This is the accepted version of the article: Alba, David M.; Rodríguez-Hidalgo, Antonio; Aouraghe, Hassan; [et al.]. «New macaque fossil remains from Morocco». Journal of human evolution, Vol. 153 (April 2021). DOI 10.1016/j.jhevol.2021.102951 This version is available at https://ddd.uab.cat/record/236871 under the terms of the license New macaque fossil remains from Morocco David M. Alba a,*, Antonio Rodríguez-Hidalgo b,c,d, Hassan Aouraghe e, Jan van der Made f, Aïcha Oujaa g, Hamid Haddoumi e, Palmira Saladié c,h,i, Al Mahdi Aissa e, Juan Marín c, Mourad Farkouch c,e, Carlos Lorenzo h,c, Said Bengamra e, Eric Delson j,k,l,m,a, M. Gema Chacón c,h,n, Robert Sala-Ramos c,h a Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193, Cerdanyola del Vallès, Barcelona, Spain b Departamento de Prehistoria, Historia Antigua y Arqueología, Universidad Complutense de Madrid, Facultad de Geografía e Historia, C/ Prof. Aranguren s/n, 28040, Madrid, Spain c Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edifici W3), Tarragona, 43007, Spain d Instituto de Evolución en África (IDEA, Madrid), C/ Covarrubias 36, 28010, Madrid, Spain e Faculté des Sciences, Département de Géologie, Université Mohamed Premier, BV Mohammed VI, Quartier Al Qods, 60000, Oujda, Morocco f Consejo Superior de Investigaciones Científicas (CSIC), Museo Nacional de Ciencias Naturales, Departamento de Paleobiología, C/ José Gutiérrez Abascal, 2, 28006, Madrid, Spain g Institut National des Sciences de l’Archéologie et du Patrimoine (INSAP), Av. Allal El-Fassi, Hay Riad, 6828, Rabat, Morocco h Universitat Rovira i Virgili, Departament d’Història i Història de l’Art, Avinguda de Catalunya 35, 43002 Tarragona, Spain 1 i Unidad Asociada al CSIC, Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales, C/ José Gutiérrez Abascal, 2, 28006, Madrid, Spain j Department of Anthropology, Lehman College of the City University of New York, 250 Bedford Park Boulevard West, Bronx, NY, 10468, USA k Department of Vertebrate Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY, 10024, USA l PhD Programs in Anthropology and Earth & Environmental Sciences, The Graduate Center of the City University of New York, 365 Fifth Avenue, New York, NY, 10016, USA m New York Consortium in Evolutionary Primatology, New York, NY, USA n UMR7194 Histoire naturelle de l’Homme préhistorique (HNHP), Museum National d'Histoire Naturelle (MNHN), CNRS, Université Perpignan Via Domitia, Alliance Sorbonne Université – Musée de l’Homme, Place du Trocadéro 17, 75016 Paris, France Corresponding author. Email: david.alba@icp.cat (D.M. Alba) Acknowledgments We thank the Jerada Government and Local Authorities of Aïn Beni Mathar and Guefaït for local permits to develop geological, archaeological and paleontological fieldworks in the region. This work has been funded by Palarq Foundation, Spanish Ministry of Culture and Sport (42-T002018N0000042853 and 170-T002019N0000038589), Direction of Cultural Heritage (Ministry of Culture and Communication, Morocco), Faculty of Sciences (Mohamed 1r University of Oujda, Morocco), INSAP (Institut National des Sciences de l'Archéologie et 2 du Patrimoine), Agencia Estatal de Investigación (Spanish Ministry of Science, Innovation and Universities: CGL2016-76431-P, CGL2016-80975-P, CGL2016-80000-P, CGL2017-82654- P, PGC2018-095489-B-I00 and PGC2018-093925-B-C31, AEI/FEDER-UE), and Research Group Support of the Generalitat de Catalunya (2017 SGR 116, 2017 SGR 836, and 2017 SGR 859). D.M.A., R.S-R, M.G.CH., P.S., C.L. and E.D.’s research is funded by CERCA Programme/Generalitat de Catalunya. A.M.A and M.F. are beneficiaries of a fellowship from the Erasmus Mundus Program to do the Master in Quaternary and Prehistory at the Universitat Rovia i Virgili (Tarragona, Spain). A.R.-H. is the beneficiary of a postdoctoral scholarship from the MICINN, Subprograma Juan de la Cierva (IJC-037447-I) and member of the consolidated research group 2017 SGR 1040 of the Generalitat de Catalunya. The Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA) has received financial support from the Spanish Ministry of Science and Innovation through the “María de Maeztu” program for Units of Excellence (CEX2019-000945-M). The µCT images were obtained in the Laboratory of Microscopy of the CENIEH-ICTS (Spain) in collaboration with CENIEH staff, and were processed by ICP technician Alejandro Serrano. We thank Eileen Westwig (American Museum of Natural History, New York), Elisabetta Cioppi (Museo di Storia Naturale, Firenze), and Lorenzo Rook (Università di Firenze) for access to comparative material; Josep Fortuny for supervising the segmentation of the CTs; Doug Boyer for providing MorphoSource DOIs; Steve Frost, Chris Gilbert, Florent Détroit and Salvador Moyà-Solà for discussion on the fossil material; and Alessandro Urciuoli for varied assitance. We also thank all the participants in the excavation of the Guefaït site, and the Editor-in- Chief (Andrea Taylor), the Associate Editor, and three anonymous reviewers for suggestions that helped to improve a previous version of this paper. 3 1 Short Communications 2 New macaque fossil remains from Morocco 3 4 Keywords: Old World monkeys; Cercopithecoidea; Papionini; Macaca; Guefaït; North Africa 5 6 1. Introduction 7 1.1. Cercopithecoids from Late Miocene and Plio-Pleistocene North Africa 8 The Late Miocene and Plio-Pleistocene record of cercopithecoids from North Africa is 9 quite meager as compared with that from tropical and southern Africa (Delson, 1973, 1974, 10 1975, 1980; Szalay and Delson, 1979; Geraads, 1987; Benefit et al., 2008; Jablonski, 2002; 11 Jablonski and Frost, 2010; Roos et al., 2019; these references are relevant to all occurrences 12 in this paragraph and the next and will not be repeated). During the Late Miocene, cf. 13 Macaca sp. and ‘Colobus’ flandrini are recorded from Menacer (=Marceau; ~7.0–5.8 Ma), 14 Algeria (Arambourg, 1959). Slightly thereafter, Macaca libyca and the colobine Libypithecus 15 markgrafi co-occur at Wadi Natrun (~6.2–5.0 Ma), Egypt (Stromer, 1913, 1920), and possibly 16 also at As Sahabi (~6.3–5.3 Ma), Libya (Meikle, 1987; Jablonski, 2002; but see Benefit et al., 17 2008 regarding uncertainty in such attributions). 18 Macaca sp. is also recorded, together with Theropithecus ‘atlanticus’1, in the earliest 19 Pleistocene of Ahl al Oughlam (~2.5 Ma), Morocco (Raynal et al., 1990; Geraads et al., 1998; 20 Alemseged and Geraads, 1998; Delson et al., 2000), and is probably present as well in the 21 Pliocene site of Garaet Ichkeul (~3 Ma), Tunisia (e.g., Szalay and Delson, 1979; Delson, 1993; 22 Fooden, 2007; but see Geraads, 1987; Alemseged and Geraads, 1998), while Macaca aff. 1 Although T. atlanticus was recognized as a distinct species by Alemseged and Geraads (1998), we do not recognize it here pending nomenclatural action in process by E.D. and D.M.A. and colleagues relating to its priority vis-à-vis subspecies of T. oswaldi. 1 23 sylvanus has been identified from the Pliocene of Aïn Brimba (~3 Ma), Tunisia (Arambourg 24 and Coque, 1959; Arambourg, 1979; Szalay and Delson, 1979; van den Brink, 1981; Fooden, 25 2007). Subsequently, Macaca sylvanus is recorded from various Middle and Late Pleistocene 26 sites in Algeria (Pomel, 1892, 1896; Fooden, 2007). In turn, T. ‘atlanticus’ has also been 27 documented from Aïn Jourdel in Algeria (~2.5 Ma; Thomas, 1884; Delson, 1993), while 28 Theropithecus oswaldi has been recovered from Tighennif (=Ternifine; ~1.0 Ma; Sahnouni 29 and van der Made, 2009) in Algeria (Jolly, 1972; Delson and Hoffstetter, 1993) and from 30 Thomas Quarries I and III (=Oulad Hamida 1 Quarry; ~700–500 ka; Raynal et al., 2010; 31 Rhodes et al., 2006; Daujeard et al., 2020) in Morocco (Geraads, 1980, 1987; Delson, 1993). 32 33 1.2. The site of Guefaït 34 An ongoing interdisciplinary Spanish-Moroccan project focused on the Aïn Béni- 35 Mathar/Guefaït region, in the northern part of the High Plateaus (Jerada Province, Eastern 36 Morocco), started in 2006 with the aim of mapping the archeological sites and establishing 37 their geochronological and paleontological context (Aouraghe et al., 2016; Chacón et al., 38 2016). Systematic surveys led to the discovery of Pleistocene archeological sites (Sala et al., 39 2016) and some paleontological localities ranging in age from the Late Miocene (Blain et al., 40 2013) to the Plio-Pleistocene (Piñero et al., 2019). The excavation in 2017–2019 of a 28 m2 41 surface trench in the locality of Guefaït-4.2 (Supplementary Online Material [SOM] Fig. S1) 42 has yielded 3269 macrovertebrate remains (Sala et al., 2017; Aouraghe et al., 2019a, 2019b; 43 Piñero et al., 2019; authors’ unpublished data), including a few cercopithecid teeth. Here we 44 describe these dental remains to provide a taxonomic assignment and discuss their 45 paleoenvironmental, paleobiogeographic, and biochronological implications. 2 46 The fossil site of Guefaït-4 (Aïn Béni Mathar-Guefaït Basin) is located at the base of Unit 2 47 of the Dhar Iroumyane (=Dhar Irroumine) stratigraphic section (Aouraghe et al., 2019a). The 48 fossiliferous layer of Guefaït-4, which consists of clays and marls with nodular calcretes, has 49 delivered a diverse faunal assemblage of vertebrates at the 4.2 locality trench (reptiles, 50 amphibians, and both small and large mammals; Hajji, 2010; Agustí et al., 2017; Aouraghe et 51 al., 2019a, 2019b; Piñero et al., 2020). Paleomagnetic analysis of the series containing 52 Guefaït-4.2 is currently underway. Although some preliminary results have been presented 53 (Alvarez Posada et al., 2015), an interpolated age for Guefaït-4.2 is not available yet. The 54 site was initially correlated broadly to the earliest Pleistocene (Agustí et al., 2016). The 55 absence of Equus (despite the abundance of hipparionin equid remains; Aouraghe et al., 56 2019a, 2019b) indicates an age older than ~2.4 Ma (Sahnouni et al., 2018), while the murid 57 assemblage further supports an age close to the 2.58 Ma Pliocene/Pleistocene boundary 58 (Agustí et al., 2017; Piñero et al., 2020). 59 60 2. Materials and methods 61 The described material includes 6 teeth and tooth fragments collected in 2018–2019 62 from Guefaït-4.2 (Table 1). The specimens are currently on loan to the Institut Català de 63 Paleoecologia Humana i Evolució Social in Tarragona, Spain but will be permanenty housed 64 in the Faculty of Science, Mohammed I University in Oujda, Morocco. 3D virtual models are 65 available from MorphoSource.org (SOM Files S1–S6; see methods in SOM S1). 66 Standard dental measurements of mesiodistal crown length (MD), buccolingual crown 67 breadth (BL), and labial crown height (H; only for the incisor) were taken to the nearest 0.1 68 mm by D.M.A. For the molars, BL was taken both at the mesial (BLm) and distal (BLd) 3 69 lophids. A breadth/length index (BLI, in %) was also computed (using BLm for the molars). 70 Body mass (BM) was estimated following Delson et al. (2000)—see SOM S2 for details. 