This is the accepted version of the journal article: Urciuoli, Alessandro; Alba, David M. «Systematics of Miocene apes : State of the art of a neverending controversy». Journal of Human Evolution, Vol. 175 (February 2023), art. 103309. DOI 10.1016/j.jhevol.2022.103309 This version is available at https://ddd.uab.cat/record/271462 under the terms of the license 1 Systematics of Miocene apes: State of the art of a neverending controversy 2 3 Alessandro Urciuoli1,2,3 & David M. Alba3,* 4 1 5 Universitat Autònoma de Barcelona, Campus de la UAB, 08193 Cerdanyola del Vallès, 6 Barcelona, Spain. 2 7 Division of Palaeoanthropology, Senckenberg Research Institute and Natural History 8 Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt am Main, Germany. 3 9 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, 10 Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, 11 Spain. 12 13 *Corresponding author. 14 E-mail address: david.alba@icp.cat (D.M. Alba). 1 Systematics of Miocene apes: State of the art of a neverending controversy 2 3 Abstract 4 Hominoids diverged from cercopithecoids during the Oligocene in Afro-Arabia, initially 5 radiating in that continent and subsequently dispersing into Eurasia. From the Late Miocene 6 onward, the geographic range of hominoids progressively shrank, except for hominins, 7 which dispersed out of Africa during the Pleistocene. Although the overall picture of 8 hominoid evolution is clear based on available fossil evidence, many uncertainties persist 9 regarding the phylogeny and paleobiogeography of Miocene apes (nonhominin hominoids), 10 owing to their sparse record, pervasive homoplasy, and the decimated current diversity of 11 this group. We review Miocene ape systematics and evolution by focusing on the most 12 parsimonious cladograms published during the last decade. First, we provide a historical 13 account of the progress made in Miocene ape phylogeny and paleobiogeography, report an 14 updated classification of Miocene apes, and provide a list of Miocene ape species-locality 15 occurrences together with an analysis of their paleobiodiversity dynamics. Second, we 16 discuss various critical issues of Miocene ape phylogeny and paleobiogeography (hylobatid 17 and crown hominid origins, plus the relationships of Oreopithecus) in the light of the highly 18 divergent results obtained from cladistic analyses of craniodental and postcranial characters 19 separately. We conclude that cladistic efforts to disentangle Miocene ape phylogeny are 20 potentially biased by a long-branch attraction problem caused by the numerous postcranial 21 similarities shared between hylobatids and hominids—despite the increasingly held view 22 that they are likely homoplastic to a large extent, as illustrated by Sivapithecus and 23 Pierolapithecus—and further aggravated by abundant missing data owing to incomplete 24 preservation. Finally, we argue that—besides the recovery of additional fossils, the retrieval 1 25 of paleoproteomic data, and a better integration between cladistics and geometric 26 morphometrics—Miocene ape phylogenetics should take advantage of total-evidence (tip- 27 dating) Bayesian methods of phylogenetic inference combining morphologic, molecular, and 28 chronostratigraphic data. This would hopefully help ascertain whether hylobatid divergence 29 was more basal than currently supported. 30 31 Keywords: Hominoidea; Evolution; Taxonomy; Phylogeny; Cladistics; Homoplasy. 32 33 1. Introduction 34 1.1. What, if anything, is a Miocene ape? 35 The term ‘ape’ is sometimes used as a synonym of ‘hominoid’—i.e., a member of the 36 superfamily Hominoidea, which includes the families Hylobatidae (gibbons and siamang) and 37 Hominidae (orangutans, gorillas, chimpanzees, and humans; Groves, 2017), plus their extinct 38 relatives. However, following the most common usage of ‘apes’ as opposed to both 39 ‘monkeys’ and ‘humans’ (e.g., Alba, 2012; Tuttle, 2014; Andrews, 2020; Almécija et al., 40 2021), we restrict the former term to hominoids exclusive of the human lineage (i.e., 41 nonhominin hominoids). Originally, the term ‘ape’ broadly referred to all nonhuman 42 anthropoids, so that nonhuman hominoids were referred to as ‘manlike apes’ (Huxley, 1863) 43 or ‘anthropomorphous apes’ (Darwin, 1871; Huxley, 1872). Huxley (1872) formalized the 44 term ‘anthropomorph’, subsequently used by other authors (e.g., Pocock, 1926; Delson, 45 1977; Szalay and Delson, 1979), but currently in disuse in the English literature. Hylobatids 46 and nonhominin hominids are customarily referred to as ‘lesser apes’ and ‘great apes’, 47 respectively, in allusion to their size differences (Tuttle, 2014). Extant hylobatids include 48 more than a dozen species classified in four genera, while hominids similarly include four 2 49 genera but a lower number of species (Groves, 2001, 2017). The restricted diversity of 50 extant hominoids contrasts with that of both extant Old World monkeys and extinct apes. 51 The latter attained a much wider geographic distribution (including Europe and mainland 52 Asia) during the Miocene (e.g., Alba, 2012; Fleagle, 2013; Begun, 2015). As a result of the 53 decimated current genus diversity of hominoids, making sense of their evolutionary history 54 constitutes a monumental challenge from both adaptive and phylogenetic viewpoints (see 55 review in Almécija et al., 2021). 56 This review focuses on apes recorded from the Miocene—the first geological epoch of the 57 Neogene period, being formally divided into Early (23.04–15.99 Ma), Middle (15.99–11.65 58 Ma), and Late (11.65–5.33 Ma) Miocene (Raffi et al., 2020). This epoch witnessed important 59 environmental and biotic changes (Zachos et al., 2001; Blois and Hadly, 2009; Raffi et al., 60 2020). The closure of the Tethys Seaway, due to the collision of the Afro-Arabian and 61 Eurasian plates, enabled intermittent intercontinental dispersals through the Middle East 62 from ~19 Ma onward (Harzhauser et al., 2007), although they were temporarily interrupted 63 during the Langhian transgression at the beginning of the Middle Miocene (~16 Ma; Rögl, 64 1999). The Mid-Miocene Climatic Optimum, a global warming event that peaked ~17–15 Ma, 65 was followed by the Middle Miocene Climate Transition, a stepwise cooling phase that 66 continued throughout the Late Miocene and had a profound impact on terrestrial 67 ecosystems and mammalian communities (Flower and Kennett, 1994; Zachos et al., 2001; 68 Kürschner et al., 2008; Foster et al., 2012; Pound et al., 2012). The geographic spread of 69 woodland and savanna biomes throughout the Old World, and the associated Pikermian 70 chronofauna adapted to more open and arid environments, started around the Middle to 71 Late Miocene transition and peaked at ~7.5 Ma (Eronen et al., 2009; Kaya et al., 2018). 72 Toward the end of the Miocene, beginning at ~6 Ma, a combination of tectonic and 3 73 glacioeustatic factors repeatedly isolated the Mediterranean Sea from the Atlantic Ocean 74 during the Messinian Salinity Crisis (Krijgsman et al., 1999), which favored the spread of 75 open landscapes around the Mediterranean and the establishment of additional dispersal 76 routes between Europe and Africa (Gibert et al., 2013). 77 As for many other groups, the factors outlined above played a major role in shaping 78 hominoid evolution and adaptation (Andrews, 1992, 1996; Andrews and Bernor, 1999; 79 Andrews and Kelley, 2007). Several books (Tuttle, 2014; Andrews, 2015; Begun, 2016) and 80 reviews (Wood and Harrison, 2011; Begun, 2013, 2015; Andrews, 2020; Almécija et al., 2021) 81 have been devoted to Miocene apes during the last decade, and the general picture is quite 82 clear. Hominoids originated in Africa during the late Oligocene, experienced a first radiation 83 in that continent during the Early and Middle Miocene, and later dispersed into Eurasia, 84 where they experienced a second radiation during the Middle to Late Miocene. 85 Subsequently, from the Late Miocene onward, many hominoid genera went extinct and the 86 geographic distribution of hominoids progressively shrank to equatorial Africa and 87 southeastern Asia—with the remarkable exception of members of the human lineage, which 88 radiated during the Plio-Pleistocene and ultimately dispersed throughout the globe. 89 Nevertheless, many uncertainties still persist, particularly regarding the origin of hylobatids 90 and crown hominids (Almécija et al., 2021). 91 92 1.2. Taxonomic scope and aims of this review 93 This review aims to synthesize current knowledge of Miocene ape diversity as well as to 94 critically review their taxonomy, phylogeny, and paleobiogeography in light of the cladistic 95 analyses published during the last decade, with emphasis on hylobatid and hominid origins. 96 A first section with the necessary historical background is followed by an updated 4 97 classification of Miocene apes and an analysis of their paleobiodiversity dynamics. Various 98 controversial issues (the origin of hylobatids, the relationships of Oreopithecus, and the 99 pongine–hominine divergence) are then discussed in the light of phylogenetic uncertainties 100 highlighted by the contradictory cladistic results obtained from craniodental and postcranial 101 data separately. We finally discuss future directions of research with emphasis on 102 phylogenetic inference methods. 103 Besides Oligocene and Plio-Pleistocene apes, which are excluded from this review by 104 definition, Early and Middle Miocene small-bodied catarrhines from Africa of uncertain 105 affinities have also been left out. These include dendropithecids, which have been variously 106 considered stem catarrhines (Harrison, 2010a, 2013; Nengo et al., 2017; Gilbert et al., 2020a) 107 or stem hominoids (Rae, 1999, 2004; Zalmout et al., 2010; Alba et al., 2015; Begun, 2015; 108 Rossie and Hill, 2018), because we consider that currently available evidence leans against 109 considering them hominoids—albeit recognizing that more complete remains would be 110 required to more conclusively assess their systematic position (see Section 3.5). 111 Two small-bodied genera from Eurasia of debated affinities have also been excluded. Kapi 112 ramnagarensis Gilbert et al., 2020a, based on an isolated M3 from the Middle Miocene 113 (13.8–12.5 Ma) of India, was originally recovered as a stem hylobatid (Gilbert et al., 2020a), 114 but subsequently reinterpreted as a pliopithecoid (Ji et al., 2022). Similarly, Pliobates 115 cataloniae Alba et al., 2015 from the Middle/Late Miocene (11.6 Ma) of Spain, known on the 116 basis of a partial skeleton, was originally considered a stem hominoid (Alba et al., 2015) but 117 alternatively interpreted as a possible pliopithecoid (Benefit and McCrossin, 2015; Nengo et 118 al., 2017; Gilbert et al., 2020a, 2020b). Pliobates displays a mosaic of plesiomorphic (stem 119 catarrhine-like) and derived (crown hominoid-like) features (Alba et al., 2015; Bouchet et al., 5 120 2021), but work in progress by the authors supports the hypothesis that it is a stem 121 catarrhine postcranially convergent with hominoids. 122 The hominin status of the Late Miocene genera Ardipithecus White et al., 1995 (~5.8–4.4 123 Ma; White et al., 1994, 2009; Haile-Selassie, 2001; Haile-Selassie et al., 2004, 2009), Orrorin 124 Senut et al., 2001 (Pickford et al., 2002; Gommery and Senut, 2006; Almécija et al., 2013), 125 and Sahelanthropus Brunet et al., 2002 (~7 Ma; Zollikofer et al., 2005; Guy et al., 2005; 126 Macchiarelli et al., 2020; Daver et al., 2022) has sometimes been questioned (Wolpoff et al., 127 2002; Macchiarelli et al., 2020; Wood and Harrison, 2011). However, here these genera have 128 been excluded based on the general view (e.g., Harcourt-Smith, 2010; Simpson, 2010, 2013; 129 Pugh, 2022)—further supported by most recent cladistic analyses (Mongle et al., 2019; Pugh, 130 2020)—that they are early hominins. 131 132 2. Historical background 133 For the purposes of the present review, we distinguish three (slightly overlapping) phases 134 of Miocene ape research: (1) from the pioneering works of earliest evolutionists until the 135 1970s; (2) a turmoil phase characterized by the molecular revolution and the cladistic 136 paradigm shift (1960s–1970s); and (3) a modern phase, characterized by an acceleration of 137 fossil discoveries and the regular application of computer-assisted methods of phylogenetic 138 inference. 139 140 2.1. From Darwin to the Ramapithecus debate 141 Ever since Darwin (and Dryopithecus) More than a century and a half ago, evidence on 142 extinct apes was very meager. However, largely based on Huxley’s (1863) studies, Darwin 143 (1871) hypothesized an African origin for the human lineage. Given the scarcity of fossil apes 6 144 known by then, Darwin (1871) recognized that the aforementioned hypothesis was little 145 more than a well-informed guess. Indeed, the discovery of Dryopithecus fontani Lartet, 1856 146 in the Miocene of France already indicated that paleobiogeographic scenarios of ape and 147 human evolution must be more complex than implied by the geographic distribution of 148 extant apes. Darwin’s (1871) hypothesis did not gain general acceptance during the 149 following decades, owing to several factors besides the initial dearth of fossil humans in 150 Africa. Even after the discovery of australopiths in South Africa (Dart, 1925), the infamous 151 Piltdown hoax from England (Dawson and Smith Woodward, 1913, 1914) contributed— 152 among other factors—to divert paleoanthropologists’ attention from Africa (Tobias, 1985, 153 1992; Lewin, 1987). The Piltdown Man became eventually sidelined in the 1940s (Harrison 154 and Howells, 2007), before the fraud was uncovered a decade later (Weiner et al., 1953a, 155 1953b). However, racist prejudices by European researchers arguably played a more 156 important role in the initial dismissal of australopiths as early human ancestors (Lewin, 1987; 157 Bowler, 1992). 158 A massacre of Miocene apes Fossil evidence from Europe, Asia, and Africa accumulated until 159 Simons and Pilbeam (1965) published a highly influential taxonomic revision of fossil apes. 160 During the 1960s, systematic thinking was dominated by Simpson’s evolutionary 161 systematics, which accepted paraphyletic and even polyphyletic taxa and was strongly 162 biased in favor of taxonomic lumping—with the proliferation of genus and species names 163 being debunked as ‘typological thinking’ (Cartmill, 2018). Following Mayr’s (1950) influential 164 contribution, Le Gros Clark (1955: 18) qualified “the somewhat arbitrary multiplication of 165 genera and species” one of “the more vexing taxonomic problems in Primate paleontology”. 166 Simons (1963) adhered to such views and put them into practice in Simons and Pilbeam’s 167 (1965) revision, which showed a marked lumping tendency—a “massacre of Miocene ape 7 168 taxa” (Cartmill, 2018: 680). They shoehorned most fossil large-bodied hominoid remains into 169 just seven species within the genus Dryopithecus Lartet, 1856, divided in three subgenera: 170 Dryopithecus (Proconsul) Hopwood, 1933 from Africa, Dryopithecus (Sivapithecus) Pilgrim, 171 1910 from Asia, and Dryopithecus s.s. from Europe. Only Gigantopithecus von Koenigswald, 172 1935 (considered an aberrant late offshoot of dryopithecines) and Ramapithecus Lewis, 173 1934 (considered an early member of the human lineage) were spared by Simons and 174 Pilbeam (1965) from being subsumed into Dryopithecus. 175 The rise and fall of Ramapithecus The notion that humans and apes (or at least African great 176 apes) diverged from a common stock deep in the Miocene or even earlier was widespread 177 during the 20th century well into the 1960s (e.g., Gregory, 1916, 1927; Keith, 1925; Osborn, 178 1930; Simpson, 1949; Leakey, 1953; Le Gros Clark, 1955, 1959)—albeit with widely divergent 179 viewpoints between Gregory and Osborn (see review in Lewin, 1987). Following Lewis’s 180 (1934) original suggestion, Simons (1961, 1964) resurrected Ramapithecus as an early 181 representative of the human lineage, contrasting to the previously prevailing view that it 182 was a dryopithecine (Simpson, 1963). By that time, Leakey (1961) made a similar proposal 183 for Kenyapithecus wickeri Leakey, 1961 from Africa, subsequently criticizing Simons and 184 Pilbeam’s (1965) proposed synonymy with Ramapithecus punjabicus (Pilgrim, 1910) as an 185 "extreme example of taxonomic lumping” (Leakey, 1967: 155). 186 As characterized in the 1960s, Ramapithecus fulfilled the expectations for a fossil human 187 relative (parabolic dental arcade, small upper incisors and canines, and orthognathous face; 188 Simons, 1961, 1964). This contention led to the so-called “Ramapithecus debate,” which 189 “had a profound and lasting effect on paleoanthropology” (Ward, 1997a: 270). According to 190 Simons and Pilbeam (1965), Ramapithecus might have evolved from an early species of 191 Dryopithecus and bounded the divergence between humans and apes to not later than 14 8 192 Ma, while extant African apes might have originated from later species of Dryopithecus, and 193 orangutans from an even older dryopithecine ancestor back in the Oligocene. Pilbeam (1966, 194 1969) even hypothesized ancestor–descendant relationships between extant great ape 195 genera and different Miocene ape species. As epitomized by the same author three decades 196 later (Pilbeam, 1997: 13–14): “When the number of taxa was limited, the number of 197 morphological characters small, phylogenetic analysis not rigorous, and when molecular 198 clocks could safely be ignored, there were few impediments to tracing extant lineages well 199 back into the Neogene, or even earlier.” 