic correlation of the composite lithos- term variation in sedimentation rates (Badgley et al., 1986). Barry 124 60 tratigraphic sequence of the ACM with the latest update of the et al.'s (2002) method tries to minimize this problem by relying 125 61 Geomagnetic Polarity Time Scale (GPTS; Ogg, 2012) is based on 366 on sedimentation rates calculated for each subchron, although 126 62 paleomagnetic samples, taken at 1e2 m intervals from the ACM and some error remains due to potentially uneven sedimentation rates 127 63 the adjacent sections (see results reported in Moya-Sol a et al., within each subchron. However, if hiatuses are randomly distrib- 128 64 2009b). In particular, Moya-Sol a et al. (2009b: Suppl. Fig. 1 and uted and short relative to interval duration, age estimates are 129 65 Suppl. Table 1; see also our Fig. 1a) reported the results for the considered reliable (Badgley et al., 1986; Barry et al., 2002). The 130

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1 66 2 67 3 68 4 69 5 70 6 71 7 72 8 73 9 74 10 75 11 76 12 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108

44 Figure 2. Panoramic views of different sectors of ACM while being excavated, illustrating the extensive Miocene outcrops and the correlation levels (denoted by white letters, see 109 45 SOM Table S1) exposed by the activity of heavy machinery, which enabled a reliable lithostratigraphic correlation between the multiple sampled local sections: (a) C5-A in May 110 46 2007; (b) ACM/C8-A in March 2011; (c) C6-A in April 2008. The use of lower or upper case letters to designate the correlation levels is arbitrary. 111 47 112 48 113 49 lateral consistency in the measured thickness of stratigraphic in- and structures of the dump were consecutively excavated toward 114 50 tervals, the lack of sharp stratigraphic contacts and the overall the northwest, over the years progressively younger sediments 115 51 uniformity of sedimentary facies across the study area suggest that were sampled. The large overlap between the extensive outcrops 116 52 steady sedimentation at the time-scale of 100 ka is reasonable. The generated by the heavy machinery enabled us to reliably correlate 117 53 latter assertion was further tested by computing differences in all the various local stratigraphic sections with one another, as well 118 54 stratigraphic thickness between consecutive correlation levels as with the magnetostratigraphic sections already sampled by 119 55 among different subsections in the composite lithostratigraphic Moya-Sol a et al. (2009b), by means of the above-described corre- 120 56 correlation panel, and using their average value multiplied by lation levels (see SOM Table S1), which could be followed across 121 57 average sedimentation rate as a proxy of the average error of the successive cells of the dump. 122 58 age estimates computed for the localities. The local composite stratigraphic sequence of ACM has a 123 59 thickness of 234 m and covers up to seven subchrons (from C5Ar.1r 124 60 4. Results to C5r.2r; Table 1), with an estimated total duration of 1.2 myr (from 125 61 12.58 to 11.38 Ma; Fig. 3). Along this sequence, 235 formally deﬁned 126 62 The strata excavated at ACM consistently displayed a moderate paleontological localities have been discovered, resulting in an 127 63 dip toward the west (Moya-Sol a et al., 2009b). In particular, average of 19.6 localities per 0.1 myr. Only 19 localities (8%) have 128 64 repeated ﬁeld measurements at different outcrops delivered values yielded primate remains, belonging to either pliopithecoids or 129 65 of strike and dip around 175 /14 W. Given that most of the cells hominoids; Table 2; Fig. 3). The estimated ages and stratigraphic Q1 130