71 Extant and fossil macaques as well as Parapapio (cf.) ado (this concept includes both P. 72 ado and P. cf. ado), ‘Parapapio’ lothagamensis, and Pliopapio alemui were included in the 73 comparative sample2. Measurements were taken from the PRIMO (PRImate Morphometrics 74 Online) database (http://primo.nycep.org) or the literature (references in SOM Table S1), or 75 taken by D.M.A. Bivariate plots of BL vs. MD and box-and-whisker plots were used to 76 visually compare dental size and proportions. Differences among some taxa from the 77 comparative sample were tested using analysis of variance (ANOVA) and Tukey’s pairwise 78 post hoc tests, while the studied specimens were compared to these taxa using z-scores (z = 79 [measurement – mean] / SD). Statistical computations and plots were performed with PAST 80 v. 4.02 for Mac (Hammer et al., 2001; 81 https://www.nhm.uio.no/english/research/infrastructure/past/). 82 83 3. Results 84 3.1. Description 85 Upper central incisor The left I1 (Fig. 1a; see Table 1 for measurements) preserves the crown 86 and a small portion of the root. The incisal edge is sharp and only slightly worn, and 87 although some enamel chips are missing from the distal and mesial aspects, MD can be 88 estimated and lingual morphology adequately ascertained. The crown is only slightly longer 89 mesiodistally than buccolingually broad, and much taller than mesiodistally long. In 90 labial/lingual views it displays a subrectangular contour (with subparallel mesial and distal 2 Other species of Parapapio, such as Parapapio jonesi and Parapapio broomi, were not included because the described molars are clearly smaller and do not overlap the range of variation of these species. 4 91 edges that converge toward the base), which is slightly asymmetrical (somewhat tilted 92 mesially). The incisal edge is horizontal except on the distal-most portion. The crown 93 progressively becomes labiolingually wider rootward. The root displays a subtriangular cross 94 section close to the cervix. The latter is V-shaped mesially and distally, and lingually it 95 extends somewhat further rootward than labially. The labial crown aspect is smooth and 96 uniformly convex, whereas the lingual side is slightly excavated and displays a distinct 97 median sulcus (flanked by two subparallel grooves) that separates the steep mesiolingual 98 crown portion from the more bulging distolingual one. The damaged distal ridge and the 99 apparently thicker mesial ridge converge basally into a narrow lingual cingulum located 100 above a protruding basal bulge. 101 Lower molars The lower molars are represented by several molar crown and germ 102 fragments plus two almost complete lower molars (see measurements in Table 1). The right 103 M1 fragment (Fig. 1b), which preserves 7.4 mm of the buccal wall, is smaller than, but shows 104 a similar morphology to, the complete M2 (Fig. 1e; see below), except for displaying a tiny 105 cuspulid-like enamel thickening in the median buccal cleft. The right mesial germ fragment 106 (Fig. 1c) preserves most of the metalophid (up to the metaconid) and, based on size, 107 probably belongs to an M2 or M3. The right distal germ fragment (Fig. 1d) preserves the 108 whole hypolophid and, given its size and the lack of a hypoconulid, must belong to an M2. 109 Both germ fragments might belong to the same tooth (although this cannot be confirmed 110 based on spatial association; Table 1), but must belong to a different individual than the 111 complete M2 and M3. 112 The right M2 (Fig. 1e) preserves the whole crown (which is moderately worn, with 113 dentine exposure at the protoconid apex) and the obliquely oriented distal root. The 114 occlusal contour is subrectangular (longer than wide, and slightly wider across the 5 115 metalophid). It displays a bilophodont occlusal pattern with a moderately deep median 116 lingual notch (down to slightly below crown midheight, far from the cervix) and some buccal 117 flare. The four main cuspids are quite bulbous, and the lingual ones are slightly more mesial 118 than the corresponding buccal cuspids. The mesial fovea is restricted, with a short 119 preprotocristid, a slightly protruding mesial marginal ridge, and a faint mesial buccal cleft. 120 The distal fovea is shorter than the trigonid basin but more spacious than the mesial fovea 121 and distally enclosed by a thicker and more protruding semicircular marginal ridge. The 122 median buccal cleft is not very wide and lacks secondary cuspulids. 123 The left M3 (Fig. 1f) preserves most of the crown (except for the missing protoconid 124 apex), which is broken at the cervix level and presents only slight wear. This molar displays 125 the same occlusal pattern as the M2, with the following differences: (1) the lingual cuspids 126 are somewhat more buccolingually compressed; (2) the median lingual notch is shallower 127 (ending above crown midheight); (3) the median buccal cleft is more restricted (narrower 128 and shallower); (4) the crown is relatively narrower, and although crown breadth at the 129 hypolophid is minimally narrower than at the metalophid, it displays a distally tapering oval 130 contour with a well-developed third lobe; and (5) the distal lobe bears a buccally-located 131 large hypoconulid (subequal in size to the remaining main cuspids and aligned with the 132 remaining buccal cuspids), as well as a smaller tuberculum sextum between the entoconid 133 and the hypoconulid (more centrally located than the two lingual cuspids). The hypoconulid 134 lobe is smaller than is usual in macaques and other papionins, although this shape is not 135 unique; the four main cuspids seem ‘normal’, with the distal portion of the tooth 136 compressed mesiodistally. 137 138 3.2. Morphological comparisons 6 139 Upper central incisor Cercopithecoid upper incisors are not particularly diagnostic. 140 Nonetheless, the I1 from Guefaït-4.2 differs from that of colobines by being taller than long 141 and by displaying a distinct median sulcus on the lingual side, which is characteristic of 142 papionins (Delson, 1973; Swindler, 2002). The continuous lingual cingulum, lacking a 143 rootward oriented excavated pit (as is typical of Papio; Delson, 1973), further agrees well 144 with the macaque I1 morphology (SOM Fig. S2), while the rootward extension of the labial 145 cervix appears less marked than in Lophocebus sp. from Koobi Fora, Kenya (1.87–1.56 Ma; 146 Jablonski et al., 2008: Figs. IV.18 and IV.20). 147 Lower molars Although the faint mesial buccal cleft (so that the mesial ridge does not 148 constitute a distinct ledge-like protrusion) is somewhat reminiscent of colobines, other 149 features of the Guefaït-4.2 specimens (moderately deep median lingual notches, moderate 150 buccal flare of the crown, and distal marginal ridge uninterrupted by the distal buccal cleft) 151 denote cercopithecine affinities (Delson, 1973). The presence of a hypoconulid on the M3 152 rules out an assignment to cercopithecins (Delson, 1973, 1975), while the presence of a 153 tuberculum sextum further supports a papionin assignment, since this feature is rare in 154 colobines (Delson, 1973). Indeed, the two lower molars display a generalized papionin 155 occlusal morphology that is compatible with both Macaca (e.g., M. sylvanus [SOM Fig. S3B– 156 F], known from this region) and dentally conservative papioninans (e.g., Parapapio, 157 Pliopapio and Papio). This morphology differs from the more derived pattern of 158 Theropithecus, which is characterized by greater occlusal relief (with deeper foveae and 159 notches), obliquely oriented lophids and trigonid, large distal accessory cuspulid on M2, and 160 less buccal flare (Delson, 1973, 1975; Szalay and Delson, 1979). In molar proportions and 161 buccal flare, the Guefaït molars do not differ much from Lophocebus sp. from Koobi Fora 162 (Frost, 2001; Jablonski et al., 2008). However, they can be distinguished from the latter by 7 163 the smaller dimensions, and the more buccally located hypoconulid and better developed 164 tuberculum sextum on the M3 (albeit variable within papionin taxa, the tuberculum sextum 165 is only incipiently developed in some Koobi Fora specimens and more consistent with an 166 attribution to Macaca). 167 168 3.3. Metrical comparisons 169 Estimated BM for the Guefaït-4.2 M2 is around 11–12.5 kg (SOM S2), more similar to 170 female (10.1 kg) than male (14.5 kg) average BM for extant M. sylvanus (Delson et al., 2000). 171 Dental measurements for the Guefaït-4.2 specimens are visually compared with those of 172 macaques and similarly-sized extinct papionins of Parapapio and Pliopapio by means of 173 bivariate plots (Fig. 2) and boxplots (Fig. 3). Statistical comparisons are provided in SOM 174 Tables S2–S4 and discussed in SOM S3. 175 Although metrical comparisons for the Guefaït-4.2 I1 are limited by the small fossil 176 samples, it comfortably fits within the variation of extant M. sylvanus, not only in length and 177 width, but also in crown height (Figs. 2A, B and 3A–C; SOM Table S2). In turn, the Guefaït- 178 4.2 molars (Figs. 2C, D and 3D–I; SOM Table S2) fit well in size with M. s. sylvanus, M. majori, 179 ‘P.’ lothagamensis, and Pl. alemui, but appear too short for P. (cf.) ado, M. s. (cf.) pliocena, 180 M. s. florentina, and M. libyca; the Guefaït-4.2 M3 is further relatively too broad as 181 compared with these fossil macaques and Pl. alemui. It is noteworthy that the Guefaït-4.2 182 M3, like those of extant M. sylvanus, not only lacks the elongation of fossil European M. 183 sylvanus but displays a particularly broad (or short) crown in relative terms (albeit within the 184 variation of extant M. sylvanus). While the I1 from Guefaït-4.2. falls toward the upper size 185 range of M. sylvanus, the lower molars display the opposite pattern, suggesting that the 186 former derives from a different (larger) individual. 8 187 188 4. Discussion and conclusions 189 The occlusal morphology of the Guefaït-4.2 upper incisor and lower molars appears 190 compatible with Macaca and other fossil papionins. However, in size and proportion the 191 Guefaït-4.2 sample only fits well with M. s. sylvanus, as well as ‘P.’ lothagamensis, which 192 despite a markedly tapering distal contour similarly displays a buccally located hypoconulid 193 and frequently a tuberculum sextum (Leakey et al., 2003). The Guefaït-4.2 lower molars are 194 somewhat unusual due to the slightly projecting mesial fovea and the unusually ‘shortened’ 195 appearance of the M3. However, the latter is not caused by a poorly developed hypoconulid 196 per se, but due to the shortness of the hypoconulid lobe as a whole. This is probably the 197 most variable dental complex among cercopithecoids (Delson, 1973). The hypoconulid has 198 been lost in cercopithecins and is reduced in smaller Asian colobines (Swindler, 1983, 2002; 199 Szalay and Delson, 1979) but is almost invariably present in papionins, where it shows some 200 degree of intraspecific variation (Swindler, 1983). It is only rarely missing in some Papio, 201 Cercocebus and Macaca individuals (Delson, 1973, 1975; Szalay and Delson, 1979; Phillips- 202 Conroy, 1982), resulting in the reduction of the hypoconulid lobe (Phillips-Conroy, 1982). 