200 The status quo set forth by Simons and Pilbeam’s (1965), particularly regarding 201 Ramapithecus (see also Pilbeam and Simons, 1965; Pilbeam, 1966), was contested by other 202 paleontologists during the following decade (see below) but temporarily led to a mainstream 203 consensus that drastically differs from current views on hominin origins. This is illustrated by 204 Campbell and Bernor’s (1976) review of ape evolution, explicitly aimed to evaluate the place 205 of origin of the human lineage. The views summarized by these authors still hold nowadays 206 regarding hominoid origins in Africa and their subsequent dispersal into Eurasia ~16 Ma. 207 However, their discussion about hominin origins was colored by the wide geographic range 208 still attributed by then to Ramapithecus, concluding that “Dryopithecines in either Africa or 209 Eurasia could have given rise to early Hominidae [currently Hominini]”, such that “neither 210 continent can be precluded as the place of origin” (Campbell and Bernor, 1976: 441). Toward 211 the end of the 1970s, Simons (1977) maintained unaltered his opinion about Ramapithecus. 212 Pilbeam and colleagues (Pilbeam et al., 1977, 1980; Pilbeam, 1979) were more amenable to 213 different interpretations—he subsequently admitted having doubts since the mid 1970s 214 (Pilbeam, 1983)—but still highlighted the distinctiveness of Ramapithecus. 9 215 In contrast, other authors voiced multiple criticisms against the interpretation of 216 Ramapithecus as an early member of the human lineage. Ramapithecus wickeri was 217 resurrected for the African remains (Andrews, 1971) and it was shown that this species 218 lacked a rounded, human-like dental arcade (Walker and Andrews, 1973; Frayer, 1976; 219 Greenfield, 1978). Von Koenigswald (1973) resurrected Kenyapithecus Leakey, 1961 for this 220 species and considered it an ape, while Greenfield (1974) argued that some remains from 221 Asia also belonged to dryopithecines and criticized the purported status of Ramapithecus as 222 a human ancestor. A more thorough rebuttal was provided by Frayer (1976), then Pilbeam 223 (1978) acknowledged that Ramapithecus lacked a parabolic arcade, and finally Greenfield 224 (1979) concluded that Ramapithecus is a junior subjective synonym of Sivapithecus—leading 225 him to favor a late divergence for the human lineage (Greenfield, 1980). During the 1980s, 226 only a few researchers (e.g., Kay, 1982; Kay and Simons, 1983) still supported ‘hominid’ 227 status of these taxa despite accepting their synonymy. 228 229 2.2. The molecular revolution and the paradigm shift of cladistics 230 The molecular revolution Early studies of hominoid phylogeny based on serological data 231 indicated that humans are more closely related to African apes than to orangutans 232 (Zuckerkandl et al., 1960; Goodman, 1962a, 1962b, 1963). Although this was recognized by 233 Simons and Pilbeam (1965), they did not see it as a challenge for an early divergence of the 234 human lineage. Studies based on protein data subsequently supported a much more recent 235 divergence between humans and African apes (Sarich and Wilson, 1967; Wilson and Sarich, 236 1969; Goodman et al., 1971; Goodman, 1974). However, paleoanthropologists were 237 unwilling to accept such a late divergence (e.g., Simons, 1969; Leakey, 1970; Uzzell and 238 Pilbeam, 1971) with just few exceptions (Washburn, 1967). Only the end of the 10 239 Ramapithecus debate enabled them to more widely reject the long held assumption that 240 apes were closely related and the human lineage divergent (Pilbeam, 1983). 241 In the following decade, molecular studies conclusively settled the identity of humans' 242 closest relatives. Initial results based on mitochondrial DNA (Ferris et al., 1981) were 243 ambiguous, but further analyses based on proteins (Goodman et al., 1983) and DNA 244 hybridization (Sibley and Ahlquist, 1984; Caccone and Powell, 1989; Sibley et al., 1990) 245 strongly supported a sister-taxon relationship between humans and chimpanzees. This was 246 confirmed by mitochondrial and nuclear DNA data during the late 1980s and 1990s 247 (Miyamoto et al., 1987, 1988; Williams and Goodman, 1989; Goodman et al., 1990, 1994; 248 Ruvolo 1994, 1997; Ruvolo et al. 1994; Arnason et al. 1996; Goodman, 1996). Current 249 estimates based on molecular data indicate that humans and chimpanzees diverged 250 sometime during the Late Miocene (~9–7 Ma; Perelman et al., 2011; Springer et al., 2012; 251 Moorjani et al., 2016). 252 The paradigm shift of cladistics Besides the paleoanthropologist’s reluctance to accept the 253 conclusions of ‘outsiders’ from another discipline in front of the ‘hard evidence’ provided by 254 fossils (see Lewin, 1987), several reasons explain the former’s adherence to an early 255 divergence of humans until the 1970s. First, the molecular revolution took several decades 256 to complete, such that the closer relationship between humans and chimpanzees did not 257 became firmly established until the 1980s (see above). Second, during the 1960s and early 258 1970s, paleoanthropologists lacked the necessary analytical methods to rigorously infer 259 phylogenetic relationships. In the 1960s and 1970s, most paleoanthropologists were still 260 anchored to the Simpsonian systematic paradigm, which allowed for paraphyletic taxa as 261 long as they were based on structural grades (e.g., Simpson, 1945). For this reason, Simons 262 and Pilbeam (1965) favored the traditional division between pongids and hominids 11 263 (Simpson, 1945, 1963) despite recognizing that humans were more closely related to African 264 apes than orangutans. 265 The conceptual cladistic toolkit was available since the publication of Hennig’s (1966) 266 book in English and molecular biologists pioneered the introduction of cladistic ideas in 267 anthropology (Goodman, 1996), as illustrated by their tendency to redefine the content of 268 the Hominidae (e.g., Goodman, 1963). However, it was not until the early 1970s that 269 cladistics started to gain ground in vertebrate paleontology, largely thanks to the work of 270 paleontologists from the American Museum of Natural History (AMNH) in New York (e.g., 271 Nelson, 1972; see review in Cartmill, 2018). In turn, the somewhat slower diffusion of the 272 cladistic paradigm in paleoanthropology during the late 1970s was promoted by Eric Delson, 273 by then already affiliated to the AMNH, and some of his colleagues there (Delson, 1977; 274 Delson et al., 1977). 275 In retrospect, the paradigm shift from Simpsonian to Hennigian systematics throughout 276 the 1970s and 1980s was relatively rapid, according to Cartmill (2018) because the cladistic 277 revolution was mostly ‘esthetic’ (sensu Kuhn, 1970)—i.e., not dictated by new facts or data 278 but by disagreements as to how phylogeny should be reflected in the classifications. 279 Certainly, unlike the molecular revolution in phylogenetic inference, the cladistic paradigm 280 shift was not driven by wealth of new data. Nevertheless, Cartmill’s (2018) account 281 downplays the profound influence that the cladistic paradigm had in the methods and 282 practice of morphology-based phylogenetic inference. Although many aspects of cladistic 283 classification are debatable (e.g., Mayr, 1974), the spread of cladistic analysis had much 284 deeper implications by prompting an explicit recognition that phylogenetic relationships 285 must be determined on the basis of shared-derived features (synapomorphies) as opposed 286 to shared-primitive characters (symplesiomorphies). 12 287 Cartmill (2018) argued that many previous systematists implicitly accepted that only 288 synapomorphies should be considered for assessing phylogeny. However, this was not the 289 case for most paleoanthropologists during the 1960s and early 1970s—as explicitly admitted 290 by Pilbeam (1986). This is also evident from the pioneering work of Delson and Andrews 291 (1975: 405), which aimed to assess the “phyletic relationships among Old World higher 292 primates in the light of the “cladistic” methodology.” These authors had to explain that “only 293 those linkages based on shared derived (“advanced,” apomorphous) characters reflect true 294 phyletic relationships” (Delson and Andrews, 1975: 406), which denotes that this was not yet 295 clearly perceived by then. This is even more clear-cut from Delson’s (1977) didactic effort to 296 explain to the paleoanthropological community the concepts, methods, merits, and pitfalls 297 of cladistics. In turn, Delson et al. (1977) put theory into practice by applying the cladistic 298 methodology to apes and humans. These contributions by Delson and colleagues played a 299 key role in promoting the application of cladistic principles and methods to 300 paleoanthropology despite being still influenced by the then prevailing paradigm that 301 considered Ramapithecus an early member of the human lineage. For example, Delson 302 (1977) adopted a very wide definition of the Hominidae (including apes and humans) but still 303 classified humans and African apes in different subfamilies (Ponginae and Homininae; see 304 also Szalay and Delson, 1979). In any event, the spread of cladistics promoted a more 305 thorough evaluation of morphological evidence (both craniodental and postcranial) in the 306 following decades. 307 308 2.3. The modern phase: Cladistics at its peak and the Sivapithecus dilemma 309 Sivapithecus and the beginning of the modern phase The mid-1970s and 1980s witnessed 310 the description of much more complete cranial remains of Miocene apes (Tekkaya, 1974; 13 311 Kretzoi, 1975; de Bonis and Melentis, 1978; Andrews and Tekkaya, 1980; Wu et al., 1981, 312 1982, 1983, 1984; Pilbeam, 1982; Ishida et al., 1984; Leakey and Walker, 1985; Leakey and 313 Leakey, 1986a, 1986b; Teaford et al., 1988; Zhang et al., 1988) than in the preceding 314 decades—leading to the appreciation that Miocene apes were far more diverse than 315 previously considered (Pilbeam, 1986). Campbell and Bernor’s (1976) attempt of 316 paleobiogeographic synthesis soon became outdated but was followed by Bernor’s (1983) 317 renewed efforts, which could not yet include the determinant discovery of a partial cranium 318 of Sivapithecus that showed many orangutan-like features (Pilbeam, 1982; see also Andrews, 319 1982). By then, persuaded by similarities between material from Turkey (now in 320 Ankarapithecus Ozansoy, 1957) and Sivapithecus (Andrews and Tekkaya, 1980), Andrews 321 had also independently arrived to the conclusion that the latter was an orangutan relative 322 and that the molecular divergence times were correct (Andrews and Cronin, 1982). 323 The discovery of the Sivapithecus cranium led to many new lines of research (Ward, 324 1997a), such as a detailed anatomical analysis of hominoid subnasal morphology (Ward and 325 Kimbel, 1983; Ward and Pilbeam, 1983; McCollum et al., 1993; McCollum and Ward, 1997). 326 These and other analyses led to the recognition of Sivapithecus as the Miocene ape most 327 clearly related to orangutans (Preuss, 1982; Ward and Kimbel, 1983; Ward and Pilbeam, 328 1983; Pilbeam and Smith, 1984; Pilbeam, 1985; Shea, 1985; Ward and Brown, 1986; Brown 329 and Ward, 1988). By this time, the synonymy between Ramapithecus and Sivapithecus 330 became widely accepted (Andrews, 1982; Andrews and Cronin, 1982; Kay, 1982; Lipson and 331 Pilbeam, 1982) and paleoanthropologists finally embraced the molecular-based late 332 divergence between humans and chimpanzees. As noted by Lewin (1987) and Pilbeam 333 (1997), besides the fossil evidence itself this shift in opinion about Ramapithecus was deeply 14 334 influenced by the molecular data, even if this was not widely acknowledged at the time (but 335 see Greenfield, 1980). 336 The establishment of a late-diverging chimpanzee–human clade thus prompted a critical 337 rethinking of the available morphological evidence, while Sivapithecus served to calibrate 338 the hominoid molecular clock. The confluence of these factors with the spread of cladistics 339 gave rise to a new consensus in hominoid phylogenetics during the 1980s (Andrews and 340 Cronin, 1982; Pilbeam, 1984) and a brand-new phase of interpretation of Miocene ape 341 evolution. Thus, since the mid-1980s, most researchers (e.g., Andrews, 1985, 1992; Alba, 342 2012; Fleagle, 2013; Groves, 2017; Almécija et al., 2021), only with few exceptions (e.g., 343 White, 2002; Tuttle, 2014), have distinguished only two extant hominoid families: 344 Hylobatidae and Hominidae, with the latter subdivided into Ponginae and Homininae. The 345 former distinction (e.g., Simpson, 1945) between Pongidae (for apes or great apes) and 346 Hominidae (for humans) became untenable because, from a cladistic viewpoint, paraphyletic 347 groups are unnatural. 348 Nevertheless, as researchers struggled to make compatible the molecular data with the 349 morphological evidence of extant hominoids, this consensus temporarily led to a plethora of 350 widely divergent morphology-based cladistic hypotheses supporting the monophyly of either 351 all great apes (Kluge, 1983), African apes (Andrews, 1987; Andrews and Martin, 1987a), 352 humans and chimpanzees (Groves, 1986), or even humans and orangutans (Schwartz, 1984a, 353 1984b). This is attributable to several factors: (1) some paleoanthropologists had yet to 354 embrace the view that molecular phylogenies were more reliable than morphology-based 355 ones; (2) the implementation of maximum-parsimony computer algorithms for cladistic 356 analysis was still underway; (3) there is an inherent arbitrariness in the selection of 357 characters and the definition of character states (Pilbeam and Young, 2001; Cartmill, 2018); 15 358 and (4) there is a lot of ‘phylogenetic noise’ caused by homoplasy (false homology; e.g., 359 Larson, 1998). While the two first problems were solved during the next decade, the latter 360 two issues have proven more enduring and difficult to resolve. 361 The Sivapithecus dilemma Despite difficulties in retrieving the molecular phylogeny of 362 hominoids based on morphological data, the resolution of the Ramapithecus debate and the 363 discovery of the Sivapithecus cranium soon led to renewed optimism. During the 1980s, it 364 seemed as if a new synthesis of Miocene ape evolution based on the “judicious use of 365 paleontological and neontological data sets” (Pilbeam, 1985: 51) was going to emerge as 366 new fossils were discovered and carefully analyzed. Then, in the blink of an eye, the view 367 that Sivapithecus is a member of the orangutan clade was questioned owing to the discovery 368 of new postcranial remains of this taxon. In particular, two humeri of Sivapithecus indicated 369 that it displayed pronograde locomotor behaviors unlike those of modern hominoids 370 (Pilbeam et al., 1990)—as was further corroborated by additional postcranials during the 371 following decades (Madar et al., 2002; Morgan et al., 2015). More complete postcranial 372 material of other Miocene apes was also discovered throughout the 1980s and 1990s 373 (Walker and Pickford, 1983; Ward et al., 1993; Moyà-Solà and Köhler, 1996), indicating that 374 Early Miocene apes (with some possible exceptions; Gebo et al., 1997) were predominantly 375 pronograde (Ward, 1993; Ward et al., 1993), whereas at least some Late Miocene apes 376 appeared suspensory (Moyà-Solà and Köhler, 1996). 377 In the early 1990s, the consensus emerging from the previous decade was probably too 378 immature to readily comprehend the implications of a Miocene ape combining an 379 orangutan-like cranium with primitive postcranials closely resembling those of the Early 380 Miocene taxa. No matter how Sivapithecus was interpreted from a phylogenetic viewpoint, 381 it was evident there must be a considerable amount of homoplasy involved—implying an 16 382 independent evolution of its orangutan-like cranial morphology, an evolutionary reversal of 383 its postcranium, or the independent acquisition of postcranial similarities by crown 384 hominoids. This conundrum was dubbed the ‘Sivapithecus dilemma’ (Pilbeam and Young, 385 2001; Young, 2003), and has ever since permeated debates about Miocene ape phylogeny. 386 Andrews (1992) reviewed the Miocene apes known by then from a cladistic perspective 387 and favored the view that Sivapithecus was closely related to Pongo Lacépède, 1799, which 388 is the interpretation favored by most subsequent authors until the present (e.g., Ward, 389 1997a, 2015; Larson, 1998; Kelley, 2002; Alba, 2012; Begun, 2015; Pugh, 2022). However, 390 during the 1990s an alternative interpretation was championed by Pilbeam (1996, 1997), 391 who reacted to the Sivapithecus dilemma by closely adhering to the postcranial evidence— 392 incidentally qualifying the profession as “craniophilic” (Pilbeam, 1996: 162). On this basis, he 393 concluded that most Miocene apes—including Middle and Late Miocene ones, such as 394 Sivapithecus, and with only the exception of Oreopithecus Gervais, 1872 and maybe some 395 dryopithecines—were most likely ‘archaic’ hominoids that diverged before the radiation of 396 hominoids of ‘modern aspect’. 