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1 Table 1 66 2 Subchrons correlated with the ACM composite section, including their boundaries and duration according to the GMPTS (Ogg, 2012), their stratigraphic position and thickness 67 in the composite section, and their computed within-subchron sedimentation rate. 3 68 4 Subchron Bottom (Ma) Top (Ma) Duration (myr)a Bottom (m) Top (m) Thickness (m) Sedimentation (cm/kyr)a 69 5 C5r.2r 11.592 11.188 0.404 204.2 261.2b 57.0a 14.1b 70 6 C5r.2n 11.657 11.592 0.065 177.1 204.2 27.1 41.7 71 7 C5r.3r 12.049 11.657 0.392 84.1 177.1 93.0 23.7 72 8 C5An.1n 12.174 12.049 0.125 68.6 84.1 15.5 12.4 73 C5An.1r 12.272 12.174 0.098 47.2 68.6 21.4 21.8 9 C5An.2n 12.474 12.272 0.202 15.9 47.2 31.3 15.5 74 10 C5Ar.1r 12.735 12.474 0.261 e 15.9 16 e 75 11 76 a myr ¼ millions of years; kyr ¼ thousands of years. 12 b Only the lower portion of this subchron (30 m) is recorded within the ACM, but the top is recorded in the nearby sequence of Camí de Can Vila. 77 13 78 14 79 15 80 16 81 17 82 18 83 19 84 20 85 21 86 22 87 23 88 24 89 25 90 26 91 27 92 28 93 29 94 30 95 31 96 32 97 33 98 34 99 35 100 36 101 37 102 38 103 39 104 40 105 41 106 42 107 43 108 44 109 45 110 46 111 47 112 48 113 49 114 50 115 51 116 52 117 53 118 54 119 55 120 56 121 Figure 3. Correlation of the composite local magnetostratigraphy of ACM and nearby areas with the Geomagnetic Polarity Time Scale (GPTS; after Ogg, 2012). European Land 57 Mammal Ages (ELMA; boundaries after Hilgen et al., 2012), Mammal Neogene units (MN) and local biozones of the Valles-Pened es Basin (based on Casanovas-Vilar et al., 2016) are 122 58 shown on the left. The stratigraphic positions of the ACM primate-bearing localities (reported in Table 2) are shown to the right on the composite lithostratigraphic column. Note 123 59 that the bottom boundary of the lowermost local biozone is unknown. 124 60 125 61 126 62 positions in the composite stratigraphic sequence of the primate- of all formally deﬁned ACM localities is reported in SOM Table S2. 127 63 bearing localities are reported in Table 2, together with remarks Interpolated ages are subject to two kinds of errors resulting from 128 64 on the recovered remains and their taxonomic assignment. A variations in accumulation rate in both time and space. Variations 129 65 complete list with the stratigraphic position and the estimated ages through time are obvious when sedimentation rates are computed 130