203 However, the development of the hypoconulid in the Guefaït-4.2 M3 is comparable to that 204 of M. sylvanus (SOM Fig. S3). Furthermore, its overall proportions fit well within the 205 variation of extant M. sylvanus from northern Africa, while it can be metrically distinguished 206 from those of European Plio-Pleistocene M. sylvanus (which generally display better 207 developed hypoconulid lobes). 208 ‘Parapapio’ lothagamensis is a stem papionin (Gilbert, 2013) from the Late Miocene and 209 earliest Pliocene of eastern Africa (Leakey et al., 2003). Therefore, an assignment to Macaca 210 is favored on biogeographical and chronological grounds. Given that M. sylvanus is the only 9 211 macaque species previously recognized in the African Plio-Pleistocene (see Introduction), an 212 assignment to this species seems warranted—although, given the small available sample, 213 we prefer to use open nomenclature to keep our assignment tentative (i.e., M. cf. sylvanus). 214 Other authors might prefer an assignment to cf. Macaca or even Papionini indet. (Benefit et 215 al., 2008). However, an attribution to M. cf. sylvanus is in agreement with standard 216 taxonomic practice for European Plio-Pleistocene macaques, which following Szalay and 217 Delson (1979) are regularly allocated to M. sylvanus despite the lack of more diagnostic 218 cranial material (e.g., Alba et al., 2011, 2014, 2018). 219 The roughly coeval Moroccan site of Ahl al Oughlam (~2.5 Ma; Geraads, 2006, 2010) also 220 records the presence of Theropithecus (Alemseged and Geraads, 1998; Geraads et al., 1998; 221 Geraads, 2006), which raises the question as to why this genus is not recorded at Guefaït. 222 The fauna from Ahl al Oughlam displays close affinities with both eastern and southern 223 Africa (Geraads, 2006), indicating that—even if the Sahara belt acted as a biogeographic 224 barrier that promoted the divergence between Macacina and Papionina in the Late Miocene 225 (Delson, 1973; Roos et al., 2019)—it did not completely prevent the dispersal of 226 Theropithecus and other mammals from lower latitudes into northern Africa sometime 227 during the Late Pliocene (Geraads, 2006). During the Late Pliocene, eastern Africa 228 experienced a progressive aridification trend punctuated by more humid episodes, including 229 a brief humidity peak at 2.7–2.5 Ma (Trauth et al., 2007), which might have facilitated the 230 northward dispersal of Theropithecus, as it roughly coincides with the appearance in eastern 231 Africa of T. oswaldi oswaldi (Jablonski and Frost, 2010). Hypothetically, Guefaït-4.2 might 232 predate the dispersal of Theropithecus into northern Africa only if it is somewhat older than 233 Ahl al Oughlam—as tentatively suggested by the presence of some Late Pliocene murids 234 that are absent from the latter site (Agustí et al., 2017; Piñero et al., 2019). 10 235 Alternatively, the absence of Theropithecus from Guefaït-4.2 might reflect 236 paleoenvironmental differences relative to Ahl al Oughlam. Preliminary accounts of the 237 Guefaït 4.2 paleoenvironment based on the micromammal and paleoherpetological 238 assemblage suggest an open woodland habitat characterized by dry conditions within a 239 rocky landscape and with a permanent (lacustrine or riverine) water body nearby (Agustí et 240 al., 2017; Sala-Ramos et al., 2017). Such a paleoenvironmental reconstruction seems highly 241 suitable for macaques, based on the preferences of extant M. sylvanus, which is dependent 242 on water availability but occupies a wide variety of seasonal habitats—from forests to 243 grasslands and rocky escarpments—as long as there are trees nearby (Deag, 1974; Camperio 244 Ciani et al., 2001; Ménard, 2002; Fooden, 2007). However, this is unlikely to rule out the 245 presence of Theropithecus—particularly T. o. oswaldi, which was better adapted to open 246 habitats than its predecessor T. oswaldi darti (Leakey, 1993; Teaford, 1993; Jablonski et al., 247 2002; Folinsbee, 2008). Furthermore, based on the accompanying fauna, Ahl al Oughlam has 248 similarly been reconstructed as recording an open (and probably seasonal) environment 249 (Geraads et al., 1998; Geraads, 2006). 250 Given the rarity of primate finds and the restricted sample currently available for some 251 large mammals (such as suids and giraffids) from Guefaït-4.2, the absence of Theropithecus 252 might simply be a sampling artifact. In any case, it is noteworthy that the macaques from 253 Ahl al Oughlam and Guefaït-4.2 represent the last known record of Macaca in Africa until its 254 reappearance in the late Middle Pleistocene of Aïn Mefta (~200 ka; Pomel, 1892, 1896; 255 Geraads, 1987). This gap of more than 2 Myr contrasts with the more continuous record of 256 macaques in Europe throughout the Pleistocene and their subsequent greater abundance in 257 the Late Pleistocene of North Africa (e.g., Fooden, 2007, and references therein). It is 258 uncertain whether this pattern is also attributable to the biases of the record or whether it 11 259 might represent a local extinction followed by a subsequent redispersal of macaques from 260 elsewhere into northern Africa toward the end of the Middle Pleistocene. Additional 261 fieldwork would be required to clarify further papionin evolution and paleobiogeography 262 during the Pleistocene of northern Africa. 263 264 References 265 Agustí, J., Blain, H.-A., Piñero, P., Hajji, N.-e., Aouraghe, H., Haddoumi, H., El Hammouti, K., 266 El Harradji, A., 2016. Les petits vertébrés des sites de Guéfait : Biostratigraphie et 267 paléoenvironment. In: 10 Ans de Recherches & de Coopérations Scientifiques Maroco- 268 Espagnole dans la Province de Jerada (Maroc Oriental). Séminaire International sur le 269 Patrimoine Archéologique de la Province de Jerada. 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Fr. 3(2), 1–53. 462 Trauth, M.H., Maslin, M.A., Deino, A.L., Strecker, M.R., Bergner, A.G.N., Dühnforth, M., 463 2007. High- and low-latitude forcing of Plio-Pleistocene East African climate and human 464 evolution. J. Hum. Evol. 53, 475–486. 465 van den Brink, S., 1981. A propos du Cercopithecidae (Mammalia, primate) villafranchien de 466 l'Aïn Brimba, Tunisie. Géobios 14, 421–425. 467 468 Figure captions 20 469 470 Figure 1. Teeth of Macaca cf. sylvanus from Guefaït-4.2 (views from left to right for each 471 specimen): a) GFT4.2’18-1-Q14-70, left I1 crown in occlusal, occusal, lingual, mesial, labial, 472 and distal views; b) GFT4.2’18-1-Q14-13, right M1 lingual crown fragment in occlusal, buccal, 473 and lingual (cross-sectional) views; c) GFT4.2’18-1-P13-191, right M2 or M3 mesial germ 474 fragment in oblique occlusal and mesial views; d) GFT4.2’19-1-R15-45, right M2 distal germ 475 fragment in occlusal and distal views; e) GFT4.2’19-1-S15-65, right M2 crown with distal root 476 in occusal, lingual, mesial, buccal, and distal views; f) GFT4.2’19-1-R13-63, left M3 crown in 477 occusal, lingual, mesial, buccal, and distal views. 3D virtual models are available from 478 MorphoSource (see SOM Files S1–S6). 479 480 Figure 2. Bivariate plots of dental proportions in Macaca cf. sylvanus from Guefaït-4.2 (star) 481 compared with extant Barbary macaques (Macaca sylvanus sylvanus), fossil macaques from 482 the Plio-Pleistocene of Europe (Macaca sylvanus subspp. and Macaca majori), fossil Macaca 483 libyca and Macaca aff. sylvanus from Aïn Brimba, and selected stem papioninans from 484 eastern Africa (Parapapio [cf.] ado, ‘Parapapio’ lothagamensis, and Pliopapio alemui): A) I1 485 MD vs. H; B) I1 BL vs. MD; C) M2 MD vs. BL; D) M3 MD vs. BL. The variation of extant M. s. 486 sylvanus is denoted by the light gray convex hull, while that of extinct M. sylvanus subspp. 487 from Europe is denoted by a semitransparent red convex hull. Data sources are reported in 488 SOM Table S1. Abbreviations: H = maximum recorded labial height (mm); BL = buccolingual 489 breadth (BLm, mesial lophid for the molars; mm); MD = mesiodistal length (mm). 490 491 Figure 3. Box-and-whisker plots of MD (A, D, G), BL (B, E, H), and BLI (C, F, I) in extant 492 macaques and some fossil papionins compared with Macaca cf. sylvanus from Guefaït: a–c) 21 493 I1; d–f) M2; g–i) M3. Vertical lines denote the median; boxes denote the 25 and 75 494 percentiles (Q1 and Q3), i.e., the interquartile range (IQR); whiskers denote the maximum- 495 minimum values. Data sources are reported in SOM Table S1. Abbreviations: BL = 496 buccolingual breadth (mm; BLm, mesial lophid for the molars); BLI = breath/length index 497 (%); MD = mesiodistal length (mm). 22 Table 1 Provenance information and measurements of the Macaca cf. sylvanus teeth from Guefaït 4.2. Catalog No.a Tooth Level Square Collection date x y z MD H BL/BLm BLd BLI GFT4.2’18-1-P13-191 R M2/3 mesial germ fragment 1 P13 May 5, 2018 — — 101 — — >7.0 — — GFT4.2’18-1-Q14-70 L I1 crown 1 Q14 May 10, 2018 — — 104 (6.8)b 10.0 6.2 — 91.2 GFT4.2’18-1-Q14-13 R M1 lingual crown fragment 1 Q14 May 10, 2018 — — 89 — — — — — GFT4.2’19-1-R15-45 R M2 distal germ fragment 1 R15 April 25, 2019 18 22 105 — — — >7.1 — GFT4.2’19-1-S15-65 R M2 crown with distal root 1 S15 April 27, 2019 17 69 106 8.6 — 7.1 6.8 82.6 GFT4.2’19-1-R13-63 L M3 crown 1 R13 April 19, 2019 43 88 88 10.0 — 7.6 7.5 76.0 Abbreviations: BL = maximum buccolingual (or labiolingual) crown breadth (mm); BLI = breadth/length index (%), computed as BL / MD × 100 (based on BLm for the molars); BLm = buccolingual crown breadth (mm) at the mesial lophid (only for molars); BLd = buccolingual crown breadth (mm) at the distal lophid (only for molars); H = buccal crown height (mm; only for the incisor); L = left; MD = mesiodistal crown length (mm); R = right. a Each specimen is denoted by its accession No., including site acronym (GFT) and locality number (4.2.), followed by year of collection (e.g., ’19 = 2019) and, separated by hyphens, the stratigraphic level, square within the excavation grid (uppercase letter followed by number), and 1 field No. in the catalog of this locality for each given year. The x, y and z columns indicate the spatial position of the specimen in the locality grid. b Actual measurement is 6.5 mm, but given damage on the mesiolabial side it is estimated that measurement should be increased by ca. 5%. 