397 Pilbeam’s (1996) interpretation of the postcranial similarities between hylobatids and 398 hominids as synapomorphic was a logical consequence of accepting the cladistic paradigm 399 established during the 1980s—contrasting with the prevailing view during the 1950s–1970s, 400 according to which such features would have been developed independently a number of 401 times in different lineages (see discussion in Harrison and Rook, 1997). Similar views were 402 expressed by the latter authors, who referred to previous papers by Harrison (1986, 1987a, 403 1987b, 1991) when concluding that “the postcranial features and character complexes 404 shared by extant hominoids are so detailed and so pervasive that they are extremely unlikely 405 to be the product of convergent evolution” (Harrison and Rook, 1997: 331). This quotation 17 406 illustrates that that adoption of the cladistic paradigm in paleoanthropology during the 407 1980s implied an increased reliance on postcranial features, which some researchers were 408 reluctant to abandon in favor of the old-fashioned greater reliance on craniodental remains. 409 The Sivapithecus dilemma was eventually solved in favor of postcranial homoplasy 410 following a highly influential paper by Larson (1998), who advocated the plausibility of many 411 purported postcranial synapomorphies of hominoids having evolved in parallel multiple 412 times along the various extant lineages. This claim was still a matter of intense debate during 413 the 2000s, including various attempts to measure whether the craniodental or the 414 postcranial data displayed a greater degree of homoplasy (Finarelli and Clyde, 2004; Young, 415 2005). In any event, the discoveries made during the last two decades (e.g., Moyà-Solà et al., 416 2004) have failed to support Pilbeam’s (1996, 1997) prediction that the discovery of 417 additional skeletons would strengthen the view that most Miocene apes are unrelated to 418 the modern radiation. Rather the contrary, Larson’s (1998) views have been vindicated 419 further, supporting the contention that many postcranial similarities among extant ape 420 lineages are indeed homoplastic and that reconstructing last common ancestors based on 421 extant apes alone is totally unreliable (Alba, 2012; Ward, 2015; Almécija et al., 2021). 422 Cladistics in the Computer Age Despite having the problem of postcranial homoplasy in 423 mind, since the 1990s paleoanthropologists have increasingly performed morphology-based 424 cladistic analyses based on both craniodental and postcranial characters to decipher the 425 phylogenetic relationships among the ever-increasing list of Miocene ape taxa. These 426 analyses have been prompted by the important discoveries and reanalyses of Miocene ape 427 crania and postcrania that have taken place during the last three decades (e.g., de Bonis et 428 al., 1990; de Bonis and Koufos, 1993; Moyà-Solà and Köhler, 1993, 1996; Gebo et al., 1997; 429 Nakatsukasa et al., 1998; Ward et al., 1999; Kordos and Begun, 2001; Moyà-Solà et al., 2004, 18 430 2009a, 2009b; Kunimatsu et al., 2007; Suwa et al., 2007; Böhme et al., 2019). The more or 431 less informal cladistic attempts of the early 1990s (Begun, 1992a; Moyà-Solà and Köhler, 432 1995) were soon replaced by formal analyses based on larger taxon-character matrices and 433 performed with the aid of computer algorithms (Begun, 1994, 1995; Shoshani et al., 1996; 434 Begun et al., 1997, 2012; Cameron, 1997a; Finarelli and Clyde, 2004; Rossie and MacLatchy, 435 2005; Zalmout et al., 2010; Stevens et al., 2013; Alba et al., 2015; Nengo et al., 2017; Rossie 436 and Hill, 2018; Gilbert et al., 2020a; Pugh, 2022; Ji et al., 2022). 437 The cladistic analyses performed during the 1990s started to find support for the 438 chimpanzee–human clade on morphological grounds (e.g., Begun, 1992a; Shoshani et al., 439 1996; Begun et al., 1997) and the debate about human origins switched toward the 440 reconstruction of the chimpanzee–human last common ancestor (for recent reviews, see 441 Andrews, 2020 and Almécija et al., 2021). However, it soon became obvious that 442 uncertainties about Miocene ape phylogeny hindered the resolution of this question, as 443 determining the ancestral hominin morphotype cannot be properly done without the aid of 444 the Miocene ape fossil record (e.g., Andrews and Harrison, 2005; Andrews, 2020; Almécija et 445 al., 2021). 446 The changing views on Miocene ape phylogenetic relationships have also given rise to 447 new paleobiogeographic scenarios. In the late 1990s, paleobiogeographic discussion was 448 focused on vicariance, either by assessing alternative phylogenetic hypotheses or by 449 explicitly favoring one of these hypotheses (Begun, 1994, 1995; Agustí et al., 1996; Andrews 450 and Bernor, 1999; Begun et al., 1997). In the 2000s, the recognition that Kenyapithecus was 451 recorded both in Africa and Turkey during the Middle Miocene (Kelley et al., 2008) deserves 452 particular mention, as it has interesting paleobiogeographic implications for hominoid 453 dispersal events (Andrews and Kelley, 2007; Moyà-Solà et al., 2009b; Casanovas-Vilar et al., 19 454 2011; Alba, 2012). In any case, given the diverging opinions on Miocene ape phylogeny, 455 several competing paleobiogeographic scenarios are still subject to ongoing debates 456 (Almécija et al., 2021). 457 458 3. Systematics of Miocene apes 459 Systematics classifies organisms based on evolutionary relationships but has also a 460 utilitarian function so that there is no single true classification to be discovered (Benton, 461 2000)—see Supplementary Online Material (SOM) S1.1 for further details. The classification 462 of Miocene apes presented in this article (Tables 1 and 2) relies on the use of Linnean ranks 463 and other the provisions of the International Code of Zoological Nomenclature; ICZN, 1999). 464 Unless we abandon Linnean nomenclature (see SOM S1.2 for further discussion), paraphyly 465 is ultimately inescapable at the species and genus ranks (Sarmiento et al., 2002), but our 466 systematic scheme assumes that the distinguished family-group taxa are monophyletic (i.e., 467 clades). If the paraphyletic status suspected for some of these taxa was more conclusively 468 supported in the future, our classification should be changed accordingly. Even within the 469 framework of phylogenetic systematics (which aims to faithfully reflect phylogeny and avoid 470 paraphyletic taxa), constructing a classification is a subjective exercise that depends on the 471 taxonomist’s decisions about what phylogenetic hypotheses are best supported, as well as 472 which clades must be denoted as taxa and what ranks should be attributed to each. 473 Therefore, we do not expect our systematic proposal to be uncritically adopted by other 474 scholars. We rather conceive it as a utilitarian construct to transmit our interpretation of 475 current knowledge about Miocene ape evolution. 476 Given that the true phylogeny of any group is scientifically unknowable, phylogenetic 477 inference plays a central role in systematics. In the case of Miocene apes, most studies have 20 478 used cladistic analysis of morphological data based on maximum parsimony to test 479 competing phylogenetic hypotheses (for the epistemological basis of cladistics, see SOM 480 S1.3). Only a few studies have relied on other methods, including Finarelli and Clyde’s (2004) 481 analysis based on stratocladistics—an alternative method of phylogenetic inference that 482 combines morphological and chronostratigraphic data (Fisher, 2008; see SOM S1.4)—and 483 the most recent exploration of Bayesian analyses (see SOM S1.5) by Pugh (2022). We mostly 484 relied on the results of recent cladistic analyses (Nengo et al., 2017: Fig. 5; Gilbert et al., 485 2020a: Fig. 4; Pugh, 2022: Fig. 5), coupled with the synthetic cladogram hypothesized by 486 Gilbert et al. (2020b: Fig. 17.1) and the cladistic results by other authors (Begun et al., 1997: 487 Fig. 1, 2012: Fig. 9; Alba et al., 2015: Fig. 8; Rossie and Hill, 2018: Fig. 5; Ji et al., 2022: Fig. 488 11). The first three recent cladistic analyses mentioned above were performed by the same 489 authors, and hence do not represent independent attempts at Miocene ape phylogenetic 490 reconstruction. However, Pugh’s (2022) analyses were more focused on hominids and hence 491 are more comprehensive regarding the phylogenetic relationships inferred for this group. 492 The phylogenetic and nomenclatural rationale underpinning our classification of Miocene 493 apes, together with taxonomic remarks for particular taxa, are provided below (see SOM S2 494 for nomenclatural remarks). 495 496 3.1. A stem-based definition of the Hominoidea 497 Since Simpson’s (1945) seminal paper, most authors have classified apes and humans 498 within a single superfamily Hominoidea (e.g., Szalay and Delson, 1979; Groves, 1986), with 499 only a few exceptions (e.g., Delson and Andrews, 1975; Thenius, 1981). Simpson (1945) 500 distinguished two families (Pongidae for apes and Hominidae for humans) but other authors 501 restricted pongids to the great apes and distinguished the Hylobatidae for lesser apes (e.g., 21 502 Fiedler, 1956). The latter scheme was widely used until the molecular revolution led to an 503 expanded concept of the Hominidae including the African apes (Goodman, 1974; Andrews 504 and Cronin, 1982), all the great apes (Goodman, 1974; Schwartz et al., 1978; Andrews, 1985; 505 Groves, 1986), or even both lesser and great apes (Delson and Andrews, 1975; Delson 1977; 506 Szalay and Delson, 1979; Goodman et al., 1990, 1998; Goodman, 1996). Groves’ (1986) 507 classification of extant hominoids in two families (Hylobatidae and Hominidae) and hominids 508 in two subfamilies (Ponginae and Homininae) has been very influential, being subsequently 509 adopted by most paleoanthropologists (Andrews, 1992; Andrews et al., 1996; Shoshani et al., 510 1996; Delson, 2000; Wood and Richmond, 2000; Begun, 2002a, 2010; Kelley, 2002; Ward and 511 Duren, 2002; Wood and Harrison, 2011; Alba, 2012; Fleagle, 2013; Almécija et al., 2021). The 512 traditional restricted usage of the Hominidae has been favored recently by several researchers 513 (e.g., White, 2002; White et al., 2009; Tuttle, 2014; Schwartz, 2015). However, such an 514 arrangement can only prevent paraphyly by distinguishing gorillas and chimpanzees at the 515 family rank (i.e., Gorillidae Frechkop, 1943 and Panidae Delson, 1977, respectively; e.g., 516 Schwartz, 1986), which is not favored here. 517 In accordance with many previous contributions (e.g., Andrews, 1992; Alba, 2012; Nengo 518 et al., 2017; Gilbert et al., 2020a; Almécija et al., 2021), here we adopt a stem-based 519 definition of the Hominoidea (Fig. 1) and other included family-group taxa (families, 520 subfamilies, and tribes; see SOM S1.6 for further details). This implies that taxa are defined 521 as putatively monophyletic groups on the basis of the subtaxa included within them, with 522 extant taxa defining the crown group and the extinct members equally related to all of them 523 constituting the stem lineage—for the concepts of ‘total group,’ ‘crown group,’ and ‘stem 524 lineage’, see Figure 1 and SOM S1.7. Thus, crown hominoids are defined as the clade 525 composed by hylobatids and hominids, whereas stem hominoids constitute a paraphyletic 22 526 assemblage of extinct taxa more closely related to both hylobatids and hominids than to the 527 extant sister taxon of hominoids (i.e., cercopithecoids). In turn, crown hominids include 528 pongines and hominines, whereas stem hominids include crown hominoids more closely 529 related to both pongines and hominines than to hylobatids. 530 The theoretical distinction between hominoids and stem catarrhines is straightforward 531 but determining the systematic status of many extinct catarrhines is complicated by 532 uncertainties regarding their branching order relative to cercopithecoids (see Section 1.2). 533 The same applies to the distinction between stem and crown hominoids. Begun et al. (1997) 534 and Begun (2001) informally employed the terms ‘eohominoids’ and ‘euhominoids’ to 535 distinguish what Pilbeam (1996, 1997) termed hominoids of ‘archaic aspect’ and ‘modern 536 aspect,’ respectively. This distinction was subsequently formalized by Begun (2009, 2015) by 537 distinguishing the superfamilies Proconsuloidea and Hominoidea within a magnafamily 538 Hominidea. Indeed, these terms are equivalent to stem and crown hominoids, respectively, 539 and hence unnecessary (Alba, 2012). A distinction of a superfamily Proconsuloidea would 540 only make sense if the included taxa are considered stem catarrhines preceding the 541 cercopithecoid-hominoid split (Harrison, 2002, 2010a). However, the latter view is at odds 542 with the results of formal cladistic analyses (e.g., Begun et al., 1997, 2012; Zalmout et al., 543 2010; Stevens et al., 2013; Alba et al., 2015; Nengo et al., 2017; Rossie and Hill, 2018; Gilbert 544 et al., 2020a) and not followed here. 545 Several family-group taxa (of questionable monophyly) have been recently distinguished 546 within the Hominoidea at the family and/or subfamily rank (Alba, 2012; Fleagle, 2013; Nengo 547 et al., 2017; Gilbert et al., 2020b). Here we follow Gilbert et al. (2020b) in distinguishing 548 three families of putative stem hominoids (Proconsulidae Leakey, 1963, Afropithecidae 549 Andrews, 1992, and Nyanzapithecidae Harrison, 2002), which correspond to the three 23 550 subfamilies formerly distinguished by Harrison (2002, 2010a) within the Proconsuloidea. 551 Some cladistic results (Nengo et al., 2017, Gilbert et al., 2020a) support a basalmost 552 divergence of proconsulids within the hominoid stem lineage, followed by a clade including 553 the paraphyletic Afropithecidae and the Nyanzapithecidae, sister to crown hominoids (Fig. 554 2a). On this basis, Gilbert et al. (2020b) depicted afropithecids and nyanzapithecids in a 555 trichotomy with crown hominoids. In contrast, Rossie and Hill (2018) recovered a basalmost 556 divergence of nyanzapithecids (together with dendropithecids), followed by the paraphyletic 557 proconsulids and finally afropithecids as the sister-taxon of crown hominoids (Fig. 2b). Most 558 recently, Pugh (2022) supported the more basal status of equatorine afropithecids 559 compared with hylobatids. However, the fact that proconsulids were employed as an 560 outgroup (i.e., assumed a priori to be more basal than afropithecids) and that neither 561 afropithecines nor nyanzapithecids were included in Pugh’s (2022) analyses makes it 562 uncertain the early branching topology and putative monophyly of stem hominoid families. 563 Both proconsulids and nyanzapithecids are represented among the earliest known 564 hominoids from the Oligocene of Africa. Proconsulids are recorded by Kamoyapithecus 565 hamiltoni Leakey et al., 1995 from Kenya, ~28–24 Ma, formerly interpreted as a stem 566 catarrhine (Harrison, 2010a, 2013; Zalmout et al., 2010; Stevens et al., 2013) but already 567 showing the distinctive canine morphology of Proconsul (Hammond et al., 2019). 568 Nyanzapithecids are first recorded by Rukwapithecus fleaglei Stevens et al., 2013 from 569 Tanzania at 25.2 Ma and an indeterminate species that co-occurs with Ka. hamiltoni 570 (Hammond et al., 2019). Although these Oligocene hominoids are only recorded from scarce 571 dentognathic material, recent cladistic analyses supported the nyanzapithecid affinities of 572 Rukwapithecus (Nengo et al., 2017; Gilbert et al., 2020a). Together with the fact that 573 afropithecids are not recorded until several million years later, from the Early Miocene, 24 574 these results support an early diversification of stem hominoids well within the Oligocene— 575 in rough agreement with average estimates of the cercopithecoid–hominoid divergence 576 between 32 Ma (Perelman et al., 2011) and 25 Ma (Springer et al., 2012)—as well as a less 577 basal status for afropithecids. Thus, although Nengo et al. (2017) and Gilbert et al. (2020a) 578 recovered afropithecids as paraphyletic, we prefer to keep them as a distinct family until 579 their phylogenetic relationships are clarified further. 580 581 3.2. Proconsulidae 582 Proconsulids include multiple species from the Early Miocene of Kenya and Uganda (~21– 583 16 Ma; Tables 2 and 3). The members of this family show a remarkable body mass disparity 584 (from ~5 to 50 kg; Ruff et al., 1989; Rafferty et al., 1995; Harrison, 2010a) and retain multiple 585 plesiomorphic features compared with crown hominoids (Harrison, 2010a; Begun, 2015), such 586 as well-developed molar cingula, an open palatine fenestra, a short tubular ectotympanic with 587 a deep V-shaped notch in the external margin, and a deep subarcuate fossa. Postcranially, 588 they display a mosaic of primitive (mostly platyrrhine-like) and derived (hominoid-like) 589 postcranial features, overall indicative of a pronograde body plan suggestive of generalized 590 arboreal quadrupedalism and powerful-grasping cautious climbing (Rose, 1983, 1997; Ward, 591 1993, 1997b, 2015; Ward et al., 1993; Kelley, 1997; Walker, 1997; Harrison, 2010a; Daver and 592 Nakatsukasa, 2015). The hominoid status of proconsulids is highlighted by the lack of an 593 external tail (Ward et al., 1991; Kelley, 1997; Nakatsukasa et al., 2003), although they display 594 other more subtle features derived toward the hominoid condition, such as an incipient distal 595 radioulnar diarthrosis (Daver and Nakatsukasa, 2015). Proconsulids display a lower degree of 596 encephalization than great apes (being more comparable in this regard to extant hylobatids; 597 Walker et al., 1983; Alba, 2010) but perhaps—based on crown formation time—already 25 598 possessed a somewhat slower life history than cercopithecoids (Kelley, 1997, 2004; Kelley and 599 Smith, 2003). 