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1 Table 2 66 2 Primate-bearing localities of Abocador de Can Mata, in chronological order from youngest to oldest (top to bottom), including their estimated ages, stratigraphic positions 67 along the synthetic ACM section, and the recorded primate taxa with main descriptive references.a 3 68 4 Locality Age (Ma) Position (m) Taxon Description Referencesb 69 5 ACM/C8-A4c 11.6 194 Pliobates cataloniae Cranium with partial skeleton 1 70 6 ACM/C5-D1 11.6 188 Pliobates cataloniae Maxillary fragment and isolated dental remains 0 71 7 ACM/C5-D1 11.6 188 Dryopithecinae indet. Eroded fragment of distal humeral epiphysis 0 72 8 ACM/C8-B* 11.7 178.5 Dryopithecinae indet. Talus 0 73 ACM/C4-A1c 11.7 166 Pliopithecus canmatensis Male mandible and isolated dental remains 2 9 ACM/C5-A8 11.8 150 Pliopithecus canmatensis Male and female maxillary fragments, and 0, 2, 3 74 10 isolated dental remains 75 11 ACM/C5-C2 11.8 146 Pliopithecus canmatensis Upper deciduous premolar 2 76 12 ACM/C4-Cb 11.8 145 Pliopithecus canmatensis Female (?) mandibular fragments and isolated 2 77 dental remains 13 78 ACM/C5-C4 11.8 133 Pliopithecus canmatensis Partial phalanx 0, 3 14 ACM/C5-C3 11.9 129 Pliopithecus canmatensis Maxillary fragment, male mandible, isolated dental 0, 2, 3 79 15 remains, cranial fragment, humeral and femoral shafts, 80 16 partial metatarsal, and phalangeal fragments 81 17 ACM/C5-C3 11.9 129 Dryopithecinae indet. Partial manual middle phalanx 82 ACM/C3-Ae 11.9 125 Dryopithecus fontani Muzzle 4, 5 18 ACM/C4-Ap 11.9 124 Dryopithecus fontani Male upper canine and two phalangeal fragments 0, 6 83 19 ACM/BCV5 11.9 117 Pliopithecus canmatensis Phalangeal fragment 0 84 20 ACM/C3-Az 11.9 110 cf. Dryopithecus fontani Partial femur 4, 5 85 e 21 ACM/BCV4 11.9 110 cf. Pierolapithecus catalaunicus Mandibular fragment 0 86 ACM/C4-Cpf 12.0 108 Anoiapithecus brevirostris Male upper canine and cheek teeth 0 22 87 ACM/BCV1c 12.0 105 Pierolapithecus catalaunicus Face with partial skeleton 5, 7e11 23 ACM/C3-Ajc 12.0 104 Anoiapithecus brevirostris Male face with mandible 5, 12, 13 88 24 ACM/C3-B2 12.1 83 Pliopithecoidea indet. Upper third molar 14 89 25 ACM/C1-E* 12.4e12.3d 34e46 Anoiapithecus brevirostris Maxillary fragments 13 90 26 a Localities with a designation ending in a number (e.g., ACM/BCV1) also have micromammal remains, whereas asterisks (e.g., ACM/C1-E*) denote isolated primate finds. See 91 27 SOM Table S2 for further details and for the estimated ages of localities without primates. 92 b 28 Legend for references: 0, Material unpublished or only partly published; 1, Alba et al., 2015;2,Alba et al., 2010;3,Alba and Moya-Sola, 2014;4,Moya-Sola et al., 2009b;5, 93 Alba, 2012;6,Alba and Moya-Sol a, 2012b;7,Moya-Sol a et al., 2004;8,Almecija et al., 2009;9,Perez de los Ríos et al., 2012; 10, Hammond et al., 2013; 11, Pina et al., 2014; 12, 29 94 Moya-Sol a et al., 2009a; 13, Alba et al., 2013; 14, Alba et al., 2012. 30 c Type localities. 95 31 d Uncertainties in stratigraphic position preclude a more secure dating. 96 32 e Attributed to Hominidae indet. by Alba (2012), but potentially belonging to Pierolapithecus catalaunicus, based on occlusal and endostructural similarities (Fortuny et al., 97 33 2014 and unpublished data). 98 f Attributed to Hominidae indet. by Alba (2012), but assigned here to Anoiapithecus based on canine morphology (for further details, see Alba and Moya-Sol a, 2012b; Alba 34 et al., 2013). 