2 A 8 I1 7 MD (mm) 6 5 4 4 5 6 7 8 9 10 11 12 13 H (mm) C 10 M2 B 7 I1 9 BL (mm) 6 8 BL (mm) 5 7 4 6 4 5 6 7 8 7 8 9 10 11 12 13 MD (mm) MD (mm) D 11 M3 10 Guefaït M. s. sylvanus M. s. (cf.) pliocena 9 M. s. florentina M. s. prisca BL (mm) M. aff. sylvanus M. majori M. libyca 8 P. (cf.) ado ‘P.’ lothagamensis Pl. alemui 7 6 8 9 10 11 12 13 14 15 16 MD (mm) M 2 MD a d M 3 MD I 1 MD g 10.50 12.00 12.75 13.50 14.25 15.00 10.50 12.00 12.75 9.00 9.75 6.75 7.50 8.25 9.00 9.75 4.0 4.5 5.0 5.5 6.0 6.5 7.0 7.5 8.0 11.25 11.25 Guefaït Guefaït Guefaït M. s. sylvanus M. s. sylvanus M. s. sylvanus M. s. pliocena M. s. pliocena M. s. florentina M. s. pliocena M. s. florentina M. s. prisca M. s. prisca M. s. florentina M. majori M. majori M. majori M. libyca M. libyca P. (cf.) ado P. (cf.) ado P. (cf.) ado ‘P.’ lothagamensis ‘P.’ lothagamensis ‘P.’ lothagamensis Pl. alemui Pl. alemui Pl. alemui M 3 BL M 2 BL I1 BL e b h 10.50 10.0 6.00 6.75 7.50 8.25 9.00 9.75 6.0 6.5 7.0 7.5 8.0 8.5 9.0 9.5 4.0 4.5 5.0 5.5 6.0 6.5 7.0 11.25 Guefaït Guefaït Guefaït M. s. sylvanus M. s. sylvanus M. s. sylvanus M. s. pliocena M. s. pliocena M. s. florentina M. s. florentina M. s. pliocena M. s. prisca M. s. prisca M. s. florentina M. majori M. majori M. majori M. libyca M. libyca P. (cf.) ado P. (cf.) ado P. (cf.) ado ‘P.’ lothagamensis ‘P.’ lothagamensis ‘P.’ lothagamensis Pl. alemui Pl. alemui Pl. alemui i f c M 3 BLI M 2 BLI I 1 BLI 105.0 67.5 75.0 82.5 90.0 97.5 112.5 100 70 75 80 85 90 95 52.5 60.0 67.5 75.0 82.5 90.0 97.5 105.0 Guefaït Guefaït Guefaït M. s. sylvanus M. s. sylvanus M. s. sylvanus M. s. pliocena M. s. pliocena M. s. florentina M. s. pliocena M. s. florentina M. s. prisca M. s. prisca M. s. florentina M. majori M. majori M. majori M. libyca M. libyca P. (cf.) ado P. (cf.) ado P. (cf.) ado ‘P.’ lothagamensis ‘P.’ lothagamensis ‘P.’ lothagamensis Pl. alemui Pl. alemui Pl. alemui Supplementary Online Material (SOM): New macaque fossil remains from Morocco David M. Alba a,*, Antonio Rodríguez-Hidalgo b,c,d, Hassan Aouraghe e, Jan van der Made f, Aïcha Oujaa g, Hamid Haddoumi e, Palmira Saladié c,h,i, Al Mahdi Aissa e, Juan Marín c, Mourad Farkouch c,e, Carlos Lorenzo h,c, Said Bengamra e, Eric Delson j,k,l,m,a, M. Gema Chacón c,h,n, Robert Sala-Ramos c,h a Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193, Cerdanyola del Vallès, Barcelona, Spain b Departamento de Prehistoria, Historia Antigua y Arqueología, Universidad Complutense de Madrid, Facultad de Geografía e Historia, C/ Prof. Aranguren s/n, 28040, Madrid, Spain c Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edifici W3), Tarragona, 43007, Spain d Instituto de Evolución en África (IDEA, Madrid), C/ Covarrubias 36, 28010, Madrid, Spain e Faculté des Sciences, Département de Géologie, Université Mohamed Premier, BV Mohammed VI, Quartier Al Qods, 60000, Oujda, Morocco f Consejo Superior de Investigaciones Científicas (CSIC), Museo Nacional de Ciencias Naturales, Departamento de Paleobiología, C/ José Gutiérrez Abascal, 2, 28006, Madrid, Spain g Institut National des Sciences de l’Archéologie et du Patrimoine (INSAP), Av. Allal El-Fassi, Hay Riad, 6828, Rabat, Morocco h Universitat Rovira i Virgili, Departament d’Història i Història de l’Art, Avinguda de Catalunya 35, 43002 Tarragona, Spain i Unidad Asociada al CSIC, Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales, C/ José Gutiérrez Abascal, 2, 28006, Madrid, Spain j Department of Anthropology, Lehman College of the City University of New York, 250 Bedford Park Boulevard West, Bronx, NY, 10468, USA k Department of Vertebrate Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY, 10024, USA 1 l PhD Programs in Anthropology and Earth & Environmental Sciences, The Graduate Center of the City University of New York, 365 Fifth Avenue, New York, NY, 10016, USA m New York Consortium in Evolutionary Primatology, New York, NY, USA n UMR7194 Histoire naturelle de l’Homme préhistorique (HNHP), Museum National d'Histoire Naturelle (MNHN), CNRS, Université Perpignan Via Domitia, Alliance Sorbonne Université – Musée de l’Homme, Place du Trocadéro 17, 75016 Paris, France Corresponding author. Email: david.alba@icp.cat (D.M. Alba) 2 SOM S1 Scanning methods To derive 3D virtual models of the studied teeth from Guefaït-4.2, the teeth were scanned at the Laboratory of Microscopy of the Centro Nacional de Investigación sobre la Evolución Humana–Unique Scientific & Technical Infrastuctures (Burgos, Spain) with a high- resolution X-ray μCT scanner phoenix v|tome|x s 240 (GE Sensing & Inspections Technologies) with the following parameters: voltage = 120 kV, current = 130 μA, filter = 0.2 mm Cu, resulting in an isometric voxel size of 16.30027 µm. Segmentation was performed with Avizo 7.0 (Visualization Sciences Group, Mérignac) using a combination of automatic and manual segmentation by a technican at the Institut Català de Paleontologia Miquel Crusafont. SOM S2 Body mass estimation Body mass (BM, in g) was estimated using Delson et al.’s (2000: Table 7) allometric regression for M2 mesial buccolingual width (BLm, in mm) for all cercopithecids: ln BM = 1.839 ln (M2 BLm) + 5.694, n = 68, SEE = 0.383, QMLE = 1.076 where SEE is the standard error of estimate and QMLE is the quasi-maximum likelihood estimator to correct for logarithmic transformation bias. Following Ruff (2003), 50% and 95% confidence intervals (CI) were computed as ±SEE × t(100-CI)(2),df × QMLE, where df = n - 2. For df = 66, t = 0.68 for 50% CI and 2.00 for 95% CI, resulting in CIs being calculated as ±0.280 and ±0.824 for 50% and 95% confidence levels, respectively. The resulting estimate is 11,753 g, with 50% CI = (11,473, 12,033) and 95% CI = (10,929, 12.577). SOM S3 Statistical comparisons ANOVA results indicate significant differences among the comparative sample for all variables except I1 H (SOM Table S3), but Tukey’s post hoc tests (SOM Table S4) indicate few significant pairwise differences for incisor dimensions (probably due to low statistical power associated with the small fossil samples). Therefore, below we focus on the greater number 3 of significant differences for the lower molars (see SOM Table S4), despite considerable overlap. Parapapio (cf.) ado displays on average significantly larger molars than ‘Parapapio’ lothagamensis, Pliopapio alemui, and macaques—except for Macaca libyca in most instances and also sometimes Macaca sylvanus pliocena and M. sylvanus florentina. ‘Parapapio’ lothagamensis and Pl. alemui are more similar to macaques in molar dimensions. However, the molars of ‘P.’ lothagamensis are significantly shorter than in M. s. (cf.) pliocena. In turn, Pl. alemui displays a narrower M2 than Macaca libyca, as well as an M3 intermediate in length between Macaca majori and fossil M. sylvanus from Europe; the M3 of Pl. alemui is also narrower than in extant M. sylvanus and M. majori. Among macaques, M. libyca does not display significant differences in molar dimensions compared to M. sylvanus sspp., but shows larger M2 and broader M3 than M. majori. The latter further differs in some molar dimensions from extant and fossil M. sylvanus. Macaca s.(cf.) pliocena and M. s. florentina do not significantly differ from one another in molar size or proportions, but they both differ from extant M. s. sylvanus by displaying on average a relatively narrower M3, which is also significantly longer in M. s. (cf.) pliocena. 4 SOM Figure S1. Location maps and pictures showing Guefaït-4 site and Guefaït-4.2 locality. A) Partial map of the Kingdom of Morocco (corresponding to the red rectangle overlapping the inset with the African continent); the area of interest (dotted box) is enlarged in panel B; data extracted from Map Tile 7_61/62/63-50 & 7_61/62/63-510 (CC BY-SA). OpenStreetMap© licensed under ODdL 1.0 (https://www.openstreetmap.org/copyright) by the OpenStreetMap Foundation (OSMF). ©OpenStreetMap contributors 5 (https://www.openstreetmap.org/). B) Location map of the Guefaït-4 site (base map taken from ASTER GDEM v. 3, https://ssl.jspacesystems.or.jp/ersdac/GDEM/E/). C–E) Different views of the Guefaït-4 site and Guefaït-4.2 locality (pictures by Alfonso Benito-Calvo and A.R.H). 6 SOM Figure S2. Left I1 crown (GFT4.2’18-1-Q14-70) of Macaca cf. sylvanus from Guefaït-4.2 (A) compared with two I1 of extant M. sylvanus (right specimens, mirrored for comparison): AMNH-M 70269 (B) and AMNH-M 185277 (C). All specimens are shown in lingual (left) and labial (right) views. 7 SOM Figure S3. Right M2 (GFT4.2’19-1-S15-65) and left M3 (GFT4.2’19-1-R13-63, mirrored for comparison) of Macaca cf. sylvanus from Guefaït-4.2 (A) compared with the M2–M3 of three extant M. s. sylvanus individuals (B–D), fossil M. aff. sylvanus from Aïn Brimba (E, only M2), M. s. florentina (F) and Macaca majori (G) from Italy, Parapapio cf. ado from Kanapoi (H), and Pliopapio alemui (only M3) from Gona (I) and Aramis (J): B) AMNH-M 19014 (M3 mirrored); C) AMNH-M 185277; D) AMNH-M 2060/5484; E) MNHN-F 1958-14-237; F) IGF 13084 (mirrored); G) IGF 2035 (mirrored); H) KNM-KP 286 (reproduced from Frost et al., 2020a: Fig. 1); I) GWM-9m/P256 (mirrored; reproduced from Frost et al., 2020b: Fig. 4); J) ARA-VP-1/2553 (mirrored; reproduced from Frost et al., 2020b: Fig. 4). All specimens are shown in occlusal view. 8 SOM Table S1 Dental measurements of Macaca cf. sylvanus from Guefaït-4.2 and the comparative sample, including: maximum recorded crown height (H, in mm; only for incisors), mesiodistal length (MD, in mm), buccolingual (or labiolingual) breadth (BL, in mm; both at the mesial [BLm] and distal [BLd] lophids for the molars), and breadth/length index (BLI, in %; computed based on BLm for molars). Values within parentheses are estimates. Catalog No. Species Site Tooth Sex MD BL or BLm BLd or H BLI Source 1 GFT4.2’18-1-Q14-70 Macaca cf. sylvanus Guefaït-4.2 I ? (6.8) 6.2 10.0 91.2 This study 1 AMNH-M 202286 Macaca sylvanus extant I M 7.3 6.5 12.8 89.0 PRIMO 1 AMNH-M 185277 Macaca sylvanus extant I F 5.72 5.17 7.35 90.4 PRIMO 1 AMNH-M 19014 Macaca sylvanus extant I F 5.98 4.73 9.39 79.1 PRIMO 1 AMNH-M 2060/5484 Macaca sylvanus extant I F 6.95 6.4 — 92.1 D.M.A. 1 AMNH-M 19178 Macaca sylvanus extant I F 6.1 5.7 — 93.4 D.M.A. 1 AMNH-M 70269 Macaca sylvanus extant I ? 6.9 5.25 — 76.1 D.M.A. 1 MNHN-ZM-AC 1910.166 Macaca sylvanus extant I M 6.9 5.9 11.7 85.5 PRIMO 1 MNHN-ZM-AC 1874.343 Macaca sylvanus extant I M 6.6 6.1 12.1 92.4 PRIMO 1 NHMUK-Z ZD.1843.5.27.1 Macaca sylvanus extant I M 6.9 5.8 8.6 84.1 PRIMO 1 NHMUK-Z ZD.1858.4.5.1 Macaca sylvanus extant I M 7.0 5.8 11.0 82.9 PRIMO 1 NHMUK-Z ZD.1911.11.4.1 Macaca sylvanus extant I F 6.2 5.3 8.1 85.5 PRIMO 1 NHMUK-Z ZD.1919.8.19.1 Macaca sylvanus extant I M 6.1 4.9 7.2 80.3 PRIMO 1 NHMUK-Z ZD.1939.1048 Macaca sylvanus extant I M 6.8 5.5 10.6 80.9 PRIMO 1 NMNH 196984 Macaca sylvanus extant I F 6.35 5.71 9.4 89.9 PRIMO 1 NMNH 476782 Macaca sylvanus extant I F 6.18 4.89 8.5 79.1 PRIMO 1 NMNH 476786 Macaca sylvanus extant I F 5.24 4.98 5.3 95.0 PRIMO 1 NMNH 476787 Macaca sylvanus extant I F 4.95 5.24 4.6 105.9 PRIMO 1 NMNH 341870 Macaca sylvanus extant I F 5.44 5.61 9.03 103.1 PRIMO 1 NMNH 476783 Macaca sylvanus extant I F 5.59 4.88 5.84 87.3 PRIMO 9 NMNH 476784 Macaca sylvanus extant I1 M 6.17 5.42 8.91 87.8 PRIMO 1 NMNH 255979 Macaca sylvanus extant I M 6.9 5.36 9.74 77.7 PRIMO 1 MCZ 5964 Macaca sylvanus extant I F 7.15 5.65 — 79.0 D.M.A. 1 MCZ 5084 Macaca sylvanus extant I F 7.0 5.75 — 82.1 D.M.A. 1 IPH UNCAT Macaca sylvanus pliocena Vallonnet I ? 6.8 5.2 10.1 76.5 PRIMO 1 HUJ-ESE UB 323 Macaca sylvanus florentina 'Ubeidiya I ? 7.06 5.16 9.23 73.1 PRIMO 1 MHNL 164295 Macaca sylvanus florentina Saint-Vallier I ? 7.2 5.4 11.1 75.0 PRIMO 1 NMB TY 5203 Macaca majori Capo Figari I F 6.0 4.6 9.0 76.7 PRIMO 1 NMB TY 5273 Macaca majori Capo Figari I ? 5.9 6.1 9.7 103.4 PRIMO 1 NMB TY 5274 Macaca majori Capo Figari I ? 6.2 4.4 8.5 71.0 PRIMO 1 NMB TY 5275 Macaca majori Capo Figari I ? 6.0 4.9 8.1 81.7 PRIMO 1 NMB TY 5279 Macaca majori Capo Figari I ? 5.7 4.2 6.6 73.7 PRIMO 1 NMB TY 5316 Macaca majori Capo Figari I ? 