600 We recognize the genus Ekembo McNulty et al., 2015 for two species—Ekembo nyanzae 601 (Le Gros Clark and Leakey, 1950) and Ekembo heseloni (Walker et al., 1993)—formerly 602 included in Proconsul (e.g., Harrison, 2010a). According to McNulty et al. (2015), Ekembo 603 displays some derived features relative to Proconsul, but thus far cladistic analyses have not 604 recovered the two genera as distinct subclades (Nengo et al., 2017; Rossie and Hill, 2018; 605 Gilbert et al., 2020a). However, it is noteworthy that Pugh (2022) refrained from analyzing 606 the two Ekembo species separately because of the difficulties to delimit their respective 607 hypodigms. We also follow most previous authors (MacLatchy and Rossie, 2005; Harrison 608 and Andrews, 2009; Harrison, 2010a, McNulty et al., 2015) in considering that 609 Ugandapithecus Senut et al., 2000, originally erected for Proconsul major Le Gros Clark and 610 Leakey, 1950 and subsequently expanded by some authors (Pickford and Kunimatsu, 2005; 611 Pickford et al., 2009a) to include Proconsul meswae Harrison and Andrews, 2009, Proconsul 612 gitongai (Pickford and Kunimatsu, 2005), and Proconsul legetetensis (Pickford et al., 2009a), 613 is a junior subjective synonym of Proconsul. However, Ugandapithecus remains potentially 614 available for P. major if other species are eventually shown to be more closely related to it 615 than to Proconsul africanus Hopwood, 1933. Moreover, following Pickford et al. (2020, 2021) 616 we find the synonymy favored by McNulty et al. (2015) between P. legetetensis (originally 617 described in Ugandapithecus) and P. major unconvincing (as they remarked the 618 distinctiveness of the holotype mandible) and tentatively prefer to keep the species distinct. 619 We also include in this family the genus Kalepithecus Harrison, 1988—whose type species, 620 Kalepithecus songhorensis (Andrews, 1979), was left as incertae sedis by Harrison (2010a)— 621 because most recent analyses (Nengo et al., 2017; Gilbert et al., 2020a) have supported the 26 622 proconsulid affinities previously noted for this species (Rae, 1997). Kalepithecus differs from 623 small-bodied stem catarrhines in the inferiorly broader nasal aperture and relatively deep 624 clivus (Harrison, 1988, 2002, 2010a; Rae, 1997), supporting its hominoid status. Proconsulids 625 may be thus more diverse than currently recognized, as it seems plausible that other small- 626 bodied catarrhines from Africa, mostly known from dentognathic material, might ultimately 627 be shown to belong to this family when more complete cranial material becomes available. 628 629 3.3. Nyanzapithecidae and Oreopithecini 630 Nyanzapithecidae The Miocene species and genera included here in the Nyanzapithecidae 631 follow Harrison’s (2010a) concept of Nyanzapithecinae with the addition of Nyanzapithecus 632 alesi Nengo et al., 2017 and Samburupithecus kiptalami Ishida and Pickford, 1997. The latter 633 species, known from the Late Miocene (~8.5 Ma) of Kenya on the basis of a maxillary fragment, 634 only slightly postdates the otherwise last occurrence of the group at ~10 Ma (Kunimatsu et 635 al., 2007) but is much larger than other nyanzapithecid described genera (~10–15 kg for males; 636 Harrison, 2010a; Fleagle, 2013). Samburupithecus Ishida and Pickford, 1997 was originally 637 considered a stem hominine (Ishida et al., 1984; Andrews, 1992; Ishida and Pickford, 1997; 638 Pickford and Ishida, 1998) but subsequently interpreted by as a late surviving stem hominoid 639 by other authors (Begun, 2001, 2013, 2015; Olejniczak et al., 2009; Almécija et al., 2012; Begun 640 et al., 2012)—see discussion in Harrison (2010a). Pugh’s (2022) analyses linked 641 Samburupithecus with Oreopithecus based on dental similarities shared with nyanzapithecids. 642 The nyanzapithecid-like dental features of Samburupithecus had been already noted by 643 Harrison (2010a), who nevertheless left the genus as incertae sedis. Based on the meager 644 evidence available, we consider that an inclusion of Samburupithecus within the 645 Nyanzapithecidae is warranted. Other large nyanzapithecids are recorded in Africa but remain 27 646 indeterminate due to the scarcity of material. They include Early/Late Miocene (~16 Ma) 647 remains from South Africa that represent the southernmost record of Miocene apes (Senut et 648 al., 1997; Harrison, 2010a). This species might be related to the younger and somewhat 649 smaller nyanzapithecid from the Middle Miocene of Kenya (Fort Ternan and Kapsibor, 13.7 650 Ma; Leakey, 1968; Harrison, 1986, 1992), which likely represents a new genus and species 651 (Harrison, 2010a). 652 Nyanzapithecids possess a distinctive dental morphology (Harrison, 2013; Nengo et al., 653 2017; Rossie and Cote, 2002) and, based on Nyanzapithecus Harrison, 1986 and 654 Turkanapithecus Leakey and Leakey, 1986b, also cranial similarities with hylobatids, which are 655 generally interpreted as homoplastic (Nengo et al., 2017). Nyanzapithecus displays a fully 656 ossified external acoustic meatus (Nengo et al., 2017), which appears more extant-catarrhine- 657 like than in Ekembo and stem cercopithecoids—despite being fully ossified, in the latter taxa 658 the ectotympanic is shorter and lacks a completely closed ventral tip (Alba et al., 2015), 659 implying some degree of independent evolution of ectotympanic ossification, as previously 660 suggested (Begun, 2002b). The postcranial morphology of nyanzapithecids suggests that they 661 were above-branch, pronograde quadrupeds broadly similar to proconsulids, but perhaps 662 with enhanced climbing abilities (Harrison, 2010a). 663 Recent cladistic analyses have supported nyanzapithecids as more basal than proconsulids 664 (Rossie and Hill, 2018) or as less basal than both proconsulids and afropithecids (Nengo et al., 665 2017; Gilbert et al., 2020a)—see Figure 2. We follow Harrison (2010a) in considering that 666 Xenopithecus Hopwood, 1933 is distinct from Proconsul (see also Pickford and Kunimatsu, 667 2005) and shows nyanzapithecid affinities based on dental morphology. Mabokopithecus von 668 Koenigswald, 1969 shows more unambiguous nyanzapithecid affinities, and while some 669 authors have favored a generic distinction from Nyanzapithecus (Benefit et al., 1998), Harrison 28 670 (2002, 2010a) tentatively supported their synonymy—although he did not formalize it, 671 pending the description of unpublished material. As remarked by Harrison (2002, 2010a), 672 Mabokopithecus has priority over Nyanzapithecus, so the species currently included in the 673 latter genus might have to be eventually transferred to the former. In any event, this would 674 not affect the validity of family-group taxa based on Nyanzapithecus. Pending further 675 clarification of the internal phylogenetic relationships of the Nyanzapithecidae and their 676 possible link with Oreopithecus (see below), we refrain from distinguishing nyanzapithecid 677 subfamilies. 678 Oreopithecini Oreopithecus, from the Late Miocene of Europe (~8–7 Ma; Rook et al., 2011), 679 is the most completely preserved Miocene ape, being known from dental, cranial, and 680 postcranial remains. However, due to a unique combination of features, the phylogenetic 681 relationships of Oreopithecus have been controversial for a century and a half (see review in 682 Delson, 1986). Oreopithecus has been recognized as a hominoid mostly based on its derived 683 postcranium (Harrison, 1987a, 1991; Sarmiento, 1987; Harrison and Rook, 1997), being 684 considered a close relative of nyanzapithecids from Africa (Harrison, 1986, 1987a; Benefit et 685 al., 1998; Benefit and McCrossin, 2001; Rossie and Cote, 2022) or a descendent of 686 dryopithecines from Europe (Harrison and Rook, 1997; Moyà-Solà and Köhler, 1997). 687 Former cladistic analyses recovered a basalmost stem hominid position for Oreopithecus 688 (Begun et al., 1997, 2012), but more recent ones supported its nyanzapithecid affinities 689 (Nengo et al., 2007; Gilbert et al., 2020a) and hinted at a possible relationship with 690 Samburupithecus (Pugh, 2022). The combined results of these cladistic analyses, largely 691 driven by dental similarities, could justify the inclusion of Oreopithecus in the 692 Nyanzapithecidae—which, as noted by Gilbert et al. (2020b), would imply that the correct 693 name for the family is Oreopithecidae Schwalbe, 1915, as already used for these taxa in 29 694 previous decades (e.g., Harrison, 1986, 1987a; Benefit et al., 1998; Benefit and McCrossin, 695 2001). Here we take a conservative approach and refrain from formalizing the inclusion of 696 Oreopithecus in the Nyanzapithecidae, classifying it instead in a tribe of its own, which is left 697 as incertae sedis within the Hominoidea (Gilbert et al., 2020b). 698 Oreopithecus is larger-bodied (>30 kg in males; Jungers, 1987) than most nyanzapithecids 699 except Nyanzapithecidae nov. from Fort Ternan and Kapsibor as well as Samburupithecus. 700 The postcranial material of Oreopithecus further differs from the scarce postcranials 701 available for nyanzapithecids by possessing multiple adaptations for antipronograde 702 positional behaviors (Harrison, 1987a, 1991; Jungers, 1987; Sarmiento, 1987), which have 703 been subject to different interpretations. Some features have been interpreted as indicative 704 of terrestrial bipedalism (Straus, 1963; Köhler and Moyà-Solà, 1997; Rook et al., 1999; Moyà- 705 Solà et al., 2005a), in agreement with the possession of human-like hand proportions 706 suitable for refined manipulation (Moyà-Solà et al., 1999a, 2005a; Almécija et al., 2014). In 707 contrast, other authors have emphasized the possession of adaptations for vertical climbing 708 (Sarmiento, 1987; Sarmiento and Marcus, 2000; Hammond et al., 2020) and suspensory 709 behaviors (Jungers, 1987; Harrison, 1991; Harrison and Rook, 1997; Susman, 2004; Begun, 710 2007; Deane and Begun, 2008; Russo and Shapiro, 2013). Part of the debate around the 711 locomotion of Oreopithecus stems from focusing on artificial locomotor categories instead of 712 positional repertoires with different behaviors displayed at varying frequencies. The lower 713 torso of Oreopithecus lacks features related to stabilization during bipedalism as well as the 714 stiffness characteristic of extant great apes, being rather reminiscent of hylobatids 715 (Hammond et al., 2020). Coupled with its manual proportions and large body mass, this 716 evidence suggests that Oreopithecus might have been an orthograde arboreal ape that 30 717 specialized in slow climbing but was committed neither to bipedalism or suspension, even if 718 these behaviors might have been part of its positional repertoire. 719 720 3.4. Afropithecidae 721 As noted above, the monophyly of afropithecids as conceived here (Tables 1 and 2) 722 remains to be better ascertained, even if most analyses concur that they are less basal than 723 proconsulids (Nengo et al., 2017; Rossie and Hill, 2018; Gilbert et al., 2020a; Pugh, 2022), in 724 agreement with their younger chronostratigraphic range. Afropithecids include medium- to 725 large-bodied species recorded from the Early to Middle Miocene (from~21–20 to ~12 Ma) of 726 Kenya, Uganda, Namibia, and Saudi Arabia (Table 3). Two afropithecid subfamilies are 727 recognized here (Afropithecinae Andrews, 1992 and Equatorinae Cameron, 2004), which are 728 separately discussed below. The phylogenetic relationships of some Miocene apes generally 729 considered to be more or less closely related to Afropithecus Leakey and Leakey, 1986a— 730 namely Heliopithecus Andrews and Martin, 1987b and Otavipithecus Conroy et al., 1992 731 (Andrews, 1992; Andrews and Kelley, 2007; Harrison, 2010a; Alba, 2012; Begun, 2013, 732 2015)—is particularly uncertain because they have not been included in recent cladistic 733 analyses. Heliopithecus, from the Early Miocene (16 Ma) of Saudi Arabia, has been 734 tentatively included in the Afropithecinae based on dental similarities (Andrews and Martin, 735 1987b; Harrison, 2010a). In turn, the medium-sized (14–20 kg) Otavipithecus, from the 736 Middle Miocene (12 Ma) of Namibia, has been included in the Afropithecidae given the 737 affinities with Afropithecus suggested by several authors (e.g., Andrews, 1992; Harrison, 738 2010a) and further supported by a cladistic analysis of mandibular characters (Singleton, 739 2000). However, Otavipithecus has been left as subfamily incertae sedis given the lack of 740 cladistic studies evaluating its relationships with other afropithecids. Otavipithecus is 31 741 recorded by craniodental and some postcranial remains (the latter being indicative of 742 arboreal locomotion) and represents one of the southernmost occurrences of Miocene apes 743 (Conroy et al., 1992; Mocke et al., 2022). 744 Afropithecinae Afropithecids are first recorded by afropithecines, which display a primitive 745 nasoalveolar morphology (Begun and Gülec, 1998; Brown et al., 2005; Nakatsukasa and 746 Kunimatsu, 2009; Begun, 2015) and differ from proconsulids in craniodental features 747 probably related to sclerocarpic feeding (Leakey and Walker, 1997; Begun, 2015; Deane, 748 2017). The postcranial morphology of afropithecines is less thoroughly known than that of 749 proconsulids, but similarly indicative of a pronograde body plan (Ward, 2015), except for 750 some postcranial remains attributed to Morotopithecus bishopi Gebo et al., 1997, which are 751 suggestive of orthograde behaviors (Sanders and Bodenbender, 1994; Gebo et al., 1997; 752 MacLatchy et al., 2000, 2019; MacLatchy, 2004; Nakatsukasa, 2008). This species has 753 sometimes been recovered as a stem hominid by cladistic analyses (Young and MacLatchy, 754 2004), but the most recent ones recovered it as a stem hominoid more basal than (Nengo et 755 al., 2017; Gilbert et al., 2020a) or as basal as (Rossie and Hill, 2018) Afropithecus. 756 The distinctiveness of the genus Morotopithecus Gebo et al., 1997 has been disputed by 757 several authors, who considered its type and only species (M. bishopi) a junior synonym of 758 Afropithecus turkanensis Leakey and Leakey, 1986a (Pickford, 2002, 2021; Pickford et al., 759 2003, 2017; Patel and Grossman, 2006; Harrison, 2010a; Pickford et al., 2017; Van Couvering 760 and Delson, 2020). Under this view, the cranial differences between the two genera (Gebo et 761 al., 1997; MacLatchy et al., 2000; Begun, 2015; Deane, 2017) might be attributable to 762 pathological remodeling in the holotype of M. bishopi and diagenetic deformation in that of 763 A. turkanensis (Pickford, 2002; Pickford et al., 2017), which other authors also consider to be 764 badly distorted (Begun, 2015). Based on the dental differences between M. bishopi and A. 32 765 turkanensis pointed out by MacLatchy et al. (2019), we tentatively favor the distinction of 766 these taxa at least to the species rank, but remain skeptical about the generic distinction for 767 several reasons. First, the putative differences in facial morphology—including the seemingly 768 more restricted palatine fenestra of Afropithecus (Gebo et al., 1997; Brown et al., 2005; 769 MacLatchy et al., 2019)—are uncertain in the light of the aforementioned arguments about 770 remodeling and distortion. Second, MacLatchy et al.’s (2019) contention that the age 771 difference between the two taxa (>3 Myr, assuming a dating of 21 Ma is correct, see footnote 772 in Table 2) supports their distinction is not particularly relevant from a taxonomic viewpoint 773 and applies to Miocene ape species but not genera. 774 The original diagnosis of Morotopithecus was partly based on its purported derived (crown 775 hominoid-like) postcranial features (Gebo et al., 1997). Nevertheless, it has subsequently been 776 argued that more than a single hominoid is present at Moroto. MacLatchy et al. (2019) 777 recognized M. bishopi and a smaller proconsulid, while Jansma and MacLatchy (2015) further 778 reported a nyanzapithecid. In contrast, Pickford et al. (2017) and Pickford (2021) recognized 779 A. turkanensis, P. gitongai, and Nacholapithecus kerioi Ishida et al., 1999. Pickford (2021) even 780 discussed the possibility that the Moroto vertebrae assigned to M. bishopi (Walker and Rose, 781 1968; Sanders and Bodenbender, 1994; Gebo et al., 1997; Nakatsukasa, 2008) might date to 782 the Plio-Pleistocene. Such a claim would have important implications for the earliest evidence 783 of orthogrady in the hominoid fossil record (Gebo et al., 1997; MacLatchy, 2004; Young and 784 MacLatchy, 2004). However, it is based on very circumstantial evidence—basically, that the 785 fossils were surface-collected from sediments that have also yielded some Plio-Pleistocene 786 fossils and that differences in preservation hint at a different taphonomic history for the 787 vertebrae. Therefore, unless geochemical analyses eventually prove the contrary, a Miocene 788 age seems much more likely. Nevertheless, the impossibility to demonstrate a close spatial 33 789 association between the holotype and the postcranial remains casts some doubts on their 790 attribution to the same taxon (MacLatchy et al., 2019; Pickford, 2021). Pending future analyses 791 that might clarify the number and identity of large hominoid taxa present at Moroto and the 792 composition of the M. bishopi hypodigm, we consider it prudent to keep the genus distinct. 