99 35 100 36 101 37 separately for each subchron (Table 1), which contributes signifi- MN7þ8 (the boundary between these zones being locally esti- 102 38 cantly in minimizing errors in estimated age, as explained in the mated at 12.4 Ma, as defined by the first appearance of Mega- 103 39 Materials and methods. Although there is some variation in sedi- cricetodon ibericus; Casanovas-Vilar et al., 2016). According to the 104 40 mentation rates between subchrons, the resulting average sedi- latest rodent-based local biozonation of the Valles-Pened es Basin 105 41 mentation rate of 19.5 cm/kyr for the ACM sequence is very similar (Casanovas-Vilar et al., 2015, 2016), the ACM sequence spans three 106 42 to the average of 20 cm/kyr computed by Garces et al. (1996) for the subzones of the Hispanomys Assemblage Zone: the lower-most part 107 43 Vallesian sections of the same basin. of the sequence is correlated with the Megacricetodon 108 44 With regard to errors in estimated ages due to spatial variation crusafonti þ Democricetodon larteti Concurrent Range Subzone 109 45 in sedimentation rates, they are minimized by the fact that all the (12.6e12.38 Ma), followed by the M. crusafonti e Democricetodon 110 46 correlated local sections come from a very restricted geographic crusafonti Interval Subzone (12.38e11.87 Ma), and finally the 111 47 area (less than 0.5 km2). This notwithstanding, the correlated local D. crusafonti e Hippotherium Interval Subzone (11.87e11.18 Ma). 112 48 stratigraphic sections do show some minor variations in strati- The upper part of the last, shortly after the middle/late Miocene 113 49 graphic thickness between synchronous pairs of points, which transition at 11.625 Ma (Hilgen et al., 2012), is not recorded at ACM, 114 50 inevitably results in some degree of uncertainty in estimated ages although it would be represented by the classical locality of Can 115 51 interpolated from the composite magnetostratigraphic correlation. Mata 1 (ca. 11.2e11.1 Ma; Casanovas-Vilar et al., 2011a). The Dry- 116 52 The average error in estimated ages due to this factor can be opithecus molar found by Guerín in the 1920s is also probably 117 53 approximated based on differences in stratigraphic thickness be- younger than the whole ACM sequence, whereas the outcrops of 118 54 tween consecutive correlation levels. We computed these differ- the classical locality of Can Vila are most likely equivalent to sector 119 55 ences for 221 pairs of consecutive correlation levels, resulting in an ACM/BCV, with an estimated age of ca. 12.0 Ma (Alba et al., 2013). 120 56 average stratigraphic distance of 5.3 m (range 1e12 m), which on 121 57 the basis of the average sedimentation rate computed for the whole 5. Discussion and conclusions 122 58 composite sequence would represent an average error in estimated 123 59 age of ca. 27 kyr (range 5e62 kyr). Even if this figure should be The primate succession at ACM is summarized in Figure 4. The 124 60 increased to some extent because of intrasubchron temporal vari- oldest primate specimen recorded at ACM corresponds to maxillary 125 61 ation in sedimentation rate, it strongly suggests that our estimated remains of Anoiapithecus brevirostris, with an estimated age of 126 62 ages are reliable at least to the nearest 0.1 Ma. 12.4e12.3 Ma, followed by an indeterminate pliopithecoid at 127 63 When biostratigraphic data are also taken into account, it 12.1 Ma. Slightly later, most of the remaining great ape finds are 128 64 emerges that the whole ACM sequence is late Aragonian in age, its clumped in a short time interval between 12.0 and 11.9 Ma, 129 65 lower-most portion being correlated to MN6 and the rest to including Anoiapithecus (12.0 Ma), Pierolapithecus (12.0 Ma and, 130