6.2 4.2 8.6 67.7 PRIMO 1 NMT LAET 77-4595 Parapapio ado Laetoli I F 6.4 5.3 12.2 82.8 PRIMO 1 KNM-KP 30149 Parapapio cf. ado Kanapoi I ? 7.5 5.8 11.3 77.3 Frost et al. (2020) 1 KNM-LT 23163 Parapapio lothagamensis Lothagam I ? 7.5 6.3 — 84.0 Leakey et al. (2003) 1 KNM-LT 24096 Parapapio lothagamensis Lothagam I ? 7.9 6.1 — 77.2 Leakey et al. (2003) 1 KNM-LT 24100 Parapapio lothagamensis Lothagam I ? 7.3 6.4 — 87.7 Leakey et al. (2003) 1 KNM-LT 26406 Parapapio lothagamensis Lothagam I ? 6.8 6.1 11.5 89.7 Leakey et al. (2003) 1 KNM-LT 28792 Parapapio lothagamensis Lothagam I F 7.3 6.4 11.7 87.7 Leakey et al. (2003) 1 NME GWM-3/P26 Pliopapio alemui Gona I ? 6.4 6.3 10.4 98.4 Frost et al. (2020b) 1 NME GWM-9m/P264 Pliopapio alemui Gona I ? 5.8 — 8.5 — Frost et al. (2020b) 1 NME GWM-9s/P414 Pliopapio alemui Gona I ? 5.1 5.7 6.4 111.8 Frost et al. (2020b) 1 NME GWM-44/P1 Pliopapio alemui Gona I ? — 5.8 9.3 — Frost et al. (2020b) 1 NME GWM-67/P55 Pliopapio alemui Gona I ? 6.3 6.3 8.0 100.0 Frost et al. (2020b) 1 NME ARA-VP 1/816 Pliopapio alemui Aramis I F 5.81 5 6.54 86.1 PRIMO GFT4.2’19-1-S15-65 Macaca cf. sylvanus Guefaït-4.2 M2 ? 8.6 7.1 6.8 82.6 This study MNHN-F 1958-14-237 Macaca aff. sylvanus Aïn Brimba M2 ? 9.3 7.8 7.0 83.9 PRIMO 10 AMNH-M 202286 Macaca sylvanus sylvanus extant M2 M 9.1 7.6 7.5 83.5 PRIMO MNHN-ZM-AC 1910.166 Macaca sylvanus sylvanus extant M2 M 10.1 8.3 7.3 82.2 PRIMO MNHN-ZM-AC 1900.244 Macaca sylvanus sylvanus extant M2 F 10.2 8.0 6.8 78.4 PRIMO MNHN-ZM-AC 1874.343 Macaca sylvanus sylvanus extant M2 M 9.5 7.9 6.6 83.2 PRIMO MNHN-ZM-AC A-1376 Macaca sylvanus sylvanus extant M2 F 9.1 6.9 6.1 75.8 PRIMO NHMUK-Z ZD.1843.5.27.1 Macaca sylvanus sylvanus extant M2 M 9.3 7.1 6.1 76.3 PRIMO NHMUK-Z ZD.1911.11.4.1 Macaca sylvanus sylvanus extant M2 F 10.0 8.3 6.7 83.0 PRIMO NHMUK-Z ZD.1919.8.19.1 Macaca sylvanus sylvanus extant M2 M 9.2 6.8 6.2 73.9 PRIMO NHMUK-Z ZD.1939.1048 Macaca sylvanus sylvanus extant M2 M 10.3 8.5 7.5 82.5 PRIMO AMNH-M 185277 Macaca sylvanus sylvanus extant M2 F 8.65 7.6 6.27 87.9 PRIMO AMNH-M 19014 Macaca sylvanus sylvanus extant M2 F 8.08 6.96 6.33 86.1 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M2 ? 8.8 8.4 7.3 95.5 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M2 M 9.0 7.4 6.7 82.2 PRIMO NMNH 196984 Macaca sylvanus sylvanus extant M2 F 8.79 6.88 6.16 78.3 PRIMO NMNH 476782 Macaca sylvanus sylvanus extant M2 F 8.68 6.98 6.62 80.4 PRIMO NMNH 341870 Macaca sylvanus sylvanus extant M2 F 7.24 6.57 6.2 90.7 PRIMO NMNH 341872 Macaca sylvanus sylvanus extant M2 F 7.05 6.65 6.3 94.3 PRIMO NMNH 476783 Macaca sylvanus sylvanus extant M2 F 8.01 6.77 6.47 84.5 PRIMO NMNH 476784 Macaca sylvanus sylvanus extant M2 M 9.12 6.72 6.45 73.7 PRIMO NMNH 255979 Macaca sylvanus sylvanus extant M2 M 9.76 8.91 8.1 91.3 PRIMO AMNH-M 2060/5484 Macaca sylvanus sylvanus extant M2 F 10.2 8.2 7.4 80.4 D.M.A. AMNH-M 19178 Macaca sylvanus sylvanus extant M2 F 11.1 8.3 7.3 74.8 D.M.A. AMNH-M 70269 Macaca sylvanus sylvanus extant M2 ? 9.65 6.85 6.15 71.0 D.M.A. MCZ 5964 Macaca sylvanus sylvanus extant M2 F 10.0 8.3 6.6 83.0 D.M.A. MCZ 7098 Macaca sylvanus sylvanus extant M2 M 8.6 (8.2) (7.3) (95.3) D.M.A. MCZ 5084 Macaca sylvanus sylvanus extant M2 F 9.4 7.75 6.8 82.4 D.M.A. MNHN-ZM-MO 1939.1118 Macaca sylvanus sylvanus extant M2 M 9.35 8.15 6.85 87.2 PRIMO MNHN-ZM-MO 1945.173 Macaca sylvanus sylvanus extant M2 F 10.0 7.7 7.1 77.0 PRIMO 11 MNHN-ZM-MO 1962.1473 Macaca sylvanus sylvanus extant M2 M 8.9 7.6 6.6 85.4 PRIMO MNHN-ZM-MO 1962.1470 Macaca sylvanus sylvanus extant M2 F 9.85 8.7 7.0 88.3 PRIMO MNHN-ZM-MO 1931.835 Macaca sylvanus sylvanus extant M2 M 9.5 7.85 6.6 82.6 PRIMO MNHN-ZM-MO 1940.1201 Macaca sylvanus sylvanus extant M2 M 9.4 7.95 6.7 84.6 PRIMO MNHN-ZM-MO 1962.1469 Macaca sylvanus sylvanus extant M2 M 10.0 7.9 6.9 79.0 PRIMO MNHN-ZM-MO 1926.299 Macaca sylvanus sylvanus extant M2 M 10.0 7.7 7.2 77.0 PRIMO MNHN-ZM-MO 1942.180 Macaca sylvanus sylvanus extant M2 F 8.5 7.5 6.7 88.2 PRIMO MNHN-ZM-AC 1907.59 Macaca sylvanus sylvanus extant M2 M 10.5 8.45 7.35 80.5 PRIMO MNHN-ZM-AC A1415 Macaca sylvanus sylvanus extant M2 M 9.0 7.3 6.25 81.1 PRIMO MNHN-ZM-AC 1892.78 Macaca sylvanus sylvanus extant M2 F 9.0 9.1 7.15 101.1 PRIMO MNHN-ZM-AC 1876.286 Macaca sylvanus sylvanus extant M2 M 9.8 7.5 7.0 76.5 PRIMO MNHN-ZM-AC 1879.133 Macaca sylvanus sylvanus extant M2 M 9.5 7.9 6.85 83.2 PRIMO MNHG 446.36 Macaca sylvanus sylvanus extant M2 F 9.2 7.9 7.0 85.9 PRIMO MAC 283/4 Macaca sylvanus sylvanus extant M2 M 11.1 9.3 7.7 83.8 PRIMO NHMW 32990 Macaca sylvanus sylvanus extant M2 M 8.9 6.95 6.15 78.1 PRIMO NHMW 2614/B3976 Macaca sylvanus sylvanus extant M2 M 10.2 8.2 7.4 80.4 PRIMO NHMW B 3849 Macaca sylvanus sylvanus extant M2 F 9.5 7.4 6.6 77.9 PRIMO NHMW 4408 Macaca sylvanus sylvanus extant M2 M 10.3 8.5 7.3 82.5 PRIMO NHMW 2616 Macaca sylvanus sylvanus extant M2 F 9.2 7.7 6.9 83.7 PRIMO NHMW 2615 Macaca sylvanus sylvanus extant M2 F 9.1 7.6 6.4 83.5 PRIMO NHMW 4407 Macaca sylvanus sylvanus extant M2 F 9.2 7.8 6.6 84.8 PRIMO NHMW 34222 Macaca sylvanus sylvanus extant M2 F 8.7 7.4 6.6 85.1 PRIMO MAU-T no number Macaca sylvanus sylvanus extant M2 F 8.4 7.35 7.45 87.5 PRIMO NHMUK-Z ZD.1984.1734 Macaca sylvanus sylvanus extant M2 M 8.7 6.3 6.1 72.4 PRIMO NHMUK-Z ZD.1858.5.4.247 Macaca sylvanus sylvanus extant M2 M 9.8 7.7 6.7 78.6 PRIMO NHMUK-Z ZD.1975.569 Macaca sylvanus sylvanus extant M2 M 9.9 7.4 6.5 74.7 PRIMO NHMUK-Z ZD.1977.3119 Macaca sylvanus sylvanus extant M2 M 9.9 7.8 7.0 78.8 PRIMO Bonifay UNCAT Macaca sylvanus pliocena St. Estève Janson M2 M 10.5 8.5 8.2 81.0 PRIMO 12 NMP UNCAT Macaca sylvanus pliocena Zlaty Kun C718 M2 ? 9.5 7.6 7.3 80.0 PRIMO NMP UNCAT Macaca sylvanus pliocena Zlaty Kun C718 M2 M 10.2 7.8 7.3 76.5 PRIMO ORGNAC 353 Macaca sylvanus pliocena Orgnac-3 M2 ? 8.8 7.9 7.6 89.8 PRIMO NMP UNCAT Macaca sylvanus pliocena Gombasek M2 M 10.4 8.5 8.0 81.7 PRIMO IG-R UNCAT Macaca sylvanus pliocena Monte Sacro, Rome M2 M 9.7 7.7 6.9 79.4 PRIMO Harlé UNCAT Macaca sylvanus pliocena Montsaunès M2 ? 9.7 7.2 6.9 74.2 PRIMO IPH UNCAT Macaca sylvanus pliocena Vallonnet M2 ? 10.2 7.9 6.9 77.5 PRIMO IAPU-P UNCAT Macaca sylvanus pliocena Valdemino M2 M? 9.4 7.4 6.9 78.7 PRIMO LPHP-S 777/G1 & 771 Macaca sylvanus pliocena Gajtan M2 F? 8.9 7.3 7.3 82.0 PRIMO LPHP-S 775 Macaca sylvanus pliocena Gajtan M2 ? 10.6 7.5 7.2 70.8 PRIMO SIP-V unknown Macaca sylvanus pliocena Cova Negra M2 ? 10.8 8.5 — (78.7) PRIMO BGDG LzII.M15.-271.300 Macaca sylvanus cf. pliocena Lezetxiki II M2 F 9.6 8.0 7.8 83.3 Castaños et al. (2011) MCSNBS PA11136 Macaca sylvanus cf. pliocena Quecchia Quarry M2 ? 9.5 7.8 7.0 82.1 Bona et al. (2016) UNIARQ ARO17-J8-Xa-934 Macaca sylvanus cf. pliocena Gruta da Aroeira M2 ? 9.45 6.55 6.5 69.3 Alba et al. (2019) HUJ-ESE UB 314 Macaca sylvanus florentina 'Ubeidiya M2 F 9.41 6.82 6.34 72.5 PRIMO HUJ-ESE UB 331 Macaca sylvanus florentina 'Ubeidiya M2 F? 9.36 7.01 6.85 74.9 PRIMO HUJ-ESE UB 334 Macaca sylvanus florentina 'Ubeidiya M2 ? 9.35 7.28 6.82 77.9 PRIMO HUJ-ESE UB 347 Macaca sylvanus florentina 'Ubeidiya M2 ? 9.49 7.57 6.79 79.8 PRIMO HUJ-ESE UB 346 Macaca sylvanus florentina 'Ubeidiya M2 ? 8.9 6.7 6.54 75.3 PRIMO IGF 10034 Macaca sylvanus florentina Upper Valdarno M2 M 9.8 8.0 7.3 81.6 PRIMO IGF 10035 Macaca sylvanus florentina Upper Valdarno M2 M 10.2 8.0 7.5 78.4 PRIMO IGF 12868 Macaca sylvanus florentina Upper Valdarno M2 F 9.6 8.3 7.5 86.5 PRIMO IGF 13084 Macaca sylvanus florentina Upper Valdarno M2 ? 9.7 7.7 6.7 79.4 PRIMO IGF 13087 Macaca sylvanus florentina Upper Valdarno M2 ? 9.0 6.8 6.6 75.6 PRIMO NMB VA 352 Macaca sylvanus florentina Upper Valdarno M2 M 9.0 7.5 7.1 83.3 PRIMO NMB VA 1088 Macaca sylvanus florentina Upper Valdarno M2 ? 9.4 7.3 7.4 77.7 PRIMO NMB VA 1415 Macaca sylvanus florentina Upper Valdarno M2 M? 9.7 7.4 7.2 76.3 PRIMO NMB VA 2058 Macaca sylvanus florentina Upper Valdarno M2 M 9.5 7.2 6.7 75.8 PRIMO 13 MAP 10 Macaca sylvanus florentina Upper Valdarno M2 M 9.9 7.9 7.4 79.8 PRIMO RGM-L 47424 Macaca sylvanus florentina Tegelen M2 M 11.0 8.3 7.9 75.5 PRIMO RGM-L 53158 Macaca sylvanus florentina Tegelen M2 ? 9.5 8.3 7.8 87.4 PRIMO ZMA 12717 Macaca sylvanus florentina Tegelen M2 ? 9.8 7.5 7 76.5 PRIMO MUPE GCP-CV4062 Macaca sylvanus florentina Quibas M2 ? 10.2 8.5 7.6 83.3 Alba et al. (2011) MUPE GCP-CV4060 Macaca sylvanus florentina Quibas M2 ? 9.4 (7.6) 7.2 (80.9) Alba et al. (2011) UM Q03-E-4a Macaca sylvanus florentina Quibas M2 ? 10.0 8.2 6.8 82.0 Alba et al. (2011) MSNC Mugello B Macaca sylvanus florentina Mugello M2 ? 9.3 6.85 6.7 73.7 Zanaga (1998) MNHN-F UNCAT Macaca sylvanus prisca Montpellier M2 M 8.8 6.8 6.8 77.3 PRIMO FSL 40191 Macaca sylvanus prisca Montpellier M2 ? 8.0 6.8 6.0 85.0 PRIMO MPC 8116 Macaca majori Is Oreris M2 ? 8.47 6.93 5.96 81.8 Zoboli et al. (2016) IGF 2035 Macaca majori Capo Figari M2 M 8.9 7.0 6.9 78.7 PRIMO IGF 2036 Macaca majori Capo Figari M2 M 8.7 6.8 6.5 78.2 PRIMO NMB TY 5202 Macaca majori Capo Figari M2 F 7.6 6.4 6.5 84.2 PRIMO NMB TY 5209 Macaca majori Capo Figari M2 ? 8.8 7.2 7.0 81.8 PRIMO NMB TY 5210 Macaca majori Capo Figari M2 ? 8.7 7.0 6.8 80.5 PRIMO NMB TY 12454 Macaca majori Capo Figari M2 ? 8.6 7.0 6.6 81.4 PRIMO NMB TY 12460 Macaca majori Capo Figari M2 M 8.6 7.1 6.9 82.6 PRIMO NMB TY 12475 Macaca majori Capo Figari M2 ? 8.8 6.9 6.3 78.4 PRIMO GPI-F 1652 Macaca libyca Wadi Natrun M2 ? 9.6 8.5 8.6 88.5 PRIMO YPM 21551 Macaca libyca Wadi Natrun M2 F 9.4 8.1 7.8 86.2 PRIMO YPM 21552 Macaca libyca Wadi Natrun M2 ? 9.8 9.4 — (95.9) PRIMO BSPG 1920 I 505 Macaca libyca Wadi Natrun M2 ? 10.9 7.9 7.3 72.5 PRIMO MB Ma 42444 Parapapio ado Laetoli M2 M 11.4 9.0 8.0 78.9 PRIMO MB Ma 42445 Parapapio ado Laetoli M2 ? 11.3 8.6 8.1 76.1 PRIMO MB Ma 42451 Parapapio ado Laetoli M2 ? 10.0 7.9 7.1 79.0 PRIMO MB Ma 42452 Parapapio ado Laetoli M2 ? 9.6 7.3 7.5 76.0 PRIMO MB Ma 42453 Parapapio ado Laetoli M2 ? 10.8 8.6 8.0 79.6 PRIMO 14 MB Ma 42442 (& 42446) Parapapio ado Laetoli M2 M 11.0 9.8 9.2 89.1 PRIMO NMT LAET 74-223 Parapapio ado Laetoli M2 F 10.8 8.7 8.8 80.6 PRIMO NMT LAET 74-243-4 Parapapio ado Laetoli M2 F 10.6 8.3 7.8 78.3 PRIMO NMT LAET 75-483 Parapapio ado Laetoli M2 ? 11.3 8.8 7.9 77.9 PRIMO NMT LAET 75-812 Parapapio ado Laetoli M2 ? 12.5 9.8 8.8 78.4 PRIMO NMT LAET 75-3035 Parapapio ado Laetoli M2 F 10.6 8.2 8.9 77.4 PRIMO NMT LAET 77-4595 Parapapio ado Laetoli M2 F 11.3 9.1 9.4 80.5 PRIMO KNM-KP 26942 Parapapio cf. ado Kanapoi M2 ? 11.1 9.8 8.7 88.3 Frost et al. (2020a) KNM-KP 29306 Parapapio cf. ado Kanapoi M2 M 9.3 8.3 7.9 89.2 Frost et al. (2020a) KNM-KP 29312 Parapapio cf. ado Kanapoi M2 ? 10.8 9.5 8.7 88.0 Frost et al. (2020a) KNM-KP 30538 Parapapio cf. ado Kanapoi M2 F 9.5 7.9 7.2 83.2 Frost et al. (2020a) KNM-KP 53085 Parapapio cf. ado Kanapoi M2 M 10.7 8.9 8.6 83.2 Frost et al. (2020a) KNM-ER 3122 Parapapio cf. ado Koobi Fora M2 ? 8.8 7.4 6.7 84.1 Jablonski et al. (2008) KNM-ER 17587 Parapapio cf. ado Koobi Fora M2 ? 8.9 8.9 7.9 100.0 Jablonski et al. (2008) KNM-ER 18881 Parapapio cf. ado Koobi Fora M2 ? 9.4 8.3 8.1 88.3 Jablonski et al. (2008) KNM-ER 36983 Parapapio cf. ado Koobi Fora M2 ? 11.0 8.2 8 74.5 Jablonski et al. (2008) KNM-ER 36998 Parapapio cf. ado Koobi Fora M2 ? 