793 Equatorinae Equatorines have been inferred to display, like afropithecines, adaptations to 794 hard-object feeding (McCrossin and Benefit, 1993, 1997; Nakatsukasa and Kunimatsu, 2009). 795 Nacholapithecus Ishida et al., 1999 differs from afropithecines by possessing slight 796 premaxillary-maxillary overlap (Ishida et al., 2004) as well as an obliterated subarcuate fossa 797 (Kunitmatsu et al., 2019). These features, which cannot be ascertained in Equatorius Ward et 798 al., 1999, have given rise to different phylogenetic interpretations for Nacholapithecus (Alba, 799 2012; Kunimatsu et al., 2019; Pugh, 2022). The postcranial morphology of equatorines is 800 better known than that of afropithecines and similar to that of proconsulids, being indicative 801 of a pronograde body plan without external tail (as in Nacholapithecus; Nakatsukasa et al., 802 2003). However, Nacholapithecus possesses some features indicative of increased forelimb- 803 dominated arboreal behaviors (climbing and clambering; Ishida et al., 2004; Nakatsukasa 804 and Kunimatsu, 2009), while Equatorius shows evidence of semiterrestriality (McCrossin and 805 Benefit, 1997; Patel et al., 2009). 806 Following the description of Equatorius, this genus has been considered distinct from 807 both Kenyapithecus and Griphopithecus Abel, 1902 (here included in the Hominidae, see 808 below) by most (e.g., Alba, 2012; Begun, 2015) but not all (Begun, 2000, 2001, 2002a; 809 Benefit and McCrossin, 2000; Güleç and Begun, 2003; Kunimatsu et al., 2004; Mocke et al., 810 2022) researchers. Begun (2002a) even formally proposed to synonymize the genus 811 Equatorius with Griphopithecus, but subsequently abandoned this view (e.g., Begun et al., 812 2012; Begun, 2015). We concur with most authors (e.g., Kelley et al., 2000, 2002; Ward and 34 813 Duren, 2002; Andrews and Kelley, 2007; Kelley et al., 2008; Harrison, 2010a; Alba, 2012; 814 Begun, 2015) that Equatorius is sufficiently distinct from Kenyapithecus, with the latter 815 differing, among other features, by possessing derived hominid synapomorphies such as a 816 higher zygomatic root (Pickford, 1985; Harrison, 1992). 817 The systematic position of Equatorius and Nacholapithecus has been subject to different 818 interpretations. Ward et al. (1999) considered Equatorius to be a stem hominoid less derived 819 than Kenyapithecus and more closely related to Afropithecus, but Ward and Duren (2002) 820 included them all within the Hominidae (albeit in different subfamilies). Alba (2012), in 821 contrast, considered only Afropithecus to be a stem hominoid and included the other genera 822 within the Hominidae, distinguishing two tribes within the Kenyapithecinae Andrews, 1992. 823 Following the recent cladistic results indicating that Equatorius is a stem hominoid, perhaps 824 even more basal than Afropithecus (Nengo et al., 2017; Gilbert et al., 2020a), Gilbert et al. 825 (2020b) left Equatorius and Nacholapithecus as family incertae sedis—even though 826 Nacholapithecus was not included in these analyses. Pugh’s (2022) results recovered 827 Equatorius and Nacholapithecus as a clade of stem hominoids, but her analyses did not 828 include Afropithecus, thereby leaving unresolved the relationships between afropithecines 829 and equatorines. Based on current knowledge, we prefer to group Equatorius and 830 Nacholapithecus in a single subfamily (Equatorinae) within the Afropithecidae, while 831 recognizing that the latter family might ultimately prove to be paraphyletic. 832 833 3.5. Hylobatidae 834 The origin of hylobatids is one of the most enduring problems in hominoid phylogenetics. 835 As noted in Section 1.2, the Middle Miocene small catarrhine Kapi Gilbert et al., 2020a from 836 India was originally interpreted as a stem hylobatid (see also Gilbert et al., 2020b) but has 35 837 more recently been considered a pliopithecoid (Ji et al., 2022). This conclusion partly relies 838 on the interpretation that Kapi possesses an incipiently developed mesial arm of the 839 pliopithecine triangle, coupled with other dental similarities (Ji et al., 2022). The homology of 840 the weakly developed crest displayed by Kapi with the pliopithecine triangle is debatable, 841 but in the lack of additional material we concur that the stem hylobatid status of Kapi is 842 insufficiently supported at present. In contrast, Yuanmoupithecus xiaoyuan Pan, 2006, from 843 the Late Miocene (~8.0–7.0 Ma) of China, is known from multiple teeth and a partial lower 844 face (Pan, 2006; Harrison et al., 2008; Harrison, 2016; Ji et al., 2022) and more 845 unambiguously supported as a stem hylobatid by cladistic analyses (Gilbert et al., 2020a; Ji et 846 al., 2022). For this reason, we classify this genus in a monotypic subfamily within the 847 Hylobatidae, Yuanmoupithecinae subfam. nov. (type genus: Yuanmoupithecus), so as to 848 reserve the Hylobatinae for crown hylobatids. Ji et al. (2022) also recovered the 849 dendropithecid Micropithecus Fleagle and Simons, 1978 as a stem hylobatid but they 850 interpreted this result as an artifact caused by functional dental convergence. This 851 interpretation is also favored here but, given that some cladistic analyses have supported a 852 stem hominoid status for dendropithecids (see Section 1.2) and that the Early Miocene 853 forerunners of hylobatids are unknown, the possibility that Micropithecus is more closely 854 related to hylobatids than to other dendropithecids from Africa (see also Section 4.3) should 855 be investigated further. 856 857 3.6. Stem Hominidae 858 Kenyapithecinae We distinguish a hominid subfamily (Kenyapithecinae) for Kenyapithecus 859 and Griphopithecus, which is equivalent to Alba’s (2012) tribe Kenyapithecini and Gilbert et 860 al.’s (2020b) family Kenyapithecidae. The latter authors considered unresolved the 36 861 systematic position of this taxon (stem vs. crown hominoid) but, based on Pugh’s (2022) 862 results, a stem hominid status seems more likely. Although there has been some confusion 863 about the taxon name for the clade including both Kenyapithecus and Griphopithecus, 864 Kenyapithecinae is to be preferred over Griphopithecinae (Casanovas-Vilar et al., 2011; see 865 SOM S2.1 for further details). 866 Kenyapithecines are medium to large-bodied apes (~27–48 kg; Fleagle, 2013) known from 867 isolated teeth, dentognathic fragments, and a few postcranial remains, the latter consistent 868 with a pronograde body plan adapted to semiterrestrial quadrupedalism (Begun, 1992c, 869 2002a, 2015; Ersoy et al., 2008; Harrison, 2010a; Alba, 2012). The two kenyapithecid genera 870 are distinguished mostly on the basis of dentognathic features (Kelley et al., 2008). 871 Kenyapithecus is here considered the oldest known hominid from Africa (13.7 Ma) and also 872 the only one recorded both in Africa and Eurasia, where it is represented by a slightly older 873 species from Turkey (~14.5–14.0 Ma). Griphopithecus also comprises two species from the 874 Middle Miocene, one from Turkey (~14.5–13.4 Ma) that co-occurs with Kenyapithecus, and 875 another from Central Europe (Germany and Slovakia) that is probably younger (~13.8–12.7 876 Ma). There are two species names available for the latter species, but Griphopithecus suessi 877 Abel, 1902, as the type species of the genus, is to be preferred for the reasons explained in 878 SOM S2.2. Middle Miocene kenyapithecines, given their African and Eurasian geographic 879 distribution, play a key role for understanding the dispersal and subsequent radiation of 880 Miocene apes in Eurasia. Their oldest representatives predate the appearance of both 881 dryopithecines and pongines in Eurasia ~13.0–12.5 Ma (Alba, 2012; Alba et al., 2017, 2022; 882 Gilbert et al., 2020b), and thus represent plausible potential ancestors for these groups in 883 chronological terms. 37 884 In the past, some cladistic analyses indicated a stem hominoid status for Kenyapithecus 885 s.l. (before Equatorius was distinguished; Begun et al., 1997) or Kenyapithecus s.s. (Finarelli 886 and Clyde, 2004), although the alternative stratocladistic analysis by the latter authors 887 supported Kenyapithecus as the last common ancestor of crown hominids (see SOM S1.4 888 regarding stratocladistics). Indeed, there is a broad (even if not universal) consensus that 889 Kenyapithecus is likely a stem hominid (Harrison, 1992, 2010a; Ward et al., 1999; Kelley et 890 al., 2002, 2008; Ward and Duren, 2002; Andrews and Kelley, 2007; Alba, 2012), as further 891 supported by recent cladistic analyses (Begun et al., 2012; Pugh, 2022). Kenyapithecus 892 already displays a clear hominid synapomorphy in the possession of a high zygomatic root 893 (Pickford, 1986; Harrison, 1992, 2010a; Kelley et al., 2008; Alba, 2012; Begun, 2015). Other 894 features that generally distinguish Kenyapithecus from Equatorius (such as the configuration 895 of the maxillary sinus) are found in pongines but not hominines (Harrison, 1992, 2010a), and 896 indeed some recent analyses recovered Kenyapithecus as crown hominid more closely 897 related to pongines (Nengo et al., 2017; Gilbert et al., 2020a). Nevertheless, Pugh’s (2022) 898 cladistic analyses—specifically focused on great apes—supported instead the stem hominid 899 status of Kenyapithecus, in agreement with Harrison’s (1992, 2010a) interpretation that the 900 features shared with pongines are symplesiomorphic. 901 The hominid status of Griphopithecus is less clear-cut, as it has not been frequently 902 included in cladistic analyses. Begun et al. (2012) recovered it as a stem hominoid but Pugh’s 903 (2022) analyses supported its stem hominid status as well as its inclusion within the same 904 clade as Kenyapithecus—albeit only in some cases as sister taxa and in most other instances 905 together with the dryopithecines Pierolapithecus Moyà-Solà et al., 2004 and Anoiapithecus 906 Moyà-Solà et al., 2009b, thereby only tentatively justifying the inclusion of Griphopithecus in 907 the Kenyapithecinae. The oldest record of hominoids in Eurasia consists of an upper molar 38 908 fragment from Engelswies (~16.5–16.0 Ma; Casanovas-Vilar et al., 2011) that has been 909 variously attributed to cf. Griphopithecus sp. (Heizman and Begun, 2001; Begun, 2015), 910 ?Griphopithecus sp. (Andrews et al., 1996), and aff. Griphopithecus (Begun, 2002a). 911 However, Casanovas-Vilar et al. (2011) questioned the possibility to reach an assignment 912 (even if tentative) at the genus rank, and we concur it is best attributed to cf. 913 Kenyapithecinae. 914 Dryopithecinae We follow Alba (2012) and other recent authors (e.g., Andrews, 2020; Gilbert 915 et al., 2020b) in classifying many European Middle to Late Miocene apes in a distinct 916 subfamily of putative stem hominids (Dryopithecinae Gregory and Hellman, 1939), rather 917 than a tribe (Dryopithecini) of stem hominines (Begun, 2009, 2010, 2013, 2015). We also 918 follow Alba (2012) in distinguishing the dryopithecine tribes Dryopithecini and 919 Hispanopithecini Cameron, 1997a, respectively for Middle Miocene (12.4–11.6 Ma) and 920 early Late Miocene (11.6–9.5 Ma) genera. However, we exclude the graecopithecin 921 Ouranopithecus de Bonis and Melentis, 1977 (9.7–7.6 Ma) from this subfamily (see next 922 subsection). Dryopithecines are mostly recorded from western and central Europe 923 (Casanovas-Vilar et al., 2011), but persisted until ~8 Ma in Georgia as recorded by 924 ?Udabnopithecus Burchak-Abramovich and Gabashvili, 1945 (Agustí et al., 2020), of 925 uncertain taxonomic validity, thus overlapping by more than 1 Myr with the 926 chronostratigraphic range of graecopithecins. 927 Dryopithecines are large-bodied apes that display multiple crown-hominid cranial 928 synapomorphies, such as a high zygomatic root, deep palate, and wide nasal aperture 929 (Moyà-Solà and Köhler, 1993, 1995; Begun, 1994; Moyà-Solà et al., 2004; 2009a, 2009b; 930 Alba, 2012; Gunz et al., 2020). Pierolapithecus, Hispanopithecus Villalta Comella and 931 Crusafont Pairó, 1944, and Rudapithecus Kretzoi, 1969 are the best known dryopithecine 39 932 genera, both cranially and postcranially, although some cranial remains are also available for 933 Dryopithecus and Anoiapithecus. Hispanopithecus and Rudapithecus further lack a 934 subarcuate fossa (Moyà-Solà and Köhler, 1993, 1995; Kordos and Begun, 1997, 2001; Begun, 935 2015) but the importance of this feature as a hominid synapomorphy is currently uncertain 936 (Pugh, 2022; see Subsection 4.5). Postcranial remains are known for all dryopithecine genera 937 except Anoiapithecus and ?Udabnopithecus. Dryopithecus is not very well-known 938 postcranially, but available specimens (Pilbeam and Simons, 1971; Moyà-Solà et al., 2009b; 939 Alba et al., 2011; Almécija et al., 2012) are suggestive of powerful grasping above-branch 940 quadrupedalism and cautious climbing without evidence of an orthograde body plan (Pina et 941 al., 2019). In contrast, the roughly coeval Pierolapithecus represents the oldest unambiguous 942 evidence of an orthograde body plan in the fossil hominoid record (Moyà-Solà et al., 2004; 943 Alba, 2012), despite the lack of specific adaptations to suspensory behaviors (Moyà-Solà et 944 al., 2004, 2005b; Almécija et al., 2009; Alba et al., 2010; contra Begun and Ward, 2005; 945 Deane and Begun, 2008, 2010; Begun et al., 2012). Overall, the postcranial evidence for 946 Pierolapithecus indicates a forelimb-dominated locomotor repertoire with a significant 947 component of vertical climbing despite retaining adaptations for above-branch powerful- 948 grasping quadrupedalism (Moyà-Solà et al., 2004, 2005b; Almécija et al., 2009; Alba et al., 949 2010; Alba, 2012; Hammond et al., 2013). Danuvius Böhme et al., 2019 similarly displays an 950 orthograde body plan but already shows suspensory adaptations (Böhme et al., 2019; 951 Williams et al., 2020; Almécija et al., 2021), which are even more clearly expressed in 952 Hispanopithecus (Moyà-Solà and Köhler, 1996; Almécija et al., 2007; Alba et al., 2010, 2012b; 953 Alba, 2012; Susanna et al., 2014; Pina et al., 2012) and Rudapithecus (Morbeck, 1983; Begun, 954 1988, 1992c, 1993, 1995). Nevertheless, at least Hispanopithecus still retains some features 40 955 functionally related to above-branch quadrupedalism (Almécija et al., 2007; Alba et al., 2010, 956 2012b). 957 It is generally considered that dryopithecines are less basal hominids than 958 kenyapithecines (e.g., Moyà-Solà and Köhler, 1993, 1995, 1996; Moyà-Solà et al., 2004, 959 2009a, 2009b; Begun, 2009, 2010, 2015; Alba et al., 2010; Casanovas-Vilar et al., 2011; Alba, 960 2012; Gilbert et al., 2020b), as supported by cladistic analyses (Begun et al., 2012; Pugh, 961 2022). However, dryopithecines have been variously interpreted as pongines (e.g., Moyà- 962 Solà and Köhler, 1995), hominines (e.g., Begun et al., 2012), or stem hominids (e.g., Alba, 963 2012; Alba et al., 2015; Pugh, 2022)—see discussion in Almécija et al. (2021). Such 964 uncertainties led Gilbert et al. (2020b) to place dryopithecines in a trichotomy with crown 965 hominids, in further agreement with some other cladistic analyses (Nengo et al., 2017; 966 Gilbert et al., 2020a). Dryoptihecine genera differ from one another in cranial and 967 postcranial features but are dentally conservative, although hispanopithecins appear 968 somewhat more derived in the latter regard (e.g., the C1 morphology of the of male 969 individuals and the more peripheral dentine horns of the upper molars; Alba, 2012; Fortuny 970 et al., 2021). Pugh’s (2022) analyses did not find clear support that dryopithecins are less 971 basal than kenyapithecines and, hence, the monophyly of dryopithecines as conceived here 972 remains uncertain (Alba, 2012; Almécija et al., 2021; Pugh, 2022). Given the lack of a cladistic 973 analysis including Danuvius, we group it with the hispanopithecins because it appears 974 somewhat more derived than dryopithecins (Almécija et al., 2021). 