D.M. Alba et al. / Journal of Human Evolution xxx (2016) 1e9 7

1 66

)

2 t 67

M r Stratigraphic ranges

(

3 r 68

4 Local of ACM primates 69

Epoch ELMA

biozone

A 5 Subchron 70 6 1n 71 7 2r-1 72 8 73 2r 9 r 74 10 5 75

atensis 11 LATE M. 2n 76

Hippotherium 12 D. crusafonti - 77 13 78

3r Pliobates cataloniae

14 MN7+8 79 15 Assembage Zone 80 12 Dryopithecinae indet. 16 1n 81

Pliopithecus canm

n 17 Dryopithecus fontani 82 1r

18 D. crusafonti 83

late Aragonian

M. crusafonti

Pierolapithecus catalaunicus

Hispanomys

19 2n Pliopithecoidea indet. 84

C 20 85

Anoiapithecus brevirostris 21 C5Ar.1r 86

D. larteti

22 MN6 87

MIDDLE MIOCENE

M. crusaf. 23 + 88 24 Figure 4. Approximate stratigraphic ranges of the various primate taxa recovered at ACM based on their first and last occurrences at ACM (in gray), with locality occurrences of each 89 25 taxon superimposed (in black). Occurrences have been depicted based on interpolated ages reported for primate-bearing localities in SOM Table S2 to the nearest 0.01 Ma, but 90 26 ranges further take into account an age uncertainty of ±0.05 Ma for first and last occurrences of each taxon. It is uncertain whether the indeterminate dryopithecine finds, depicted 91 27 here within a single range, represent the same taxon or not (they might belong to one or more of the species recorded at ACM). The light gray background signals the time span 92 represented by the ACM composite sequence. 28 93 29 94 30 95 31 tentatively, 11.9 Ma), and Dryopithecus (12.0e11.9 Ma). Only the last pliopithecoid and hominoid remains come from the very same 96 32 taxon and fragmentary indeterminate dryopithecine remains stratigraphic horizon. Here we show that, through the ACM 97 33 minimally overlap with the lower stratigraphic range of the plio- sequence, pliopithecoids and hominoids generally do not co-occur 98 34 pithecoid Pliopithecus canmatensis, which is recorded in seven lo- within a single locality (with a single exception), in spite of close 99 35 calities that span from 11.9 to 11.7 Ma. Subsequently, two isolated similarities in the geological age of the material. This resembles the 100 36 dryopithecine postcranial bones are recorded at ca. 11.7e11.6 Ma, case previously documented for the complex of karstic sites from La 101 37 whereas the stem ape Pliobates cataloniae is the youngest primate Grive-Saint-Alban (France), where two pliopithecoids and a hom- 102 38 from the ACM sequence, being recorded in two localities (one un- inoid are recorded from different fissure fillings (Mein and 103 39 published), both with an age of 11.6 Ma. Co-occurrence of two Ginsburg, 2002; Perez de los Ríos et al., 2013). In fact, despite the 104 40 different primate taxa within a single locality has been recorded high number of primate taxa recorded at ACM (at least, by Euro- 105 41 only twice, based on very fragmentary unpublished dryopithecine pean Miocene standards), two different species (a large-bodied 106 42 remains: a partial middle phalanx from ACM/C5-C3 (11.9 Ma), too dryopithecine and a small-bodied catarrhine, either Pliopithecus 107 43 large to belong to P. canmatensis; and a distal humeral fragment or Pliobates) co-occur only twice. Some authors have previously 108 44 from ACM/C5-D1 (11.6 Ma), too large and derived in morphology to argued that the infrequent co-occurrence between hominoids and 109 45 belong to P. cataloniae. The original stratigraphic ranges of the pliopithecoids throughout the European Miocene might be, at least 110 46 recorded primate species, in all probability, must have been longer in part, a sampling artifact related to the rarity of these primates 111 47 than recorded (about an order of magnitude, judging from pub- (Andrews et al., 1996). Alternatively, the rare co-occurrence be- 112 48 lished average species durations for fossil mammals; Vrba and tween these groups at ACM and elsewhere in Europe might be 113 49 DeGusta, 2004; Alroy, 2009), and it is thus most likely that all of related to differences in the ecological preferences of each group, 114 50 them overlapped in time. However, it is noteworthy that pliopi- coupled with changing paleoenvironmental conditions through 115 51 thecoids and hominoids, despite overlapping in stratigraphic range time. According to the principle of competitive exclusion, these 116 52 at ACM (Fig. 4), only rarely co-occur at the same locality. Therefore, groups might have had different ecological preferences or occupied 117 53 it is uncertain whether these two groups habitually co-existed in slightly different ecological niches, thereby only coexisting under 118 54 both time and space, i.e., whether they frequently inhabited the particular environmental conditions (Andrews et al., 1996). Since 119 55 same type of habitat. the diet inferred for P. canmatensis overlaps with the dietary vari- 120 56 Throughout the European Miocene, pliopithecoids and homi- ation displayed by ACM hominoids as a whole (DeMiguel et al., 121 57 noids co-occur at only a few sites (Begun, 1989; Andrews et al., 2013, 2014; Alba et al., 2015), potential ecological differences be- 122 58 1996; Harrison et al., 2002; Eronen and Rook, 2004; Begun et al., tween these groups might be more likely related to paleobiological 123 59 2010; Sukselainen et al., 2015), such as Castell de Barbera in the aspects different from diet (e.g., locomotion, life history). Most 124 60 Valles-Pened es Basin (Alba et al., 2011b; Almecija et al., 2012; Alba recently, Sukselainen et al. (2015) concluded that pliopithecoids 125 61 and Moya-Sol a, 2012a), or Rudabanya in Hungary (Ungar, 2005; generally inhabited more humid environments than hominoids. 126 62 Begun et al., 2010). However, the co-occurrence of pliopithecoids These authors further tentatively suggested that localities with 127 63 and hominoids within a single stratigraphic horizon can be proven both taxa were even more humid, although these differences were 128 64 only for Rudabanya (Begun et al., 2010), whereas at most not statistically significant (which is not surprising, given the small 129 65 other sites, such as Castell de Barbera, it is unknown whether number of localities where both groups have been recorded). 130