11.1 8.4 7.9 75.7 Jablonski et al. (2008) KNM-ER 37093 Parapapio cf. ado Koobi Fora M2 ? 9.6 8.1 7.9 84.4 Jablonski et al. (2008) KNM-ER 37128 Parapapio cf. ado Koobi Fora M2 ? 9.9 7.0 7.1 70.7 Jablonski et al. (2008) KNM-LT 415 Parapapio lothagamensis Lothagam M2 ? 8.2 7.4 6.9 90.2 Leakey et al. (2003) KNM-LT 22971 Parapapio lothagamensis Lothagam M2 ? 8.8 8.4 7.8 95.5 Leakey et al. (2003) KNM-LT 23091 Parapapio lothagamensis Lothagam M2 M 9.1 7.9 7.8 86.8 Leakey et al. (2003) KNM-LT 24094 Parapapio lothagamensis Lothagam M2 ? 8.1 6.5 5.9 80.2 Leakey et al. (2003) KNM-LT 24095 Parapapio lothagamensis Lothagam M2 ? 9.1 7.4 7.7 81.3 Leakey et al. (2003) KNM-LT 24122 Parapapio lothagamensis Lothagam M2 ? 9.5 7.4 6.6 77.9 Leakey et al. (2003) KNM-LT 24135 Parapapio lothagamensis Lothagam M2 ? 8.3 7.4 7.2 89.2 Leakey et al. (2003) KNM-LT 24140 Parapapio lothagamensis Lothagam M2 ? (8.6) (7.0) (7.2) (81.4) Leakey et al. (2003) KNM-LT 26395 Parapapio lothagamensis Lothagam M2 ? 8.5 7.1 6.9 83.5 Leakey et al. (2003) 15 KNM-LT 28728 Parapapio lothagamensis Lothagam M2 ? 10.3 9.0 8.4 87.4 Leakey et al. (2003) NME GWM-3/P84 Pliopapio alemui Gona M2 F 9.0 7.0 7.4 77.8 Frost et al. (2020b) NME GWMS-11/P65 cf. Pliopapio alemui Gona M2 ? 9.1 7.4 7.0 81.3 Frost et al. (2020b) NME ARA-VP 1/816 Pliopapio alemui Aramis M2 F 9.47 7.58 7.6 80.1 PRIMO NME ARA-VP 1/1006 Pliopapio alemui Aramis M2 F 9.22 6.88 7.05 74.7 PRIMO NME ARA-VP 1/1349 Pliopapio alemui Aramis M2 F 8.74 6.49 6.67 74.2 PRIMO NME ARA-VP 6/8 Pliopapio alemui Aramis M2 ? 8.76 6.38 6.82 72.8 PRIMO GFT4.2’19-1-R13-63 Macaca cf. sylvanus Guefaït-4.2 M3 ? 10.0 7.6 7.5 76.0 This study AMNH-M 202286 Macaca sylvanus sylvanus extant M3 M 12.2 8.2 7.6 67.2 PRIMO MNHN-ZM-AC 1910.166 Macaca sylvanus sylvanus extant M3 M 12.5 9.3 7.5 74.4 PRIMO MNHN-ZM-AC 1900.244 Macaca sylvanus sylvanus extant M3 F 11.1 8.0 6.7 72.1 PRIMO MNHN-ZM-AC 1874.343 Macaca sylvanus sylvanus extant M3 M 12.2 8.4 6.3 68.9 PRIMO MNHN-ZM-AC A-1376 Macaca sylvanus sylvanus extant M3 F 11.6 8.3 6.4 71.6 PRIMO NHMUK-Z ZD.1843.5.27.1 Macaca sylvanus sylvanus extant M3 M 11.5 7.7 6.6 67.0 PRIMO NHMUK-Z ZD.1911.11.4.1 Macaca sylvanus sylvanus extant M3 F 11.5 8.6 6.9 74.8 PRIMO NHMUK-Z ZD.1919.8.19.1 Macaca sylvanus sylvanus extant M3 M 10.6 6.8 6.0 64.2 PRIMO AMNH-M 185277 Macaca sylvanus sylvanus extant M3 F 10.7 7.89 6.44 73.7 PRIMO AMNH-M 19014 Macaca sylvanus sylvanus extant M3 F 10.51 7.67 6.19 73.0 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M3 ? 12.6 9.4 7.6 74.6 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M3 ? 11.4 8.0 6.5 70.2 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M3 ? 13.2 7.4 6.8 56.1 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M3 ? 12.4 8.3 7.2 66.9 PRIMO MNHN-F UNCAT Macaca sylvanus sylvanus extant M3 M 11.8 7.9 6.4 66.9 PRIMO NMNH 196984 Macaca sylvanus sylvanus extant M3 F 11.09 7.59 6.44 68.4 PRIMO NMNH 476782 Macaca sylvanus sylvanus extant M3 F 10.7 7.31 6.79 68.3 PRIMO NMNH 476786 Macaca sylvanus sylvanus extant M3 F 11.73 8.24 7.16 70.2 PRIMO NMNH 341870 Macaca sylvanus sylvanus extant M3 F 9.34 6.56 5.67 70.2 PRIMO NMNH 341872 Macaca sylvanus sylvanus extant M3 F 9.49 6.41 5.9 67.5 PRIMO 16 NMNH 476783 Macaca sylvanus sylvanus extant M3 F 9.81 6.79 6.67 69.2 PRIMO NMNH 476784 Macaca sylvanus sylvanus extant M3 M 10.95 7.61 6.77 69.5 PRIMO NMNH 255979 Macaca sylvanus sylvanus extant M3 M 11.77 9.26 7.92 78.7 PRIMO AMNH-M 2060/5484 Macaca sylvanus sylvanus extant M3 F 12.1 8.55 7.05 70.7 D.M.A. AMNH-M 19178 Macaca sylvanus sylvanus extant M3 F 13.05 8.95 7.55 68.6 D.M.A. MCZ 7098 Macaca sylvanus sylvanus extant M3 M 10.3 (7.6) >6.5 (73.8) D.M.A. MNHN-ZM-MO 1939.1118 Macaca sylvanus sylvanus extant M3 M 12.2 9.5 7.45 77.9 PRIMO MNHN-ZM-MO 1945.173 Macaca sylvanus sylvanus extant M3 F 12.4 8.0 6.75 64.5 PRIMO MNHN-ZM-MO 1962.1473 Macaca sylvanus sylvanus extant M3 M 11.85 8.4 6.9 70.9 PRIMO MNHN-ZM-MO 1931.835 Macaca sylvanus sylvanus extant M3 M 12.3 9.0 7.0 73.2 PRIMO MNHN-ZM-MO 1940.1201 Macaca sylvanus sylvanus extant M3 M 12.7 8.5 6.7 66.9 PRIMO MNHN-ZM-MO 1926.299 Macaca sylvanus sylvanus extant M3 M 11.3 11.4 7.9 100.9 PRIMO MNHN-ZM-MO 1942.180 Macaca sylvanus sylvanus extant M3 F 9.6 7.65 6.4 79.7 PRIMO MNHN-ZM-AC 1907.59 Macaca sylvanus sylvanus extant M3 M 14.6 9.1 7.8 62.3 PRIMO MNHN-ZM-AC A1415 Macaca sylvanus sylvanus extant M3 M 11.4 8.2 6.2 71.9 PRIMO MNHN-ZM-AC 1879.133 Macaca sylvanus sylvanus extant M3 M 12.6 8.6 6.7 68.3 PRIMO MNHG 446.36 Macaca sylvanus sylvanus extant M3 F 11.5 8.8 6.7 76.5 PRIMO NHMW 32990 Macaca sylvanus sylvanus extant M3 M 11.55 7.55 6.45 65.4 PRIMO NHMW 2614/B3976 Macaca sylvanus sylvanus extant M3 M 13.6 8.6 7.6 63.2 PRIMO NHMW B 3849 Macaca sylvanus sylvanus extant M3 F 11.8 7.8 6.6 66.1 PRIMO NHMW 4408 Macaca sylvanus sylvanus extant M3 M 13.0 8.8 7.4 67.7 PRIMO NHMW 2616 Macaca sylvanus sylvanus extant M3 F 11.2 8.0 6.7 71.4 PRIMO NHMW 2615 Macaca sylvanus sylvanus extant M3 F 12.1 8.1 6.6 66.9 PRIMO NHMW 4407 Macaca sylvanus sylvanus extant M3 F 11.5 8.4 6.8 73.0 PRIMO NHMW 34222 Macaca sylvanus sylvanus extant M3 F 10.8 7.8 7.0 72.2 PRIMO MAU-T no number Macaca sylvanus sylvanus extant M3 F 9.75 6.85 6.85 70.3 PRIMO NHMUK-Z ZD.1939.3470 Macaca sylvanus sylvanus extant M3 F 10.4 7.3 6.1 70.2 PRIMO NHMUK-Z ZD.1984.1734 Macaca sylvanus sylvanus extant M3 M 11.7 6.9 6.2 59.0 PRIMO 17 NHMUK-Z ZD.1858.5.4.247 Macaca sylvanus sylvanus extant M3 M 11.4 8.2 7.1 71.9 PRIMO NHMUK-Z ZD.1975.569 Macaca sylvanus sylvanus extant M3 M 12.3 7.8 6.8 63.4 PRIMO NHMUK-Z ZD.1977.3119 Macaca sylvanus sylvanus extant M3 M 12.5 8.1 7.3 64.8 PRIMO SMF 71-1301 Macaca sylvanus pliocena Hohensülzen M3 ? 12.1 7.3 6.7 60.3 PRIMO Bonifay UNCAT Macaca sylvanus pliocena St. Estève Janson M3 M 14.4 9.4 8.4 65.3 PRIMO NMP UNCAT Macaca sylvanus pliocena Zlaty Kun C718 M3 ? 11.7 8.0 7.5 68.4 PRIMO NMP UNCAT Macaca sylvanus pliocena Zlaty Kun C718 M3 M 13.0 8.0 7.9 61.5 PRIMO ORGNAC 723 Macaca sylvanus pliocena Orgnac-3 M3 ? 13.0 8.2 8.0 63.1 PRIMO NMP UNCAT Macaca sylvanus pliocena Gombasek M3 M 13.5 8.6 7.7 63.7 PRIMO IG-R UNCAT Macaca sylvanus pliocena Monte Sacro, Rome M3 M 13.0 8.2 7.2 63.1 PRIMO LPHP-S 777/G1 & 771 Macaca sylvanus pliocena Gajtan M3 F? 12.2 7.6 7.1 62.3 PRIMO SIP-V unknown Macaca sylvanus pliocena Cova Negra M3 ? 13.9 9.2 — 66.2 PRIMO UG unknown Macaca sylvanus pliocena Solana de Zamborino M3 ? 12.1 7.5 7.1 62.0 PRIMO BGDG LzII.M15.-271.300 Macaca sylvanus cf. pliocena Lezetxiki II M3 F 13.0 8.6 7.4 66.2 Castaños et al. (2011) NMR 999100010537 Macaca sylvanus cf. pliocena North Sea M3 ? 12.1 7.8 7.5 64.5 Reumer et al. (2018) MCSNBS PA11136 Macaca sylvanus cf. pliocena Quecchia Quarry M3 ? 12.1 8.2 — 67.8 Bona et al. (2016) HUJ-ESE UB 314 Macaca sylvanus florentina 'Ubeidiya M3 F 12.03 7.07 6.73 58.8 PRIMO HUJ-ESE UB 329 Macaca sylvanus florentina 'Ubeidiya M3 ? 10.2 7.15 6.0 70.1 PRIMO IGF 10034 Macaca sylvanus florentina Upper Valdarno M3 M 12.3 8.3 7.4 67.5 PRIMO IGF 10035 Macaca sylvanus florentina Upper Valdarno M3 M 12.8 8.2 7.9 64.1 PRIMO IGF 12868 Macaca sylvanus florentina Upper Valdarno M3 F 13.5 8.4 7.8 62.2 PRIMO IGF 13084 Macaca sylvanus florentina Upper Valdarno M3 ? 12.8 7.6 7.1 59.4 PRIMO IGF 13087 Macaca sylvanus florentina Upper Valdarno M3 ? 11.5 6.8 6.4 59.1 PRIMO NMB VA 352 Macaca sylvanus florentina Upper Valdarno M3 M 12.3 8.0 7.2 65.0 PRIMO NMB VA 2058 Macaca sylvanus florentina Upper Valdarno M3 M 12.1 7.5 6.8 62.0 PRIMO MAP 10 Macaca sylvanus florentina Upper Valdarno M3 M 14.5 7.8 7.2 53.8 PRIMO IQW 1980/16566 (Mei. 16087) Macaca sylvanus florentina Untermassfeld M3 F 11.0 7.4 7.0 67.3 Zapfe (2001) IQW 1984/20021 (Mei. 16541) Macaca sylvanus florentina Untermassfeld M3 M 13.4 8.4 7.4 62.7 Zapfe (2001) 18 MUPE GCP-CV4063 Macaca sylvanus florentina Quibas M3 ? 12.1 8.8 7.6 72.7 Alba et al. (2011) UM Q03-Ec-1 Macaca sylvanus florentina Quibas M3 ? 12.0 7.5 6.6 62.5 Alba et al. (2011) MUPE GCP-CV4060 Macaca sylvanus florentina Quibas M3 ? (12.5) 7.8 6.7 62.4 Alba et al. (2011) MSNC Mugello A Macaca sylvanus florentina Mugello M3 ? 12.8 7.3 6.7 57.0 Zanaga (1998) FSL 40137 Macaca sylvanus prisca Montpellier M3 ? 10.8 6.9 6.1 63.9 PRIMO NHMUK-P M11713 Macaca majori Capo Figari M3 ? 10.6 6.5 6.2 61.3 PRIMO IGF 2035 Macaca majori Capo Figari M3 M 10.6 7.0 6.5 66.0 PRIMO NMB TY 5209 Macaca majori Capo Figari M3 ? 9.4 7.1 6.5 75.5 PRIMO NMB TY 5210 Macaca majori Capo Figari M3 ? 10.5 7.0 6.2 66.7 PRIMO NMB TY 5211 Macaca majori Capo Figari M3 ? 11.6 7.0 6.4 60.3 PRIMO NMB TY 5214 Macaca majori Capo Figari M3 ? 9.9 7.3 6.0 73.7 PRIMO NMB TY 5304 Macaca majori Capo Figari M3 ? 9.4 7.4 6.5 78.7 PRIMO NMB TY 12454 Macaca majori Capo Figari M3 ? 10.1 7.4 6.6 73.3 PRIMO NMB TY 12460 Macaca majori Capo Figari M3 M 10.1 7.1 6.7 70.3 PRIMO NMB TY 12469 Macaca majori Capo Figari M3 ? 10.7 7.0 6.7 65.4 PRIMO NMB TY 12470 Macaca majori Capo Figari M3 ? 9.0 7.3 6.2 81.1 PRIMO NMB TY 12514 Macaca majori Capo Figari M3 ? 10.5 7.3 — 69.5 PRIMO GPI-F 1652 Macaca libyca Wadi Natrun M3 ? 11.7 8.3 7.6 70.9 PRIMO YPM 21552 Macaca libyca Wadi Natrun M3 ? 13.2 9.2 8.1 69.7 PRIMO MB Ma 42445 Parapapio ado Laetoli M3 ? 14.9 9.1 8.1 61.1 PRIMO MB Ma 42441 Parapapio ado Laetoli M3 F 14.3 8.7 8.2 60.8 PRIMO MB Ma 42442 (& 42446) Parapapio ado Laetoli M3 M 15.0 10.8 8.9 72.0 PRIMO NHMUK-P M14940 Parapapio ado Laetoli M3 F 12.9 8.1 7.7 62.8 PRIMO NMT LAET 64 Parapapio ado Laetoli M3 ? 13.6 8.4 7.2 61.8 PRIMO NMT LAET 74-223 Parapapio ado Laetoli M3 F 12.9 9.1 8.6 70.5 PRIMO NMT LAET 74-243-4 Parapapio ado Laetoli M3 F 13.9 8.7 8.1 62.6 PRIMO NMT LAET 75-483 Parapapio ado Laetoli M3 ? 14.5 9.1 8.1 62.8 PRIMO NMT LAET 75-1209 Parapapio ado Laetoli M3 F 12.5 7.8 6.5 62.4 PRIMO 19 NMT LAET 75-1390 Parapapio ado Laetoli M3 ? 14.4 9.6 8.2 66.7 PRIMO NMT LAET 75-3035 Parapapio ado Laetoli M3 F 12.9 8.3 8.0 64.3 PRIMO NMT LAET 77-4568 Parapapio ado Laetoli M3 ? 12.6 8.8 8.5 69.8 PRIMO NMT LAET 77-4595 Parapapio ado Laetoli M3 F 14.2 9.5 8.7 66.9 PRIMO KNM-KP 286 Parapapio cf. ado Kanapoi M3 M 12.6 10.1 8.6 80.2 Frost et al. (2020a) KNM-KP 26942 Parapapio cf. ado Kanapoi M3 ? 14.3 9.7 8.1 67.8 Frost et al. (2020a) KNM-KP 29305 Parapapio cf. ado Kanapoi M3 ? 11.0 8.4 7.1 76.4 Frost et al. (2020a) KNM-KP 29306 Parapapio cf. ado Kanapoi M3 ? 12.3 8.6 8.2 69.9 Frost et al. (2020a) KNM-KP 29312 Parapapio cf. ado Kanapoi M3 ? 12.2 9.1 8.5 74.6 Frost et al. (2020a) KNM-KP 30230 Parapapio cf. ado Kanapoi M3 F 12.3 8.3 7.3 67.5 Frost et al. (2020a) KNM-KP 30233 Parapapio cf. ado Kanapoi M3 M 13.0 8.9 8.1 68.5 Frost et al. (2020a) KNM-KP 30399 Parapapio cf. ado Kanapoi M3 ? 13.4 9.5 8.5 70.9 Frost et al. (2020a) KNM-KP 30483 Parapapio cf. ado Kanapoi M3 ? 13.1 9.5 8.4 72.5 Frost et al. (2020a) KNM-KP 30535 Parapapio cf. ado Kanapoi M3 ? 13.4 10.0 8.2 74.6 Frost et al. (2020a) KNM-KP 30538 Parapapio cf. ado Kanapoi M3 F 11.7 9.3 8.4 79.5 Frost et al. (2020a) KNM-KP 32806 Parapapio cf. ado Kanapoi M3 ? 10.9 7.8 7.1 71.6 Frost et al. (2020a) KNM-KP 32817 Parapapio cf. ado Kanapoi M3 ? 11.1 8.1 6.9 73.0 Frost et al. (2020a) KNM-KP 32869 Parapapio cf. ado Kanapoi M3 M (12.19) (8.75) 8.1 (71.8) Frost et al. (2020a) KNM-KP 59889 Parapapio cf. ado Kanapoi M3 ? 