975 Among dryopithecins, the distinctiveness of both Pierolapithecus and Anoiapithecus from 976 Dryopithecus has been accepted by various researchers (Pickford, 2012; Fleagle, 2013; 977 Böhme et al., 2019; Andrews, 2020) but questioned by Begun (2009, 2010, 2015), who 978 argued that the two former might be junior synonyms of Dryopithecus, mostly based on 41 979 claims of distortion (see SOM S3 for further details). However, preliminary results of a digital 980 reconstruction of the Pierolapithecus cranium (Pugh et al., 2022) are consistent with its stem 981 hominid status and difficult to reconcile with craniodental differences relative to 982 Anoiapithecus (Moyà-Solà et al., 2009b; Pérez de los Ríos et al., 2012). In our opinion, the 983 distinction of the three monotypic dryopithecin genera is supported by both dental and 984 cranial differences (Moyà-Solà et al., 2004, 2009a, 2009b; Alba, 2012; Pérez de los Ríos et al., 985 2012; Alba et al., 2013; Fortuny et al., 2021), although more complete remains would be 986 required to clarify further their phylogenetic relationships relative to both kenyapithecines 987 and hispanopithecins. ‘Sivapithecus’ occidentalis Villalta Comella and Crusafont Pairó, 1944, 988 previously considered a nomen dubium (Moyà-Solà et al., 2004; Alba, 2012) or synonymized 989 with Neopithecus brancoi (Schlosser, 1901) by Pickford (2012), is here included in the 990 Dryopithecini as a species inquirenda following Alba et al. (2020), as this nominal species is 991 potentially a junior synonym of either Pi. catalaunicus Moyà-Solà et al., 2004 or An. 992 brevirostris Moyà-Solà et al, 2009b. 993 With regard to hispanopithecins, following the opinion of Begun and colleagues (Begun 994 and Kordos, 1993; Kordos and Begun, 1997; Begun, 2002a), the binomen Dryopithecus 995 brancoi (Schlosser, 1901) was formerly considered by many authors as a taxonomically valid 996 species including the hispanopithecin remains from Hungary (e.g., Moyà-Solà and Köhler, 997 1995; Cameron, 2004, 2005; Ersoy et al., 2008; Kelley et al., 2008). In turn, the Spanish 998 hispanopithecin remains were included in one or more species of Dryopithecus (e.g., Begun 999 et al., 1990; Harrison, 1991; Begun, 1992b, 1994, 2002a; Moyà-Solà and Köhler, 1993, 1995, 1000 1996). Alternatively, Andrews et al. (1996) considered D. brancoi a nomen dubium and 1001 advocated the inclusion of the Hungarian material in Dryopithecus carinthiacus Mottl, 1957, 1002 which is here considered a junior synonym of D. fontani (Begun, 2002a; Begun et al., 2006; 42 1003 Casanovas-Vilar et al., 2011). Following most recent authors (Moyà-Solà et al., 2009a; 1004 Casanovas-Vilar et al., 2011; Begun, 2015; Alba et al., 2020; but see Pickford, 2012), both 1005 Neopithecus Abel, 1902 and Neopithecus brancoi are here considered nomina dubia. 1006 Furthermore, following Moyà-Solà et al. (2009a) and Begun (2009), Dryopithecus is restricted 1007 to Middle Miocene dryopithecines. Moyà-Solà et al. (2009a) resurrected Hispanopithecus 1008 not only for Late Miocene dryopithecines from Spain, as sometimes done by previous 1009 authors (Cameron, 1997a, 1999, 2004; Almécija et al., 2007), but also for those from 1010 Hungary, which were assigned by Begun (2009) to Rudapithecus hungaricus Kretzoi, 1969. 1011 Subsequently, Alba and coauthors (Casanovas-Vilar et al., 2011; Alba, 2012; Alba et al., 1012 2012a, 2012b) distinguished Hispanopithecus and Rudapithecus only at the subgenus rank, 1013 but following most recent authors (e.g., Begun, 2015; Gunz et al., 2020; Urciuoli et al., 1014 2021a) here both taxa are distinguished at the genus rank. The distinction of 1015 Hispanopithecus crusafonti (Begun, 1992b) from Hispanopithecus laietanus Villalta Comella 1016 and Crusafont Pairó, 1944 (Begun, 1992b, 2002a; Cameron, 1999) has been questioned by 1017 some authors (Harrison, 1991; Andrews et al., 1996; Ribot et al., 1996). However, they are 1018 here distinguished based on dental morphology (Alba, 2012; Alba et al., 2012a; Fortuny et 1019 al., 2021)—albeit recognizing that the proper genus allocation of H. crusafonti is difficult to 1020 evaluate due to the lack of more complete cranial remains. 1021 The latest occurring dryopithecine (Agustí et al., 2020), ?Udabnopithecus garedziensis 1022 Burchak-Abramovich and Gabashvili, 1945, has been variously synonymized with D. fontani 1023 (e.g., Szalay and Delson, 1979) or left as incertae sedis within the Dryopithecinae (Andrews 1024 et al., 1996). More recently, it has been considered by some a distinct species of 1025 Dryopithecus (Gabunia et al., 2001; Agustí et al., 2020). Given the limited evidence available 1026 (a maxillary fragment with P4–M1) and the current more restricted usage of Dryopithecus 43 1027 (see above), the inclusion in Dryopithecus does not appear warranted and we prefer to 1028 provisionally leave ?Udabnopithecus as incertae sedis at the tribe rank, denoting the 1029 uncertain taxonomic validity of the genus with a question mark. We also leave 1030 ‘Dryopithecus’ wuduensis, known on the basis of a partial mandible from the Late Miocene 1031 (~8–6 Ma) of China (Xue and Delson, 1988), as family incertae sedis. The inclusion of this 1032 species in Dryopithecus has been questioned (as ?Dryopithecus wuduensis) or even ruled out 1033 by several authors (Kelley, 2002; Harrison, 2005, 2006; Gilbert et al., 2020b), and most 1034 recently Pugh et al. (2020) identified closer similarities to stem hominoids. Given that the 1035 referral of this species to Dryopithecus can be discounted, we put the genus name within 1036 quotation marks until the allocation of the species is clarified. 1037 1038 3.7. Crown Hominidae 1039 Ponginae Five genera of Miocene pongines are recognized here (Table 2), being recorded 1040 from the late Middle Miocene (~13 Ma) to the latest Miocene (~6 Ma; Table 1). Besides their 1041 geographic distribution in Asia, these genera are customarily considered pongines based on 1042 the possession of derived craniodental features displayed by extant orangutans (e.g., Kelley, 1043 2002)—even though this has been disputed for Lufengpithecus Wu, 1987 during the last 1044 decade (see below). The most comprehensive cladistic analysis of crown hominids published 1045 so far (Pugh, 2022) found support for the inclusion of non-kenyapithecine hominids from 1046 Asia in the same clade as Pongo, with the single exception of a species thus far included in 1047 Lufengpithecus. To reflect the internal phylogenetic relationships of pongines, we follow 1048 Alba (2012) in distinguishing three tribes, with genera tentatively arranged in accordance 1049 with Pugh’s (2022) results. The apparently most plesiomorphic pongines (Ankarapithecus 1050 and Lufengpithecus) are included in the tribe Lufengpithecini Alba, 2012, whereas 44 1051 Indopithecus von Koenigswald, 1949 is included (together with Sivapithecus and the 1052 Pleistocene Gigantopithecus) in the Sugrivapithecini Simonetta, 1958 (regarding the year of 1053 publication of Simonetta's work, see SOM S2.3), and Khoratpithecus Chaimanee et al., 2004 1054 is grouped with extant Pongo in the Pongini Elliot, 1913. 1055 Sivapithecus, from the late Middle and Late Miocene of Indo-Pakistan (13.0–7.5 Ma), is 1056 the fossil pongine whose cranial morphology is the most complete, showing extensive 1057 similarities with orangutans; these include an airorynchous cranium, distinct supraorbital 1058 costae, tall and narrow orbits with their inferior margin well above the superior end of the 1059 nasal aperture, narrow interorbital space, and horizontal and procumbent premaxilla that 1060 considerably overlaps the hard palate, configuring a long and very narrow incisive canal as 1061 well as a smooth subnasal floor (Pilbeam, 1982; Ward and Pilbeam, 1983; Ward and Brown, 1062 1986; Brown and Ward, 1988; Ward, 1997a; Kelley, 2002; Brown et al., 2005). In contrast, 1063 the postcranium of Sivapithecus appears much less derived than that of Pongo, combining 1064 some modern hominoid-like features (such as the morphology of the distal humerus) with 1065 an otherwise plesiomorphic postcranial morphology, more consistent with a pronograde 1066 body plan suitable for emphasis on powerful-grasping and cautious, above-branch arboreal 1067 quadrupedalism (Pilbeam et al., 1990; Rose, 1997; Madar et al., 2002; Morgan et al., 2015). 1068 As a result of their cranial similarities, a close phylogenetic link between Sivapithecus and 1069 Pongo has received much support from cladistic analyses (Pugh, 2022 and references 1070 therein). The latter analysis, however, did not consistently recover a Pongo + Sivapithecus 1071 clade exclusive of all Khoratpithecus species. For this reason, we prefer to keep Sivapithecus, 1072 along with other extinct taxa presumably more closely related to it than to Pongo, in a 1073 different tribe. Although the intuitive name for this tribe would be Sivapithecini Pilbeam et 1074 al., 1977 (e.g., Andrews, 1992; Kelley, 2002), three names of the family group— 45 1075 Ramapithecini Simonetta, 1958, Bramapithecini Simonetta, 1958, and Sugrivapithecini 1076 Simonetta, 1958—have priority. The latter was used by Szalay and Delson (1979) and Alba 1077 (2012) for the tribe including Sivapithecus, but neither of them qualify as First Reviser (ICZN, 1078 1999: Art. 24.2; see SOM S2.3). We therefore act as First Reviser and choose Sugrivapithecini 1079 as the senior synonym of Ramapithecini and Bramapithecini for the tribe including 1080 Sivapithecus. 1081 We follow Kelley (2002, 2005) in tentatively distinguishing three species of Sivapithecus, 1082 while recognizing that alternative taxonomic schemes (see discussion in Kelley, 2005; 1083 Pickford, 2010; Bhandari et al., 2018; Gilbert et al., 2019) might ultimately prove to be more 1084 satisfactory, because only Sivapithecus parvada Kelley, 1988 seems at present well 1085 diagnosed from other species of the genus based on dental size and shape (Kelley, 1988, 1086 2002, 2005). For example, Kelley (2005) tentatively assigned the specimens from 1087 Haritalyangar to Sivapithecus cf. sivalensis (Lydekker, 1879), but admitted that this sample 1088 could alternatively include two different species, given the high levels of molar size variation 1089 and apparent sexual dimorphism (Scott et al., 2009). According to the present taxonomic 1090 arrangement, Sivapithecus simonsi Kay, 1982 is considered a synonym of Sivapithecus 1091 indicus Pilgrim, 1910 (Kelley, 2005). 1092 We also include in the Sugrivapithecini the genus Indopithecus, from the Late Miocene 1093 (~8.9–8.6 Ma) of India and Pakistan. Its type species, Indopithecus giganteus (Pilgrim, 1915), 1094 was originally described in Dryopithecus based on a single lower molar and later transferred 1095 to Indopithecus. Subsequently, Gigantopithecus bilaspurensis Simons and Chopra, 1969 was 1096 described based on a relatively complete mandible. The two species were synonymized by 1097 Szalay and Delson (1979), who used the combination Gigantopithecus giganteus. However, 1098 during the last decades most authors have supported the generic distinction between 46 1099 Indopithecus and Gigantopithecus blacki von Koenigswald, 1935 (Cameron, 2004; Alba, 1100 2012; Begun, 2015; Zhang and Harrison, 2017; Chaimanee et al., 2022; Pugh, 2022), which 1101 displays more strongly molarized lower premolars and other dental differences relative to 1102 Indopithecus (Kelley, 2002; Zhang and Harrison, 2017). Gigantopithecus is recorded from the 1103 Pleistocene (~2.0–0.3 Ma) of China, Vietnam, and Thailand (Zhang et al., 2014; Zhang and 1104 Harrison, 2017), and its pongine status has been recently supported by paleoproteomic data 1105 (Welker et al., 2019). Similarities in molar occlusal morphology between Indopithecus and 1106 Sivapithecus could justify their inclusion in the same genus, but the larger size and some 1107 similarities in mandibular and P3 morphology with the much younger and larger 1108 Gigantopithecus suggest that Indopithecus might be a basal member of the Gigantopithecus 1109 lineage (Kelley, 2002; Zhang and Harrison, 2017; Pugh, 2002). 1110 The subnasal morphology of Sivapithecus and orangutans is also shared to a large extent 1111 by Khoratpithecus, which includes four species from the late Middle to latest Miocene 1112 (~12.4–6.0 Ma) of southeastern Asia (Begun and Güleç, 1998; Kelley, 2002; Begun, 2015; 1113 Chaimanee et al., 2019, 2022). The nasoalveolar configuration of a palate—initially 1114 attributed to cf. Khoratpithecus (Chaimanee et al., 2019) but recently assigned by 1115 Chaimanee et al. (2022) to the type species of the genus, Khoratpithecus piriyai Chaimanee 1116 et al., 2004—closely resembles the pongine condition shared by Sivapithecus and Pongo but 1117 displays larger incisive canal and fossa, thus being less derived than that of Sivapithecus 1118 (Chaimanee et al., 2019, 2022). In contrast, Khoratpithecus has been proposed as the 1119 pongine most closely related to orangutans based on its symphyseal morphology and the 1120 lack of anterior digastric fossa (Chaimanee et al., 2003, 2004, 2006, 2019, 2022; Jaeger et al., 1121 2011). These features can be ascertained in both Kh. piriyai and Khoratpithecus 1122 ayeyarwadyensis Jaeger et al., 2011, which in our opinion supports the inclusion of the 47 1123 genus in the same tribe as orangutans. On the other hand, Pugh (2022) only recovered a 1124 sister-taxon relationship with Pongo in the case of Kh. ayeyarwadyensis—suggesting that 1125 additional cranial remains would be required to better ascertain the relationships between 1126 Pongo, Sivapithecus, and Khoratpithecus. 1127 The inclusion of other non-kenyapithecine hominid genera from the Miocene of Asia 1128 (Ankarapithecus and Lufengpithecus) in the Ponginae, and their classification within a single 1129 tribe Lufengpithecini is more debatable because the group might ultimately prove 1130 paraphyletic. Both Ankarapithecus meteai Ozansoy, 1957 from the Late Miocene (9.8 Ma) of 1131 Turkey and the species previously included in Lufengpithecus, from the late Middle to latest 1132 Miocene of China, retain a more plesiomorphic stepped subnasal configuration (Begun and 1133 Güleç, 1998; Brown et al., 2005; Kelley and Gao, 2012; Ji et al., 2013) more similar to that of 1134 dryopithecines (Begun, 1994; Moyà-Solà and Köhler, 1995; Brown et al., 2005; Moyà-Solà et 1135 al., 2009a, 2009b; Pérez de los Ríos et al., 2012; Pugh, 2022). Nevertheless, the subnasal 1136 morphology of Ankarapithecus appears somewhat derived toward the condition of other 1137 pongines, being interpreted as the plesiomorphic condition for this subfamily—as further 1138 supported by cladistic analyses (Begun and Güleç, 1998; Pugh, 2022), although it could be 1139 alternatively interpreted as compatible with a stem hominid status (Alpagut et al., 1996; 1140 Kappelman et al., 2003). 1141 The genus Lufengpithecus has traditionally been considered a pongine (e.g., Schwartz, 1142 1990, 1997; Kelley, 2002; Alba, 2012; Begun, 2015), but evidence supporting its pongine 1143 status is more debatable than in the case of Ankarapithecus and further complicated by the 1144 existence of multiple species with a complex nomenclatural history (Harrison et al., 2002). 1145 Kelley (2002) tentatively distinguished three species of Lufengpithecus: Lufengpithecus 1146 lufengensis (Xu et al., 1978), which is the type species of the genus; Lufengpithecus 48 1147 keiyuanensis (Woo, 1957); and Lufengpithecus hudienensis (Zhang et al., 1987). In contrast, 1148 Harrison et al. (2002) and Harrison (2006) favored a two species taxonomic scheme, by 1149 considering that L. hudienensis was a junior subjective synonym of L. keiyuanensis. Yet 1150 another species, Lufengpithecus yuanmouensis Zheng and Zhang, 1997 was described based 1151 on a juvenile cranium from Yuanmou as the holotype, being considered a junior subjective 1152 synonym of either L. hudienensis or L. keiyuanensis, according to Kelley (2002) and Harrison 1153 et al. (2002), respectively. The redescription of juvenile crania attributed to L. hudienensis 1154 and L. cf. lufengensis, the two younger species of Lufengpithecus (~8–6 Ma), not only 1155 showed substantial differences between these species but also the lack of crown hominoid 1156 (either pongine or hominine) synapomorphies—not being attributable to their ontogenetic 1157 stage (Kelley and Gao, 2012; Ji et al., 2013). Subsequent cladistic analysis recovered 1158 Lufengpithecus as more closely related to dryopithecines than to Pongo or Sivapithecus 1159 (Nengo et al., 2017; Gilbert et al., 2020a), which could justify transferring the Lufengpithecini 1160 into the Dryopithecinae. Nevertheless, most recently Pugh (2022) did not recover the 1161 monophyly of Lufengpithecus and supported a pongine status for L. lufengensis but not L. 1162 hudienensis, while the older species L. keiyuanensis (~12 Ma) could not be analyzed. 1163 On the basis that Lufengpithecus might be polyphyletic, Gilbert et al. (2020b) left this 1164 genus as incertae sedis within the Hominoidea. However, we consider more advisable to 1165 provisionally keep the Lufengpithecini (for L. lufengensis) within the Ponginae while 1166 transferring ‘L.’ hudienensis to a different genus, for which the nomen Sinopithecus Zhang et 1167 al., 1990 is available with this species as its type (Harrison et al., 2002; Harrison, 2006). This 1168 is consistent with the marked cranial differences previously noted between these taxa 1169 (Kelley and Gao, 2012; Ji et al., 2013; Kelley, 2017). We thus formally reassign the species to 1170 Sinopithecus hudienensis, which is left as subfamily incertae sedis until it is clarified whether 49 1171 it belongs to the Dryopithecinae, the Ponginae, or neither. The third and least well-known 1172 species of Lufengpithecus distinguished by Kelley (2002) and Ji et al. (2013) is further 1173 tentatively included in Sinopithecus, as previously done by Zhang et al. (1990), given the 1174 greater dental similarities with S. hudienensis (Harrison et al., 2002; Harrison, 2006). This 1175 arrangement must be considered very tentative until additional until additional evidence 1176 enables a better assessment of the phylogenetic relationships between Lufengpithecus, 1177 Sinopithecus, putative basal pongines (Ankarapithecus), and dryopithecines (particularly 1178 hispanopithecins). It is remarkable that the few postcranial remains of Lufengpithecus, 1179 unlike those of Sivapithecus (but similar to hispanopithecins such as Hispanopithecus and 1180 Rudapithecus, see above), are indicative (particularly based on phalangeal curvature) of 1181 suspensory behaviors (Deane and Begun, 2008; Begun, 2015; Zhang et al., 2020). 