8 D.M. Alba et al. / Journal of Human Evolution xxx (2016) 1e9

1 In order to test the two hypotheses mentioned above at ACM, References 66 2 detailed quantitative analyses (outside the scope of this paper) 67 3 would be required, including the analysis of taphonomic data (such Agustí, J., Cabrera, L., Moya-Sola, S., 1985. Sinopsis estratigrafica del Neogeno de la 68 fosa del Valles-Pened es. Paleontol. Evol. 18, 57e81. 4 as the number of specimens of each taxon recovered from each Alba, D.M., 2012. Fossil apes from the Valles-Pened es Basin. Evol. Anthropol. 21, 69 5 locality) to evaluate sampling biases. Given the heterogeneous 254e269. 70 6 nature of the formally-defined ACM localities, the role of sampling Alba, D.M., Moya-Sol a, S., 2012a. A new pliopithecid genus (Primates: Pliopithe- 71 7 coidea) from Castell de Barbera (Valles-Penedes Basin, Catalonia, Spain). Am. J. 72 biases (due to differences in collecting techniques, volume of Phys. Anthropol. 147, 88e112. 8 sediment excavated and/or screen-washed, and richness of the Alba, D.M., Moya-Sol a, S., 2012b. On the identity of a hominoid male upper canine 73 9 accumulations) cannot be ignored when comparing the taxa from the Valles-Pened es Basin figured by Pickford (2012c). Estudios Geol. 68, 74 e 10 recovered from each locality. To some extent, these biases may be 149 153. 75 Alba, D.M., Moya-Sol a, S., 2014. New fossil remains of Pliopithecus canmatensis from 11 minimized by combining localities into stratigraphic bins of equal Abocador de Can Mata, and their implications for the taxonomic validity and 76 12 (0.1 myr) duration. However, sampling effort appears more intense phylogenetic position of Epipliopithecus (Primates, Pliopithecidae). Am. J. Phys. 77 13 toward the upper part of the ACM sequence (particularly between Anthropol. 153 (S58), 64. 78 Alba, D.M., Moya-Sol a, S., Casanovas-Vilar, I., Galindo, J., Robles, J.M., Rotgers, C., 14 12.0 and 11.7 Ma and, especially, at 11.6 Ma)dat least, as roughly Furio, M., Angelone, C., Kohler,€ M., Garces, M., Cabrera, L., Almecija, S., 79 15 approximated by the number of formally defined localities per Obrado, P., 2006. Los vertebrados fosiles del Abocador de Can Mata (els Hos- 80 ~ 16 0.1 myr stratigraphic bin (see Table 2)dso that a quantitative talets de Pierola, l'Anoia, Cataluna), una sucesion de localidades del Arago- 81 niense superior (MN6 y MN7þ8) de la cuenca del Valles-Pened es. Campanas~ 17 correction for sampling biases will, nevertheless, be required. Un- 2002e2003, 2004 y 2005. Estudios Geol. 62, 295e312. 82 18 fortunately, these analyses will not be possible until a much larger Alba, D.M., Robles, J.M., Rotgers, C., Casanovas-Vilar, I., Galindo, J., Moya-Sol a, S., 83 19 proportion of ACM fossils are prepared (in the case of large verte- Garces, M., Cabrera, L., Furio, M., Carmona, R., Berto Mengual, J.V., 2009. Middle 84 Miocene vertebrate localities from Abocador de Can Mata (els Hostalets de 20 brates) or sorted out from the screen-washing residue (regarding Pierola, Valles-Pened es Basin, Catalonia, Spain): An update after the 85 21 small vertebrates). In the meantime, paleoecological analyses based 2006e2008 field campaigns. Paleolusitana 1, 59e73. 