12.9 8.5 7.8 65.9 Frost et al. (2020a) KNM-KP 59899 Parapapio cf. ado Kanapoi M3 ? 11.4 8.1 6.7 71.1 Frost et al. (2020a) KNM-ER 3122 Parapapio cf. ado Koobi Fora M3 ? 12.3 7.7 6.9 62.6 Jablonski et al. (2008) KNM-ER 19539 Parapapio cf. ado Koobi Fora M3 ? 11.6 8.1 6.6 69.8 Jablonski et al. (2008) KNM-ER 20437 Parapapio cf. ado Koobi Fora M3 ? 13.3 8.0 7.2 60.2 Jablonski et al. (2008) KNM-ER 36915 Parapapio cf. ado Koobi Fora M3 ? 12.7 8.3 7.6 65.4 Jablonski et al. (2008) KNM-ER 36921 Parapapio cf. ado Koobi Fora M3 ? 12.5 8.0 7.3 64.0 Jablonski et al. (2008) KNM-ER 36970 Parapapio cf. ado Koobi Fora M3 ? 13.2 8.4 7.2 63.6 Jablonski et al. (2008) KNM-ER 36972 Parapapio cf. ado Koobi Fora M3 ? 14.8 9.4 8.4 63.5 Jablonski et al. (2008) KNM-ER 36973 Parapapio cf. ado Koobi Fora M3 ? 13.8 8.0 7.2 58.0 Jablonski et al. (2008) 20 KNM-ER 36987 Parapapio cf. ado Koobi Fora M3 ? 13.2 8.8 7.6 66.7 Jablonski et al. (2008) KNM-ER 37013 Parapapio cf. ado Koobi Fora M3 ? 13.0 8.6 7.6 66.2 Jablonski et al. (2008) KNM-ER 39518 Parapapio cf. ado Koobi Fora M3 ? 11.9 7.6 7.3 63.9 Jablonski et al. (2008) KNM-LT 415 Parapapio lothagamensis Lothagam M3 ? 11.0 7.8 7.0 70.9 Leakey et al. (2003) KNM-LT 23066 Parapapio lothagamensis Lothagam M3 ? 12.5 7.9 7.7 63.2 Leakey et al. (2003) KNM-LT 23090 Parapapio lothagamensis Lothagam M3 F 10.5 7.5 6.6 71.4 Leakey et al. (2003) KNM-LT 23091 Parapapio lothagamensis Lothagam M3 M 11.5 8.0 7.4 69.0 Leakey et al. (2003) KNM-LT 24094 Parapapio lothagamensis Lothagam M3 ? 10.4 6.7 5.9 64.4 Leakey et al. (2003) KNM-LT 24105 Parapapio lothagamensis Lothagam M3 ? 12.5 9.2 7.6 73.6 Leakey et al. (2003) KNM-LT 24117 Parapapio lothagamensis Lothagam M3 ? 13.5 8.4 7.9 62.2 Leakey et al. (2003) KNM-LT 24122 Parapapio lothagamensis Lothagam M3 ? 11.0 7.0 6.0 63.6 Leakey et al. (2003) KNM-LT 24133 Parapapio lothagamensis Lothagam M3 ? 11.9 8.9 7.6 74.8 Leakey et al. (2003) KNM-LT 24135 Parapapio lothagamensis Lothagam M3 ? 10.4 7.4 — 71.2 Leakey et al. (2003) KNM-LT 24136 Parapapio lothagamensis Lothagam M3 M 12.4 7.8 6.8 62.9 Leakey et al. (2003) KNM-LT 24139 Parapapio lothagamensis Lothagam M3 ? 11.1 8.1 7.1 73.0 Leakey et al. (2003) KNM-LT 26391 Parapapio lothagamensis Lothagam M3 ? 10.7 7.2 6.4 67.3 Leakey et al. (2003) KNM-LT 26409 Parapapio lothagamensis Lothagam M3 ? 11.2 7.3 6.4 65.2 Leakey et al. (2003) KNM-LT 28755 Parapapio lothagamensis Lothagam M3 ? 9.7 7.3 6.3 75.3 Leakey et al. (2003) NME GWM-3/P84 Pliopapio alemui Gona M3 F 11.5 7.1 — 61.7 Frost et al. (2020b) NME GWM-3/P164 Pliopapio alemui Gona M3 ? 10.9 7.6 6.0 69.7 Frost et al. (2020b) NME GWM-3w/P183 Pliopapio alemui Gona M3 ? 10.6 7.0 5,9 66.0 Frost et al. (2020b) NME GWM-5m/P125 Pliopapio alemui Gona M3 ? 10.5 6.9 6.2 65.7 Frost et al. (2020b) NME GWM-5sw/P153 Pliopapio alemui Gona M3 ? 11.7 7.6 7.5 65.0 Frost et al. (2020b) NME GWM-5m/P281 Pliopapio alemui Gona M3 ? 10.1 7.4 6.8 73.3 Frost et al. (2020b) NME GWM-5sw/P296 Pliopapio alemui Gona M3 ? 9.5 7.0 6.7 73.7 Frost et al. (2020b) NME GWM-5m/P514 Pliopapio alemui Gona M3 ? 11.9 7.7 6.8 64.7 Frost et al. (2020b) NME GWM-9n/P89 Pliopapio alemui Gona M3 ? 11.5 7.2 6.5 62.6 Frost et al. (2020b) NME GWM-9m/P256 Pliopapio alemui Gona M3 ? 12.1 8.4 7.9 69.4 Frost et al. (2020b) 21 NME GWM-25/27/P100 Pliopapio alemui Gona M3 ? 12.3 7.7 7.0 62.6 Frost et al. (2020b) NME GWM-45/P46 Pliopapio alemui Gona M3 ? 11.6 7.5 6.8 64.7 Frost et al. (2020b) NME GWMS-11/P65 cf. Pliopapio alemui Gona M3 ? 11.4 7.2 7.2 63.2 Frost et al. (2020b) NME ARA-VP 1/816 Pliopapio alemui Aramis M3 F 11.92 7.70 7.00 64.6 PRIMO NME ARA-VP 1/1006 Pliopapio alemui Aramis M3 F 10.47 7.44 6.42 71.0 PRIMO NME ARA-VP 1/1349 Pliopapio alemui Aramis M3 F 9.70 6.79 6.36 70.0 PRIMO NME ARA-VP 1/744 Pliopapio alemui Aramis M3 M 10.75 6.70 6.14 62.3 PRIMO NME ARA-VP 1/1571 Pliopapio alemui Aramis M3 ? 11.20 6.65 6.18 59.3 PRIMO NME ARA-VP 1/1859 Pliopapio alemui Aramis M3 ? 13.14 7.80 7.43 59.4 PRIMO NME ARA-VP 1/190 Pliopapio alemui Aramis M3 ? 11.12 6.94 6.65 62.4 PRIMO NME ARA-VP 1/8 Pliopapio alemui Aramis M3 ? 10.57 6.70 5.88 63.4 PRIMO NME ARA-VP 1/1377 Pliopapio alemui Aramis M3 ? 11.86 7.42 6.76 62.6 PRIMO NME ARA-VP 1/1574 Pliopapio alemui Aramis M3 ? 11.16 6.68 6.75 59.9 PRIMO NME ARA-VP 1/289 Pliopapio alemui Aramis M3 ? 11.73 7.56 7.59 64.5 PRIMO NME ARA-VP 1/1615 Pliopapio alemui Aramis M3 ? 11.68 7.49 6.80 64.1 PRIMO NME ARA-VP 1/9 Pliopapio alemui Aramis M3 ? 11.85 7.17 6.71 60.5 PRIMO NME ARA-VP 1/492 Pliopapio alemui Aramis M3 ? 12.57 7.35 6.92 58.5 PRIMO NME ARA-VP 1/1573 Pliopapio alemui Aramis M3 ? 12.42 7.36 6.68 59.3 PRIMO NME ARA-VP 1/12 Pliopapio alemui Aramis M3 ? 10.49 7.08 6.43 67.5 PRIMO NME ARA-VP 6/8 Pliopapio alemui Aramis M3 ? 10.62 6.48 6.84 61.0 PRIMO NME ARA-VP 6/659 cf. Pliopapio alemui Aramis M3 ? 11.64 7.42 6.12 63.7 PRIMO Abbreviations: F = female; M = male; ? = unknown sex. Institutional acronyms: AMNH-M = American Museum of Natural History (Mammalogy), New York, USA; BGDG = Basque Government official repository institution for the District of Guipúzcoa, San Sebastián, Spain; UNIARQ = Centro de Arqueologia da Universidade de Lisboa, Lisbon, Portugal; Bonifay = Research collection of Dr. Marie-Francoise Bonifay, Marseille, France; BSPG = Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany; FSL = Dept. Sciences de la Terre, Faculté des Sciences, Université Claude-Bernard Lyon I, Lyon, France; GOV = 22 Soprintendenza dell'Abruzzo, Chieti, Italy; GPI = Geologisches-Paläontologisches Institut, Universität Freiburg im Breisgau, Freiburg, Germany; Harlé = Collection of M. Édouard Harlé (not located), Toulouse, France; HUJ-ESE = Hebrew University (Ecology, Systematics & Evolution), Jerusalem, Israel; IAPU-P = Istituto di Antropologia e Paleontologia Umana, Università di Pisa, Italy; IG-R = Istututo di Geologia, Universita di Roma, Italy; IGF = Museo di Storia Naturale (Sezione di Paleontologia e Geologia), Florence, Italy—formerly Istituto di Geologia e Paleontologia, Università di Firenze); IPH = Institut de Paléontologie Humaine, Paris, France; IQW = Senckenberg Research Station of Quaternary Paleontology, Weimar, Germany; KNM = National Museums of Kenya, Kenya; LPHP-S = Laboratori i Paleontologjise Humane Prehistorise, ILP "Luigi Gurakuqi", Shkoder, Albania; MAP = Museo dell'Accademia del Poggio, Montevarchi, Italy; MB = Paläontologisches Museum der Humboldt-Universität, Berlin, Germany; MAC = Museo Anatomia Comparata, Modena, Italy; MAU-T = Museo di Anatomia Umana "Luigi Rolando", Università di Torino, Turin, Italy; MCSNBS = Museo Civico di Scienze Naturali di Brescia, Italy; MCZ = Museum of Comparative Zoology, Harvard, USA; MHNL = Musée des Confluences (formerly Musée d'Histoire Naturelle de Lyon), Lyon, France; MNHN-F = Muséum National d’Histoire Naturelle (Paleontology), Paris, France; MNHN-ZM-AC = Muséum National d’Histoire Naturelle (Zoologie Mammifères – Anatomie Comparée), Paris, France; MNHN-ZM-MO = Muséum National d’Histoire Naturelle (Zoologie Mammifères – Mammifères et Oiseaux), Paris, France; MPC = Museo dei Paleoambienti Sulcitani – E.A. Martel, Carbonia, Sardinia, Italy; MSNC = Museo di Storia naturale e del Calci, Pisa, Italy; MUPE = Museu Paleontològic d’Elx, Elche, Spain; NHMUK-Z = Natural History Museum (Zoology), London, United Kingdom—formerly British Museum (Natural History) [BM(NH)-Z]; NHMUK-P = Natural History Museum (Paleontology), London, United Kingdom—formerly British Museum (Natural History) [BM(NH)-P]; NMB = Naturhistorisches Museum Basel, Switzerland; NME = National Museum of Ethiopia, Addis Ababa, Ethiopia; NMNH = National Museum of Natural History, Smithsonian Institution, Washington D.C., USA; NMP = Národní Muzeum, Prague, Czech Republic; NMR = Natural History Museum Rotterdam, The Netherlands; NMT = National Museum of Tanzania, Dar es Salaam, Tanzania; ORGNAC = Orgnac faunal collection, details uncertain; RGM-L = Rijksmuseum van Geologie en Mineralogie, Leiden, The Netherlands; SIP-V = Servicio de Investigaciones Prehistorica, Diputación Provincial de Valencia, Spain; SMF = Senckenberg 23 Forschungsinstitut und Naturmuseum (Terrestrische Zoologie), Frankfurt am Mein, Germany (formerly Senckenberg Museum Forschungstinstitut); UG = Universidad de Granada, Granada, Spain; UM = Universidad de Murcia, Spain; ZMA = Zoologisch Museum, Universiteit van Amsterdam, The Netherlands. a Data taken from the PRIMO (PRImate Morphometrics Online) database (http://primo.nycep.org), measured by one of the authors (D.M.A.), or available from the literature. 24 SOM Table S2 Descriptive statistics for the various measurements investigated in the tooth loci preserved at Guefaït-4.2, and the z-scores computed for the latter (bolded when significant at p < 0.05, i.e., z >|1.96|) based on the mean and SD of each sample.a I1 H n Mean SD Minimum Maximum z-score Guefaït 1 10.00 — — — — Macaca s. sylvanus 18 8.90 2.29 4.60 12.80 0.48 Macaca s. pliocena 1 10.10 — — — — Macaca s. florentina 2 10.17 1.32 9.23 11.10 — Macaca majori 6 8.42 1.04 6.60 9.70 — Parapapio (cf.) ado 2 11.75 0.64 11.30 12.20 — Pliopapio alemui 6 8.19 1.56 6.40 10.40 — I1 MD n Mean SD Minimum Maximum z-score Guefaït 1 6.80 — — — — Macaca s. sylvanus 23 6.37 0.66 4.95 7.30 0.66 Macaca s. pliocena 1 6.08 — — — — Macaca s. florentina 2 7.13 0.10 7.06 7.20 — Macaca majori 6 6.00 0.19 5.70 6.20 — Parapapio (cf.) ado 2 6.95 0.78 6.40 7.50 — ‘Parapapio' lothagamensis 5 7.36 0.40 6.80 7.90 — Pliopapio alemui 5 5.88 0.52 5.10 6.40 — I1 BL n Mean SD Minimum Maximum z-score Guefaït 1 6.20 — — — — Macaca s. sylvanus 23 5.50 0.48 4.73 6.50 1.47 Macaca s. pliocena 1 5.20 — — — — Macaca s. florentina 2 5.28 0.17 5.16 5.40 — Macaca majori 6 4.73 0.72 4.20 6.10 — Parapapio (cf.) ado 2 5.55 0.35 5.30 5.80 — ‘Parapapio' lothagamensis 5 6.26 0.15 6.10 6.40 — Pliopapio alemui 5 5.82 0.54 5.00 6.30 — I1 BLI n Mean SD Minimum Maximum z-score 25 Guefaït 1 91.18 — — — — Macaca s. sylvanus 23 86.90 7.78 76.09 105.86 0.55 Macaca s. pliocena 1 76.47 — — — — Macaca s. florentina 2 74.05 1.35 73.09 75.00 — Macaca majori 6 79.02 12.86 67.74 103.39 — Parapapio (cf.) ado 2 80.07 3.87 77.33 82.81 — ‘Parapapio' lothagamensis 5 85.25 4.94 77.22 89.71 — Pliopapio alemui 4 99.07 10.51 86.06 111.76 — M2 MD n Mean SD Minimum Maximum z-score Guefaït 1 8.60 — — — — Macaca s. sylvanus 55 9.35 0.80 7.05 11.10 -0.94 Macaca s. (cf. ) pliocena 15 9.82 0.61 8.80 10.80 -2.01 Macaca s. florentina 22 9.61 0.47 8.90 11.00 -2.17 Macaca s. prisca 2 8.4 0.57 8.00 8.80 — Macaca majori 9 8.57 0.39 7.60 8.90 0.07 Macaca libyca 4 9.93 0.67 9.40 10.90 — Parapapio (cf.) ado 24 10.47 0.92 8.80 12.50 -2.03 ‘Parapapio' lothagamensis 10 8.85 0.68 8.10 10.30 -0.37 Pliopapio alemui 6 9.05 0.28 8.74 9.47 — M2 BLm n Mean SD Minimum Maximum z-score Guefaït 1 7.10 — — — — Macaca s. sylvanus 55 7.69 0.66 6.30 9.30 -0.90 Macaca s. (cf. ) pliocena 15 7.74 0.53 6.55 8.50 -1.21 Macaca s. florentina 22 7.58 0.55 6.70 8.50 -0.86 Macaca s. prisca 2 6.80 0 6.80 6.80 — Macaca majori 9 6.93 0.23 6.40 7.20 0.77 Macaca libyca 4 8.48 0.67 7.90 9.40 — Parapapio (cf.) ado 24 8.53 0.76 7.00 9.80 -1.89 ‘Parapapio' lothagamensis 10 7.55 0.72 6.50 9.00 -0.63 