1182 Graecopithecini The Late Miocene genera Ouranopithecus and Graecopithecus von 1183 Koenigswald, 1972, from the Late Miocene (9.7–7.2 Ma) of Greece and Turkey, are included 1184 in the tribe Gracopithecini Cameron, 1997b, whereas an isolated upper premolar from 1185 Bulgaria (7.2 Ma; Spassov et al., 2012; Böhme et al., 2017) is assigned to Graecopithecini 1186 indet. The genus Graecopithecus has been variously considered distinct from (Begun, 2002a, 1187 2009, 2015; Koufos and de Bonis, 2005; Begun et al., 2012; Böhme et al., 2017; Fuss et al., 1188 2017) or synonymous with (Martin and Andrews, 1984; Andrews et al., 1996; Cameron, 1189 1997a, 1997b; Smith et al., 2004) Ouranopithecus, in which case the former would take 1190 precedence. Most authors favoring the latter view have indeed synonymized the respective 1191 type species, except for Cameron (1997b), who considered them distinct. In the light of 1192 recently published evidence (Fuss et al., 2017; Pugh, 2022), we favor the view that these two 1193 genera and species are distinct but likely closely related. Following the cladistic results 1194 summarized in the following paragraph, we include both Graecopithecus and 50 1195 Ouranopithecus within a single tribe Graecopithecini Cameron, 1997b, with Ouranopithecini 1196 Begun, 2009 being considered its junior subjective synonym (see SOM S2.4 for further 1197 details). 1198 Only the older graecopithecin species, Ouranopithecus macedoniensis (de Bonis et al., 1199 1974) from Greece (9.7–8.8 Ma), is known from craniodental material (de Bonis et al., 1990; 1200 de Bonis and Koufos, 1993, 1994). This species is characterized by larger body size than 1201 dryopithecines (Kappelman et al., 2003), hyperthick molar enamel (Smith et al., 2004)— 1202 interpreted as an adaptation for a sclerocarpic diet (Ungar, 1996; Begun, 2009; DeMiguel et 1203 al., 2014)—and a subnasal morphology most similar to that of dryopithecines (de Bonis and 1204 Melentis, 1987; de Bonis and Koufos, 1994; Moyà-Solà and Köhler, 1995). The phylogenetic 1205 relationships of Ouranopithecus have been much debated, being interpreted as an early 1206 hominin by some authors (de Bonis et al., 1990, 1998; de Bonis and Koufos, 1993, 1994, 1207 2004; Koufos and de Bonis, 2005), or alternatively as a stem pongine (Moyà-Solà and Köhler, 1208 1995; Agustí et al., 1996; Köhler et al., 2001), a stem hominid (Alba, 2012), or a stem 1209 hominine (Begun, 1994, 2001, 2002a, 2009, 2010, 2015; Cameron, 2004). Ouranopithecus 1210 has been recovered as a stem hominine by cladistic analyses (Begun et al., 1997, 2012; 1211 Cameron, 1997a; Nengo et al., 2017; Gilbert et al., 2020a; Pugh, 2022). However, while 1212 Begun et al. (1997, 2012) recovered Ouranopithecus and dryopithecines as successive sister 1213 taxa of extant hominines, both Cameron (1997a) and more recent cladistic analyses (Nengo 1214 et al., 2017; Gilbert et al., 2020a; Pugh, 2022) supported a hominine status only for 1215 Ouranopithecus. It has also been recently argued that Graecopithecus is a hominin (Fuss et 1216 al., 2017) but such a contention has not been supported by subsequent cladistic analyses 1217 (Benoit and Thackeray, 2017; Pugh, 2022). Nevertheless, Pugh (2022) yielded tentative 1218 support to a clade of stem hominines including both Ouranopithecus and Graecopithecus. 51 1219 Although we favor the view that these genera are closely related, following Pugh’s (2022) 1220 results, we consider it premature to classify the Graecopithecini in the Homininae. While 1221 Pugh’s (2022) analyses are thus far the most thorough in supporting the hominine status of 1222 graecopithecins, we consider that their hominine status is plausible but insufficiently 1223 supported at present (for further explanations, see SOM S4 and SOM Fig. S1), and even 1224 unlikely unless at least some dryopithecines are also considered hominines (e.g., Begun et 1225 al., 2012). Therefore, we provisionally prefer to leave graecopithecins as incertae sedis at the 1226 subfamily rank while noting that, on both morphologic and chronostratigraphic grounds, 1227 graecopithecins might have been derived from dryopithecines irrespective of whether they 1228 are closely related to Nakalipithecus Kunimatsu et al., 2007 and/or hominines (see below). 1229 Homininae The ape record from the latest Middle and Late Miocene of Africa is quite 1230 meager compared to the panoply of Eurasian taxa during the same time interval. Isolated 1231 teeth from Middle Miocene (~12.8–12.0 Ma) localities of the Ngorora Formation, Kenya 1232 have been interpreted by some authors as hominines (Bishop and Chapman, 1970; Pickford 1233 and Senut, 2005) or stem hominoids (Hill and Ward, 1998; Hill et al., 2002), and the same 1234 applies to dental specimens from the latest Miocene (~6 Ma) of the Lukeino Formation 1235 (Pickford, 1975; Hill and Ward, 1988; Pickford and Senut, 2005), considered by some related 1236 to African apes (Pickford and Senut, 2005). All these samples are too meager to reach 1237 definite conclusions beyond a probable hominid status (e.g., Harrison, 2010a). A very 1238 fragmentary mandibular fragment, tentatively dated to the Late Miocene (Pickford et al., 1239 2008, 2009b), records the presence of hominoids in Niger but does not allow further 1240 taxonomic precision (Harrison, 2010a) 1241 Additionally, two great ape genera have been recognized from the Late Miocene in Africa, 1242 being represented by fragmentary dentognathic remains: Nakalipithecus from Kenya (9.9– 52 1243 9.8 Ma) and Chororapithecus Suwa et al., 2007 from Ethiopia (~8.0 Ma). Nakalipithecus was 1244 originally proposed as a stem hominine likely related to Ouranopithecus (Kunimatsu et al., 1245 2007)—as further favored by Harrison (2010a, 2010b), who on the basis of the more 1246 plesiomorphic dental features of Nakalipithecus supported an African origin of hominines. 1247 More recently, Pugh’s (2022) cladistic results supported a stem hominine status for 1248 Nakalipithecus and, more tentatively, a possible link with European graecopithecins, but the 1249 features used in support of such links (Kunimatsu et al., 2007; Pugh, 2022) are too 1250 ambiguous (see SOM S4 for further details). Chororapithecus was originally interpreted as a 1251 member of the gorilla lineage based on some details of molar shape and overall size (Suwa 1252 et al., 2007), but this phylogenetic link has been questioned by Harrison (2010a), who 1253 suggested instead a link (and possible synonymy) with Samburupithecus. Pugh (2022) found 1254 the cladistic topology of Chororapithecus to be unstable because the available hypodigm 1255 does not preserve enough phylogenetically informative features to resolve its relationships 1256 and also owing to the fact that the purported gorillin synapomorphies identified by Suwa et 1257 al. (2007) have not been examined in a broad hominoid sample. Based on Pugh’s (2022) 1258 results for Nakalipithecus and the purported similarities between Chororapithecus and 1259 gorillas noted by Suwa et al. (2007), we tentatively include both genera in the Homininae as 1260 tribe incertae sedis, given that additional remains would be required to better support the 1261 hypothesized relationships with graecopithecins and gorillins, respectively. 1262 1263 4. Miocene ape paleobiodiversity, phylogeny, and paleobiogeography 1264 4.1. Paleobiodiversity dynamics 1265 The chronostratigraphic range and geographic distribution of Miocene ape species are 1266 summarized in Table 3, whereas species-locality occurrences with primary references and 53 1267 the age of each Miocene ape-bearing locality are reported respectively in SOM Tables S1 and 1268 S2. Up to 56 species of Miocene apes are considered valid in this work (plus two species of 1269 uncertain taxonomic validity), being included in 35 genera (plus two of doubtful or uncertain 1270 taxonomic validity). The geographic distribution of these taxa, including occurrences 1271 indeterminate at the species rank, during the Early, Middle, and Late Miocene is depicted in 1272 Figures 3, 4, and 5, respectively. In turn, the chronostratigraphic ranges reported in Table 3 1273 for Miocene ape species, together with other citations indeterminate to species rank, have 1274 been summarized at the genus level in Figure 6. 1275 The chronostratigraphic ranges reported in Table 3 simultaneously reflect the occurrence 1276 of each species in differently aged localities and the dating uncertainties for these localities, 1277 but may be used to provide a first approximation to the paleobiodiversity dynamics of 1278 Miocene apes through time (Foote, 2000; see SOM S5 and SOM Table S3 for further details). 1279 Range-through (total) diversity (Fig. 7a) and standing diversity (Fig. 7b) at the species level 1280 yield similar diversity patterns, with a diversity peak during the Early Miocene (~21–20 Ma) 1281 and an even more marked peak during the Late Miocene (~9–8 Ma). Both metrics reflect a 1282 more or less marked decrease in diversity during the late Early Miocene, followed by an 1283 irregular recovery throughout the Middle Miocene until reaching the highest diversity well 1284 within the Late Miocene, followed by an abrupt decline thereafter. Changes in diversity 1285 through time are more marked in the plot based on standing diversity (Fig. 7b), which more 1286 adequately minimizes sampling biases than total diversity counts (Fig. 7a). However, both 1287 patterns support that the Middle (Fig. 4) and Late (Fig. 5) Miocene ape radiation—largely 1288 driven by the diversification of great apes throughout Eurasia—was of comparable 1289 magnitude to (if not greater than) the Early Miocene radiation of putative stem apes (Fig. 1290 3)—confined to Afro-Arabia except for the Engelswies tooth fragment. 54 1291 It is noteworthy that the Mid-Miocene Climatic Optimum (Foster et al., 2012) is 1292 characterized by a low diversity of Miocene apes, which is followed by a progressive increase 1293 in diversity that continues into the Late Miocene despite a progressive trend toward 1294 increased cooling and seasonality initiated at ~14 Ma. This supports the interpretation that 1295 climatic 'deterioration' and associated paleoenvironmental changes might have triggered 1296 the adaptive radiation of crown hominoids in Eurasia (Casanovas-Vilar et al., 2011), as a 1297 result of progressive locomotor and dietary specialization (DeMiguel et al., 2014). A 1298 temporary drop in diversity at ~11 Ma appears artifactual to a large extent, both in Eurasia 1299 (where the poor sampling of hominoids by this time has been previously noted; Alba et al., 1300 2022) and in Africa (as indeterminate remains older than 12 Ma and the presence of several 1301 taxa from 10 Ma onward suggest a greater diversity than currently recorded around the 1302 Middle/Late Miocene boundary). In contrast, the marked decline in diversity during the 1303 latest Miocene mostly reflects the local extinction of hominoids in Western Eurasia and their 1304 progressive decline in Eastern Eurasia and probably Africa (hominins excluded). This has 1305 been interpreted as the result of changes in vegetation structure related to progressive 1306 climate cooling and aridification (Agustí et al., 2003; Casanovas-Vilar et al., 2011; Marmi et 1307 al., 2012; Almécija et al., 2021), roughly coinciding with the spread or more open and arid 1308 environments across the Old World (Kaya et al., 2018). 1309 1310 4.2. Phylogenetic uncertainties: Craniodental vs. postcranial evidence 1311 Although both climatic and paleogeographic factors seemingly played a role in Miocene 1312 ape diversification, important paleobiogeographic issues remain to be deciphered due to 1313 phylogenetic uncertainties. For example, it is unclear whether hominids and hylobatids 1314 diverged in Africa during the Early Miocene and separately dispersed into Eurasia by the 55 1315 Middle Miocene, or whether they diverged in Eurasia following a single dispersal event. It is 1316 similarly uncertain whether hominines and pongines evolved as a result of a vicariant event 1317 in Eurasia and Africa, respectively, or whether they diverged in one of these continents and 1318 subsequently dispersed to the other. Gilbert et al. (2020b) examined several recent 1319 phylogenetic hypotheses from a paleobiogeographic perspective and concluded that they 1320 implied between one and four hominoid dispersal events between Africa and Eurasia and 1321 zero to two between Europe and Asia. Later in this section, we will discuss the pros and cons 1322 of various competing paleobiogeographic scenarios in the light of phylogenetic evidence, but 1323 before doing so it is worth discussing to what extent the phylogenetic relationships favored 1324 by cladistic analyses published during the last decade (Nengo et al., 2017; Rossie and Hill, 1325 2018; Gilbert et al., 2020a; Pugh, 2022) appear reliable. 1326 Discrepancies between most parsimonious topologies and chronostratigraphic ranges are 1327 to be expected due to the multiple biases of the fossil record, but can also by caused by 1328 inaccurate phylogenies, particularly in the light of abundant postcranial homoplasy. Previous 1329 attempts to determine whether craniodental or postcranial features are more homoplastic 1330 proved rather inconclusive, finding higher levels in craniodental (Finarelli and Clyde, 2004) or 1331 postcranial (Young, 2005) characters. Any attempt to estimate craniodental and postcranial 1332 homoplasy based on a most parsimonious cladogram derived from both anatomical areas 1333 simultaneously results in circular reasoning and is biased by the number of characters in 1334 each subset and their actual degree of homoplasy (which is unknowable). An alternative way 1335 to tackle this issue is to compare the results provided by separate analyses of craniodental 1336 and postcranial features, as done by Pugh (2022). 1337 The most parsimonious cladograms obtained by Pugh (2022: Fig. 4) based on craniodental 1338 and postcranial characters separately evince important similarities and differences, and 56 1339 further highlight those phylogenetic links that are exclusively or mostly based on 1340 craniodental data. The results supported by both craniodental and postcranial evidence 1341 include the sequential branching of equatorines, kenyapithecines, and dryopithecines. 1342 Hispanopithecines are also recovered as less basal than dryopithecins + kenyapithecines in 1343 all analyses, but the craniodental evidence supports H. laietanus as a pongine and R. 1344 hungaricus as a hominine—casting even more doubts on the monophyly of dryopithecines 1345 as conceived here. The results of the total morphological evidence analysis that are only 1346 supported by craniodental data include: (1) the stem hylobatid status of the clade 1347 constituted by Oreopithecus and Samburupithecus; (2) the polyphyly of Lufengpithecus s.l. 1348 and Khoratpithecus; and (3) the recovery graecopithecins in a hominine clade also including 1349 Nakalipithecus. This is not surprising given that most of these taxa were not included in the 1350 postcranial analysis, with the exception of Oreopithecus. However, it is noteworthy that the 1351 clade including graecopithecins + Nakalipithecus is not recovered as sister to crown 1352 hominines, but as sister of hominins—supporting the view that, unless interpreted as 1353 basalmost hominins (e.g., de Bonis and Koufos, 2004), the hominine status of these taxa in 1354 the total morphological evidence analysis is influenced by craniodental convergences with 1355 hominins related to powerful mastication (see discussion in Pugh, 2022). 1356 Most noteworthy for assessing the reliability of Pugh’s (2022) results based on total 1357 morphological evidence are the differences between the postcranial and craniodental 1358 results. The cladogram based on the former recovers the monophyly of crown hominoids, 1359 hylobatids, hominids, and hominines, but excludes from these clades all the analyzed 1360 Miocene apes except Si. sivalensis and Oreopithecus, which are recovered as crown 1361 hominids. Thus, besides failing to recover the pongine status of Sivapithecus, the postcranial 1362 data support a much less basal branching for both hylobatids and Oreopithecus. As a result, 57 1363 most of the Miocene apes recovered as hominids in the craniodental analysis (equatorines, 1364 kenyapithecines, dryopithecines, and Si. sivalensis) are recovered as stem hominoids based 1365 on the postcranial data. Oreopithecus exhibits the most extreme incongruency between 1366 craniodental and postcranial data, but the contrasting results obtained for hylobatids are 1367 also worrisome for the reliability of their topology in the total morphological evidence 1368 cladogram—intermediate between equatorines and kenyapithecines + dryopithecins—given 1369 its implications for distinguishing stem from crown hominoids. As in the case of 1370 Oreopithecus, the branching topology of hylobatids appears strongly influenced by 1371 postcranial similarities with hominids, which are functionally related to orthograde 1372 behaviors and most likely evolved in parallel to a large extent, as illustrated by Sivapithecus 1373 and Pierolapithecus (e.g., Larson, 1998; Moyà-Solà et al., 2004; Alba, 2012; Ward, 2015) and 1374 further favored by Pugh (2022). 