86 22 on the ACM fauna (such as the analyses of tooth enamel stable Alba, D.M., Moya-Sola, S., Malgosa, A., Casanovas-Vilar, I., Robles, J.M., Almecija, S., 87 23 Galindo, J., Rotgers, C., Berto Mengual, J.V., 2010. A new species of Pliopithecus 88 isotopes, or mesowear analysis of ungulate teeth) may provide Gervais, 1849 (Primates: Pliopithecidae) from the Middle Miocene (MN8) of 24 helpful insights about whether ACM habitats did change signifi- Abocador de Can Mata (els Hostalets de Pierola, Catalonia, Spain). Am. J. Phys. 89 25 cantly with the passing of time and, if so, whether the inferred Anthropol. 141, 52e75. 90 26 paleoenvironmental changes are somewhat correlated with the Alba, D.M., Casanovas-Vilar, I., Robles, J.M., Moya-Sola, S., 2011a. Parada 3. El Arago- 91 niense superior y la transicion con el Vallesiense: Can Mata y la exposicion pale- 27 faunal turnovers observed among the recovered fauna (including ontologica de els Hostalets de Pierola. Paleontol. Evol. Mem. Esp. Núm. 6, 95e109. 92 28 primates). Therefore, the set of primate-bearing localities from Alba, D.M., Moya-Sol a, S., Almecija, S., 2011b. A partial hominoid humerus from the 93 29 ACM, framed within a densely-sampled sequence spanning about middle Miocene of Castell de Barbera (Valles-Penedes Basin, Catalonia, Spain). 94 Am. J. Phys. Anthropol. 144, 365e381. 30 one million years and restricted to a small geographic area, offers Alba, D.M., Moya-Sol a, S., Robles, J.M., Galindo, J., 2012. Brief Communication: The 95 31 the prospect of shedding new light, in years to come, on the most oldest pliopithecid record in the Iberian Peninsula based on new material from 96 e 32 likely cause(s) underpinning the infrequent co-occurrence between the Valles-Penedes Basin. Am. J. Phys. Anthropol. 147, 135 140. 97 Alba, D.M., Fortuny, J., Perez de los Ríos, M., Zanolli, C., Almecija, S., Casanovas- 33 large-bodied apes and small catarrhines in the European Miocene. Vilar, I., Robles, J.M., Moya-Sol a, S., 2013. New dental remains of Anoiapithecus 98 34 In this regard, the estimated ages reported here for all the fossil and the first appearance datum of hominoids in the Iberian Peninsula. J. Hum. 99 Evol. 65, 573e584. 35 vertebrate localities from the ACM stratigraphic sequence represent 100 fi Alba, D.M., Almecija, S., DeMiguel, D., Fortuny, J., Perez de los Ríos, M., Pina, M., 36 an important rst step towards the resolution of this long-standing Robles, J.M., Moya-Sol a, S., 2015. Miocene small-bodied ape from Eurasia sheds 101 37 conundrum. light on hominoid evolution. Science 350, aab2625. 102 38 Almecija, S., Alba, D.M., Moya-Sol a, S., 2009. Pierolapithecus and the functional 103 morphology of Miocene ape hand phalanges: paleobiological and evolutionary 104 39 Acknowledgments implications. J. Hum. Evol. 57, 284e297. 