Pliopapio alemui 6 6.96 0.48 6.38 7.58 — M2 BLI n Mean SD Minimum Maximum z-score 26 Guefaït 1 82.56 — — — — Macaca s. sylvanus 55 82.55 6.07 70.98 101.11 <0.01 Macaca s. (cf. ) pliocena 15 78.99 5.07 69.31 89.77 0.70 Macaca s. florentina 22 78.81 4.00 72.48 87.37 0.94 Macaca s. prisca 2 81.14 5.47 77.27 85.00 — Macaca majori 9 80.83 2.08 78.16 84.21 0.84 Macaca libyca 4 85.78 9.79 72.48 95.92 — Parapapio (cf.) ado 24 81.73 6.39 70.71 100.00 0.13 ‘Parapapio' lothagamensis 10 85.34 5.41 77.89 95.45 -0.51 Pliopapio alemui 6 76.80 3.45 72.77 81.30 — M3 MD n Mean SD Minimum Maximum z-score Guefaït 1 10.00 — — — — Macaca s. sylvanus 51 11.61 1.07 9.34 14.60 -1.50 Macaca s. (cf. ) pliocena 13 12.78 0.81 11.70 14.40 -3.41 Macaca s. florentina 16 12.36 1.01 10.20 14.50 -2.35 Macaca s. prisca 1 10.80 — — — — Macaca majori 12 10.20 0.71 9.00 11.60 -0.28 Macaca libyca 2 12.45 1.06 11.70 13.20 — Parapapio (cf.) ado 40 12.97 1.08 10.90 15.00 -2.75 ‘Parapapio’ lothagamensis 15 11.36 1.03 9.70 13.50 -1.33 Pliopapio aleumi 31 11.31 0.84 9.50 13.14 -1.56 M3 BLm n Mean SD Minimum Maximum z-score Guefaït 1 7.60 — — — — Macaca s. sylvanus 51 8.12 0.87 6.41 11.40 -0.60 Macaca s. (cf. ) pliocena 13 8.20 0.62 7.30 9.40 -0.96 Macaca s. florentina 16 7.75 0.56 6.80 8.80 -0.27 Macaca s. prisca 1 6.90 — — — — Macaca majori 12 7.12 0.25 6.50 7.40 1.92 Macaca libyca 2 8.75 0.64 8.30 9.20 — Parapapio (cf.) ado 40 8.74 0.73 7.60 10.80 -1.56 ‘Parapapio’ lothagamensis 15 7.77 0.69 6.70 9.20 -0.24 27 Pliopapio alemui 31 7.26 0.42 6.48 8.40 0.81 M3 BLI n Mean SD Minimum Maximum z-score Guefaït 1 76.00 — — — — Macaca s. sylvanus 51 70.11 6.39 56.06 100.88 0.92 Macaca s. (cf. ) pliocena 13 64.17 2.46 60.33 68.38 4.81 Macaca s. florentina 16 62.91 4.88 53.79 72.73 2.68 Macaca s. prisca 1 63.89 — — — — Macaca majori 12 70.17 6.54 60.34 81.11 0.89 Macaca libyca 2 70.32 0.88 69.70 70.94 — Parapapio (cf.) ado 40 67.60 5.32 57.97 80.16 1.58 ‘Parapapio’ lothagamensis 15 68.53 4.66 62.22 75.26 1.60 Pliopapio alemui 31 64.40 4.06 58.50 73.68 2.86 Abbreviations: H = maximum recorded crown height (mm); BL = buccolingual (or labiolingual) breadth (mm); BLm = buccolingual breadth across the mesial lophid (mm); BLI = breadth/length index (%; using BLm for molars); MD = mesiodistal length (mm). a z-scores for I1 measurements are only provided based on M. s. sylvanus, as the small samples available for the remaining taxa would make render them unreliable. For the same reason, z-scores for the molars are not provided based on M. libyca, M. s. prisca, or Pl. alemui M2, while those for M. majori and ‘P.’ lothagamensis should be interpreted with care. 28 SOM Table S3 ANOVA results for the metrical comparisons of taxa included in the comparative sample (see SOM Tables S1 and S2)a. Significant results at p < 0.05 are bolded. See SOM Table S4 for the results of pairwise comparisons. Variable F p I1 H 1.59 0.203 I1 MD 5.25 0.001 I1 BL 5.80 <0.001 I1 BLI 3.74 0.008 M2 MD 10.82 <0.001 M2 BL 9.94 <0.001 M2 BLI 3.08 0.005 M3 MD 16.58 <0.001 M3 BL 15.90 <0.001 M3 BLI 6.20 <0.001 Abbreviations: H = maximum recorded crown height (mm); BL = buccolingual (or labiolingual) breadth (mm); BLI = breadth/length index (%); MD = mesiodistal length (mm). a Macaca s. prisca was excluded due to insufficient sample size. 29 SOM Table S4 Tukey’s pairwise post hoc tests among the comparative sample (see SOM Tables S1 and S2)a. The Studentized range statistic Q is given below the diagonal, whereas p-values are given above. Significant differences at p < 0.05 are bolded. I1 H M. s. sylvanus M. s. florentina M. majori P. (cf.) ado Pl. alemui M. s. sylvanus — 0.902 0.984 0.302 0.936 M. s. florentina 1.243 — 0.802 0.923 0.723 M. majori 0.746 1.565 — 0.243 1.000 P. (cf.) ado 2.797 1.159 2.984 — 0.189 Pl. alemui 1.097 1.768 0.287 — I1 MD M. s. sylvanus M. s. florentina M. majori P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.469 0.721 0.734 0.013 0.528 M. s. florentina 2.568 — 0.174 1.000 0.997 0.119 M. majori 1.979 3.430 — 0.341 0.004 0.999 P. (cf.) ado 1.963 0.446 2.883 — 0.954 0.246 ‘P.' lothagamensis 4.992 0.681 5.566 1.214 — 0.003 Pl. alemui 2.431 3.696 0.483 3.163 5.791 — I1 BL M. s. sylvanus M. s. florentina M. majori P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.990 0.018 1.000 0.038 0.774 M. s. florentina 0.864 — 0.750 0.994 0.190 0.777 M. majori 4.816 1.924 — 0.344 0.001 0.010 P. (cf.) ado 0.188 0.776 2.874 — 0.525 0.986 30 ‘P.' lothagamensis 4.415 3.366 7.244 2.438 — 0.719 Pl. alemui 1.853 1.855 5.156 0.927 1.999 — I1 BLI M. s. sylvanus M. s. florentina M. majori P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.335 0.351 0.882 0.999 0.114 M. s. florentina 2.899 — 0.979 0.980 0.620 0.019 M. majori 2.859 1.013 — 1.000 0.830 0.010 P. (cf.) ado 1.541 1.002 0.214 — 0.977 0.129 ‘P.' lothagamensis 0.555 2.227 1.712 1.030 — 0.176 Pl. alemui 3.733 4.803 5.163 3.647 3.423 — M2 MD M. s. sylvanus M. s. (cf.) pliocena M. s. florentina M. majori M. libyca P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.346 0.834 0.061 0.785 <0.001 0.468 0.977 M. s. (cf.) pliocena 3.139 — 0.990 0.002 1.000 0.114 0.027 0.352 M. s. florentina 2.046 1.189 — 0.009 0.993 0.002 0.108 0.683 M. majori 4.247 5.790 5.164 — 0.044 <0.001 0.991 0.915 M. libyca 2.177 0.378 1.124 4.417 — 0.854 0.194 0.562 P. (cf.) ado 8.993 3.906 5.705 9.535 1.987 — <0.001 <0.001 ‘P.' lothagamensis 2.867 4.654 3.937 1.179 3.571 8.463 — 1.000 Pl. alemui 1.386 3.126 2.414 1.767 2.669 6.125 0.755 — M2 BLm M. s. sylvanus M. s. (cf.) pliocena M. s. florentina M. majori M. libyca P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 1.000 0.995 0.020 0.267 <0.001 0.997 0.122 M. s. (cf.) pliocena 0.341 — 0.994 0.052 0.451 0.005 0.995 0.174 M. s. florentina 1.065 1.098 — 0.163 0.164 <0.001 1.000 0.396 31 M. majori 4.802 4.331 3.685 — 0.002 <0.001 0.391 1 M. libyca 3.346 2.903 3.682 5.757 — 1.000 0.218 0.007 P. (cf.) ado 7.616 5.359 7.222 9.185 0.241 — 0.002 <0.001 ‘P.' lothagamensis 0.967 1.057 0.168 3.035 3.491 5.833 — 0.608 Pl. alemui 3.863 3.644 3.023 0.125 5.258 7.721 2.573 — M2 BLI M. s. sylvanus M. s. (cf.) pliocena M. s. florentina M. majori M. libyca P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.367 0.146 0.989 0.952 0.999 0.828 0.253 M. s. (cf.) pliocena 3.090 — 1.000 0.994 0.383 0.814 0.107 0.992 M. s. florentina 3.754 0.140 — 0.984 0.303 0.641 0.051 0.994 M. majori 1.208 1.104 1.296 — 0.820 1.000 0.649 0.869 M. libyca 1.581 3.054 3.249 2.085 — 0.880 1.000 0.208 P. (cf.) ado 0.851 2.102 2.503 0.579 1.901 — 0.623 0.531 ‘P.' lothagamensis 2.061 3.940 4.341 2.488 0.186 2.436 — 0.068 Pl. alemui 3.386 1.150 1.104 1.938 3.524 2.733 4.191 — M3 MD M. s. sylvanus M. s. (cf.) pliocena M. s. florentina M. majori M. libyca P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.005 0.149 <0.001 0.938 <0.001 0.989 0.876 M. s. (cf.) pliocena 5.356 — 0.953 <0.001 1.000 0.999 0.005 <0.001 M. s. florentina 3.751 1.578 — <0.001 1.000 0.447 0.096 0.015 M. majori 6.282 9.192 8.093 — 0.065 <0.001 0.057 0.027 M. libyca 1.661 0.615 0.163 4.207 — 0.996 0.826 0.760 P. (cf.) ado 9.166 0.851 2.910 12.010 1.019 — <0.001 <0.001 ‘P.' lothagamensis 1.223 5.339 3.991 4.277 2.068 7.580 — 1.000 32 Pl. alemui 1.915 6.356 4.909 4.646 2.239 9.913 0.245 — M3 BLm M. s. sylvanus M. s. (cf.) pliocena M. s. florentina M. majori M. libyca P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 1.000 0.572 0.001 0.907 0.001 0.655 <0.001 M. s. (cf.) pliocena 0.536 — 0.653 0.003 0.965 0.220 0.709 0.001 M. s. florentina 2.647 2.477 — 0.238 0.525 <0.001 1.000 0.283 M. majori 6.441 5.578 3.425 — 0.044 <0.001 0.226 0.999 M. libyca 1.804 1.493 2.745 4.408 — 1.000 0.550 0.064 P. (cf.) ado 6.046 3.478 6.881 10.459 0.032 — <0.001 <0.001 ‘P.' lothagamensis 2.474 2.357 0.088 3.459 2.692 6.618 — 0.272 Pl. alemui 7.785 5.870 3.296 0.863 4.212 12.750 3.327 — M3 BLI M. s. sylvanus M. s. (cf.) pliocena M. s. florentina M. majori M. libyca P. (cf.) ado ‘P.’ lothagamensis Pl. alemui M. s. sylvanus — 0.009 <0.001 1.000 1.000 0.336 0.973 <0.001 M. s. (cf.) pliocena 5.085 — 0.998 0.097 0.793 0.471 0.378 1.000 M. s. florentina 6.687 0.900 — 0.010 0.579 0.063 0.070 0.985 M. majori 0.052 3.989 5.062 — 1.000 0.822 0.993 0.035 M. libyca 0.081 2.157 2.632 0.054 — 0.997 1.000 0.790 P. (cf.) ado 3.162 2.857 4.220 2.080 1.002 — 0.999 0.194 ‘P.' lothagamensis 1.427 3.063 4.165 1.126 0.634 0.821 — 0.213 Pl. alemui 6.676 0.180 1.286 4.522 2.164 3.563 3.501 — Abbreviations: BL = buccolingual (or labiolingual) breadth (mm); BLm = buccolingual breadth across the mesial lophid (mm); BLI = breadth/length index (%) (using BLm for molars); MD = mesiodistal length (mm). a Macaca s. prisca was excluded due to insufficient sample size. 33 SOM File S1. Virtual model of GFT4.2’18-1-Q14-70, left I1 crown of Macaca cf. sylvanus from Guefaït-4.2, available in PLY format from MorphoSource at https://doi.org/10.17602/M2/M158035 (unique identifier: ark:/87602/m4/M158035). A digital rendering in lingual view is depicted. SOM File S2. Virtual model of GFT4.2’18-1-Q14-13, right M1 lingual crown fragment of Macaca cf. sylvanus from Guefaït-4.2, available in PLY format from MorphoSource at https://doi.org/10.17602/M2/M158036 (unique identifier: ark:/87602/m4/M158036). A digital rendering in occlusal view is depicted. 34 SOM File S3. Virtual model of GFT4.2’18-1-P13-191, right M2 or M3 mesial germ fragment of Macaca cf. sylvanus from Guefaït-4.2 available in PLY format from MorphoSource at https://doi.org/10.17602/M2/M158037 (unique identifier: ark:/87602/m4/M158037). A digital rendering in occlusal view is depicted. SOM File S4. Virtual model of GFT4.2’19-1-R15-45, right M2 distal germ fragment of Macaca cf. sylvanus from Guefaït-4.2 available in PLY format from MorphoSource at https://doi.org/10.17602/M2/M158038 (unique identifier: ark:/87602/m4/M158038). A digital rendering in occlusal view is depicted. 35 SOM File S5. Virtual model of GFT4.2’19-1-S15-65, right M2 crown with distal root of Macaca cf. sylvanus from Guefaït-4.2 available in PLY format from MorphoSource at https://doi.org/10.17602/M2/M158291 (unique identifier: ark:/87602/m4/M158291). A digital rendering in occlusal view is depicted. SOM File S6. Virtual model of GFT4.2’19-1-R13-63, left M3 crown of Macaca cf. sylvanus from Guefaït-4.2 available in PLY format from MorphoSource at https://doi.org/10.17602/M2/M158292 (unique identifier: ark:/87602/m4/M158292). A digital rendering in occlusal view is depicted. 36 SOM References Alba, D.M., Carlos Calero, J.A., Mancheño, M.Á., Montoya, P., Morales, J., Rook, L., 2011. Fossil remains of Macaca sylvanus florentina (Cocchi, 1872) (Primates, Cercopithecidae) from the Early Pleistocene of Quibas (Murcia, Spain). J. Hum. Evol. 61, 703–718. Alba, D. M., Daura, J., Sanz, M., Santos, E., Yagüe, A. S., Delson, E., Zilhão, J., 2019. New macaque remains from the Middle Pleistocene of Gruta da Aroeira (Almonda karst system, Portugal). J. Hum. Evol. 131, 40–47. Bona, F., Bellucci, L., Casali, D., Schirolli, P., Sardella, R., 2016. Macaca sylvanus Linnaeus 1758 from the Middle Pleistocene of Quecchia quarry (Brescia, Northern Italy). 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