40 Almecija, S., Alba, D.M., Moya-Sol a, S., 2012. The thumb of Miocene apes: new in- 105 41 Research reported in this work has been funded by the Spanish sights from Castell de Barbera (Catalonia, Spain). Am. J. Phys. Anthropol. 148, 106 e 42 Ministerio de Economía y Competitividad (CGL2014-54373-P, 436 450. 107 Alroy, J., 2009. Speciation and extinction in the fossil record of North American 43 CGL2014-55900-P and RYC-2013-12470 to ICV) and the Departa- mammals. In: Butlin, R.K., Bridle, J.R., Schluter, D. (Eds.), Speciation and Patterns 108 44 ment d’Innovacio, Universitats i Empresa of the Generalitat de of Diversity. Cambridge University Press, Cambridge, pp. 301e323. 109 45 Catalunya (2014 SGR 416, GRC). In turn, fieldwork at ACM has been Andrews, P., Harrison, T., Delson, E., Bernor, R.L., Martin, L., 1996. Distribution and 110 biochronology of European and Southwest Asian Miocene catarrhines. In: 46 defrayed by CESPA G.R., S.A.U., whereas preparation of the recov- Bernor, R.L., Fahlbusch, V., Mittmann, H.-W. (Eds.), The Evolution of Western 111 47 ered fossils has been partly financed by the Departament de Cultura Eurasian Neogene Mammal Faunas. Columbia University Press, New York, 112 e 48 of the Generalitat de Catalunya (2014/100609). We gratefully thank pp. 168 207. 113 e Badgley, C., Tauxe, L., Bookstein, F.L., 1986. Estimating the error of age interpolation 49 Jordi Galindo for his pioneering geological work (2002 2006) at in sedimentary rocks. Nature 319, 139e141. 114 50 ACM, and Salvador Moya-Sol a for discussions about the ACM pri- Barry, J.C., Morgan, M.E., Flynn, L.L., Pilbeam, D., Behrensmeyer, A.K., Mahmood 115 51 mates. We further acknowledge the continued collaboration and Raza, S., Khan, I.A., Badgley, C., Hicks, J., Kelley, J., 2002. Faunal and environ- 116 ’ mental change in the late Miocene Siwaliks of Northern Pakistan. Paleobiology 52 support of the Servei d Arqueologia i Paleontologia of the Gen- 28 (sp. 3), 1e71. 117 53 eralitat de Catalunya during the successive paleontological in- Bataller Calatayud, J.R., 1938. Els Ratadors Fossils de Catalunya. Impremta de la Casa 118 54 terventions performed at ACM. Finally, we are grateful to the two d'Assistencia President Macia, Barcelona. 119 55 Begun, D.R., 1989. A large pliopithecine molar from Germany and some notes on the 120 anonymous reviewers, whose constructive criticisms and sugges- Pliopithecinae. Folia Primatol. 52, 156e166. 56 Q2 tions helped us to improve a previous version of this paper. Begun, D., Nargolwalla, M.C., Kordos, L., 2010. New catarrhine fossils from 121 57 Rudabanya (Hungary): Evidence for sympatric primates in a late Miocene 122 58 Supplementary Online Material swamp forest. Am. J. Phys. Anthropol. 141 (S50), 64. 123 Cabrera, L., Roca, E., Garces, M., Porta, J. de, 2004. 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