1375 To account for the potential bias introduced by crown hominoid postcranial similarities, in 1376 Figure 8b we have depicted a cladogram that better reflects current main uncertainties 1377 about Miocene ape phylogeny while hypothesizing a more basal divergence of hylobatids 1378 (more consistent with the craniodental results) instead of the less basal divergence 1379 hypothesis currently favored by most parsimonious cladograms. The cladogram depicted in 1380 Figure 8b thus mainly differs from that of Figure 8a in the position of afropithecines and 1381 equatorines, which are considered basalmost stem hominids instead of advanced stem 1382 hominoids (leaving unresolved if they constitute a clade or successive sister taxa). Based on 1383 the consistency between craniodental and postcranial results about the more derived status 1384 of hispanopithecins, in Figure 8b we have further considered them less basal than 1385 dryopithecins while leaving unresolved if the latter constitute a clade with kenyapithecines. 58 1386 In the following subsections, we discuss some of the most vexing unresolved enigmas of 1387 Miocene ape evolution in terms of phylogeny and paleobiogeography by taking into account 1388 not only molecular estimates of divergence times and the additional evidence provided by 1389 the fossil record (chronostratigraphic ranges and geographic distribution), but also the more 1390 vs. less basal divergence hypotheses for hylobatids depicted in Figure 8. These unresolved 1391 issues are the following: (1) What are the closer phylogenetic relationships of Oreopithecus; 1392 (2) What is the evolutionary origin of hylobatids, i.e., where do we draw the cladistic 1393 boundary between stem and crown hominoids; (3) When and where did pongines and 1394 hominines diverge, and what is the role that dryopithecines and graecopithecins played in 1395 the origin and subsequent diversification of the hominid clade. 1396 1397 4.3. The perplexing enigma of Oreopithecus 1398 Uncertainties about the phylogenetic relationships of Oreopithecus, from the Late 1399 Miocene of Italy, are “especially perplexing because Oreopithecus is one of the best-known 1400 fossil primates” (Harrison and Rook, 1997: 328). A skeleton discovered in 1958 led Straus 1401 (1963) to conclude that Oreopithecus was either a ‘hominid’ s.s. or a member of a separate 1402 hominoid family, whereas Hürzeler (1958, 1968) more explicitly considered Oreopithecus a 1403 side branch of the human lineage. However, the bizarre dental morphology of Oreopithecus, 1404 coupled with its peculiar cranial morphology and modern hominoid-like postcranium led to 1405 varied interpretations over the years (see review in Delson, 1986). Harrison (1986, 1987a) 1406 advocated a phylogenetic link between Oreopithecus and nyanzapithecids, while supporting 1407 its hominoid status based on the contention that its postcranial similarities with extant apes 1408 “are so detailed that there seems little possibility that they could have been developed 1409 independently” (Harrison, 1987a: 541). Similar views were stated by Sarmiento (1987: 35): 59 1410 “given the evidence of the forelimb anatomy and associated shape of the thorax, this form 1411 [Oreopithecus] cannot be interpreted as anything but a hominoid”. Sarmiento (1987) noted 1412 that, in some aspects, Oreopithecus is more hylobatid-like than great ape-like, and put 1413 forward two possible interpretations: either Oreopithecus retained the crown-hominoid 1414 plesiomorphic condition; or Oreopithecus is a large-bodied hylobatid. 1415 Given Harrison’s (1986, 1987a, 1987b) views about the synapomorphic nature of crown 1416 hominoid postcranial similarities (see Section 2.3), Harrison and Rook (1997: 347) still 1417 maintained that “the postcranial characteristics shared by Oreopithecus and the extant 1418 hominoids are so pervasive throughout the skeleton that it is almost impossible to consider 1419 that these could have been developed independently to such a remarkable degree of detail 1420 in every anatomical region.” The same opinion was held by Pilbeam (1996, 1997), who 1421 considered Oreopithecus as the most secure modern Miocene hominoid. Nevertheless, 1422 Harrison and Rook (1997) abandoned Harrison’s (1986, 1987a) former views about a close 1423 phylogenetic link with nyanzapithecids and, like Moyà-Solà and Köhler (1997), hypothesized 1424 that Oreopithecus is a derived dryopithecine. The postcranial similarities between 1425 Oreopithecus and Hispanopithecus (Moyà-Solà and Köhler, 1996) were determinant in these 1426 proposals of a great ape status for Oreopithecus, even if also supported by some cranial 1427 features—such as the configuration of the incisive canal and the lack of subarcuate fossa 1428 (Harrison and Rook, 1997; Moyà-Solà and Köhler, 1997), whose reliability as hominid 1429 synapomorphies is currently doubtful (see next subsection). A hominid status for 1430 Oreopithecus would not only imply dental convergences with nyanzapithecids (see 1431 discussion in Rossie and Cote, 2022), but also that its cranial morphology is autapomorphic 1432 (Harrison and Rook, 1997; Moyà-Solà and Köhler, 1997; Alba et al., 2001) rather than 60 1433 symplesiomorphic for crown hominoids, as generally assumed (Harrison, 1987a; Sarmiento, 1434 1987; Begun et al., 1997; Begun, 2002a, 2013, 2015; Pugh, 2022). 1435 Begun et al.’s (1997, 2012) cladistic analyses recovered Oreopithecus as a stem hominid, 1436 and its hominid status was generally accepted until about a decade ago (e.g., Wood and 1437 Harrison, 2011; Alba, 2012), with some exceptions restating the link with nyanzapithecids 1438 (Benefit et al., 1998; Benefit and McCrossin, 2001). Subsequent cladistic analyses, including a 1439 wider representation of Early and Middle Miocene apes, recovered Oreopithecus as deeply 1440 nested within the nyanzapithecid clade (Nengo et al., 2017; Gilbert et al., 2020a)—in 1441 agreement with Harrison’s (1986, 1987a) former opinion. Most recently, Pugh’s (2022) 1442 analyses hinted at a possible close relationship between Oreopithecus and Samburupithecus 1443 and provided some support for considering both taxa as stem hylobatids. The link between 1444 Oreopithecus and Samburupithecus, given the previously noted dental similarities between 1445 the latter taxon and nyanzapithecids (Harrison, 2010a), is consistent with previous cladistic 1446 analyses supporting that Oreopithecus is a derived nyanzapithecid (Nengo et al., 2017; 1447 Gilbert et al., 2020a), in further agreement with previously noted similarities between 1448 Oreopithecus and members of this family (Leakey, 1968; Harrison, 1986, 1987a, 1992, 1449 2010a). In contrast, the stem hylobatid status recovered by Pugh (2022) must be considered 1450 poorly supported at present, given the lack of nyanzapithecids in her analyses and the highly 1451 divergent results obtained for Oreopithecus based on craniodental and postcranial 1452 characters. As noted by Pugh (2022), similarities between Oreopithecus with hylobatids 1453 include a few craniodental features present in some putative stem hominoids (e.g., elongate 1454 fourth premolars as in nyanzapithecids) and even stem catarrhines (an anteriorly protruding 1455 rhinion), thus being likely symplesiomorphic, coupled with multiple modern hominoid-like 1456 postcranial characteristics that are most likely homoplastic between crown hominoids. 61 1457 Therefore, Pugh (2022) concluded that Oreopithecus is most reasonably interpreted as a 1458 stem hominoid. 1459 That the link between Oreopithecus and hylobatids retrieved by Pugh (2022) is not 1460 entirely attributable to postcranial similarities is confirmed by the fact that craniodental 1461 features alone support this link (while postcranial characteristics favor a more advanced 1462 status toward hominids). Nevertheless, a stem hominoid status for Oreopithecus is further 1463 supported by the morphology of the inner ear semicircular canals, which despite some 1464 similarities in proportions with extant great apes is much more primitive and does not show 1465 particular similarities with that of hylobatids (Urciuoli et al., 2020, 2021a, 2021b). On the 1466 other hand, the homologies hypothesized between Oreopithecus and nyanzapithecids in 1467 lower molar cristids (Rossie and Cote, 2022) are suggestive but questionable, so it is 1468 probably too early to confidently rule out their independent evolution. Analyses of enamel– 1469 dentine junction shape (currently underway) would be required to better evaluate these 1470 dental similarities, but preliminary results indicate close morphometric affinities between 1471 Oreopithecus and stem catarrhines (Zanolli et al., 2022a), supporting further that the former 1472 is not a crown hominoid. Taken overall, current evidence lends greater support to the 1473 hypothesis that Oreopithecus is a late descendant of an ancient hominoid lineage (maybe 1474 nyanzapithecids) that dispersed from Africa into Tusco-Sardinia sometime before 8 Ma and 1475 acquired orthograde-related features independently from crown hominoids. However, given 1476 that the branching order between nyanzapithecids and crown hominoids does not appear 1477 reliable (see Section 5.1), the possibility that Oreopithecus and some nyanzapithecids are 1478 stem hylobatids deserves further consideration in the future. 1479 1480 4.4. The vexing problem of hylobatid origins 62 1481 The fossil record indicates that hominoids originated in the Oligocene (Stevens et al., 1482 2013; Hammond et al., 2019) and first radiated during the Early Miocene in Afro-Arabia 1483 (Harrison, 2010a), when this continent was isolated from Eurasia by the Tethys Seaway 1484 (Bernor, 1983; Seiffert, 2012; Begun et al., 2012). By ~19 Ma, the emergence of an 1485 intermittent land-bridge allowed intercontinental faunal exchanges between Africa and 1486 Eurasia (Harzhauser et al., 2007), even if temporarily interrupted by the Langhian 1487 transgression at ~16 Ma (Rögl, 1999), leading to the eventual dispersal of multiple catarrhine 1488 lineages into Eurasia at different times (Roos et al., 2019; Gilbert et al., 2020b). Stem 1489 catarrhines (pliopithecoids) were apparently the first group to disperse, being first recorded 1490 at ~19–18 Ma in Asia (Harrison and Gu, 1999; Begun, 2002b, 2017; Harrison, 2013; Harrison 1491 et al., 2020). Large-bodied putative hominids followed soon thereafter, being first recorded 1492 by kenyapithecines (Griphopithecus and Kenyapithecus) at ~16.5–14 in Europe and Turkey 1493 (Heizmann and Begun, 2001; Andrews and Kelley, 2007; Casanovas-Vilar et al., 2011), and 1494 subsequently by dryopithecines in Europe and pongines in Asia from ~13–12.5 Ma onward 1495 (Kelley, 2005; Alba, 2012; Begun, 2015; Gilbert et al., 2020b). However, the scarce record of 1496 Miocene hylobatids (Harrison, 2016; Ji et al., 2022), coupled with phylogenetic uncertainties 1497 (Fig. 8), hinders an adequate understanding of hylobatid origins. 1498 Molecular estimates strongly support that hylobatids and hominids diverged during the 1499 Early Miocene (i.e., before the Langhian transgression), probably ~20–17 Ma, although the 1500 large confidence intervals of some of these estimates do not completely rule out a later 1501 divergence in the early Middle Miocene: 20.3 (24.2–16.5) Ma (Perelman et al., 2011), 17.4 1502 (23.9–12.4) Ma (Springer et al., 2012), 20.3 (23.5–17.4) Ma (Finstermeier et al., 2013), and 1503 16.8 (17.6–15.9) Ma (Carbone et al., 2014). Given that their oldest unambiguous fossil 1504 record (Yuanmoupithecus) dates to the latest Miocene (8.2–7.1 Ma; Harrison, 2016; Ji et al., 63 1505 2022), hylobatids have a long ghost lineage. Furthermore, molecular data indicate that 1506 crown members of the group did not diverge from one another until the latest Miocene 1507 (Perelman et al., 2011; Springer et al., 2012; Finstermeier et al., 2013; Carbone et al., 2014), 1508 which implies that stem hylobatids might lack most of the derived features shared by 1509 Yuanmoupithecus and crown hylobatids. 1510 Decades ago, pliopithecoids were considered broadly ancestral to hylobatids (Hürzeler, 1511 1954; Zapfe, 1958, 1961; Simons and Fleagle, 1973; Andrews and Simons, 1977). However, 1512 such an assumption was based on superficial cranial and postcranial similarities and is no 1513 longer tenable given the widely-accepted stem catarrhine status of pliopithecoids (e.g., 1514 Zalmout et al., 2010; Stevens et al., 2013; Nengo et al., 2017; Gilbert et al., 2020a; Urciuoli et 1515 al., 2021b). The latter is supported by the lack of crown catarrhine synapomorphies and the 1516 retention of multiple plesiomorphic features (e.g., incompletely ossified ectotympanic, 1517 entepicondylar foramen in the distal humerus, and single hinge-like carpometacarpal thumb 1518 joint; Zapfe, 1961; Harrison, 1987b, 2005; Andrews et al., 1996; Begun, 2002b, 2017). 1519 According to this interpretation, hylobatids must represent an independent dispersal event 1520 from those of pliopithecoids. It has generally been assumed that hylobatids and large-bodied 1521 hominoids from Eurasia are the result of at least two different dispersal events from Africa 1522 (Moyà-Solà et al., 1999b; Roos et al., 2019; Gilbert et al., 2020a), either synchronous or 1523 diachronous. Gilbert et al. (2020a) suggested that this dispersal event took place from Africa 1524 into Asia just after the Mid-Miocene Climatic Optimum. However, in the light of the 1525 molecular estimates of crown hominoid divergence time reported above, hylobatids could 1526 have alternatively dispersed somewhat earlier, before than Langhian transgression (19–16 1527 Ma), like pliopithecoids (see above). Both alternatives are possible, as a pre-Langhian 64 1528 divergence time does not exclude a later dispersal event of hylobatid ancestors from Africa 1529 into Eurasia. 1530 Gilbert et al. (2020a) further hypothesized that hylobatids might have originated from a 1531 dendropithecid or purported stem hominoid (Proconsulidae s.l.) from the Early Miocene of 1532 Africa. Hypothetical basalmost hylobatids might be simply unknown or currently 1533 unrecognized as such among the multiple small-bodied catarrhine genera of uncertain 1534 affinities from Early Miocene of Africa. An origin from dendropithecids (sensu Harrison, 1535 2010a) is plausible on chronological grounds, as they are first recorded in the Early Miocene 1536 (e.g., Harrison, 2010a), but would imply that at least some dendropithecids are crown 1537 hominoids instead of the stem catarrhines (as supported by Rossie and Hill, 2018, who 1538 recovered Dendropithecus Andrews and Simons, 1977 and Simiolus Leakey and Leakey, 1987 1539 as basal nyanzapithecids). Cladistic analyses have generally recovered dendropithecids as 1540 stem catarrhines (Nengo et al., 2017; Gilbert et al., 2020a) or as stem hominoids at most 1541 (Rae, 1999, 2004; Zalmout et al., 2010; Alba et al., 2015; Rossie and Hill, 2018), such that a 1542 crown hominoid status for dendropithecids lacks cladistic support except perhaps for 1543 Micropithecus. Originally interpreted as a hylobatid relative based on facial morphology 1544 (Fleagle and Simons, 1978), Micropithecus and other dendropithecids were subsequently 1545 reinterpreted as stem catarrhines given the lack of crown catarrhine postcranial 1546 synapomorphies (e.g., Harrison, 1987b, 2002, 2010a), with cranial similarities with 1547 hylobatids being interpreted as symplesiomorphic. However, Micropithecus was tentatively 1548 considered a hominoid (‘proconsuloid’) by Begun (2015) based on its moderately developed 1549 molar cingula, and most recently recovered as a stem hylobatid more basal than 1550 Yuanmoupithecus by Ji et al. (2022). Even if the latter authors attributed this result to dietary 65 1551 convergence, the possibility that hylobatids may be closely related to some dendropithecids 1552 should be scrutinized further when more complete material becomes available. 1553 Alternatively, hylobatids (or even crown hominoids as a whole) might have evolved from 1554 putative stem hominoids from the Early or Middle Miocene of Africa, i.e., proconsulids or 1555 nyanzapithecids. The Oligocene record of both proconsulids and nyanzapithecids (Stevens et 1556 al., 2013; Hammond et al., 2019) supports an early divergence between these families, while 1557 afropithecids are not recorded until later. Although some recent cladistic analyses recovered 1558 a monophyletic nyanzapithecid clade that nests within afropithecids (Nengo et al., 2017; 1559 Gilbert et al., 2020a; but see Rossie and Hill, 2018 for a different topology), the Oligocene 1560 age of Rukwapithecus and another indeterminate nyanzapithecids (Stevens et al., 2013; 1561 Hammond et al., 2019) is at odds with its nesting well within the nyanzapithecid radiation. 1562 This suggests that the relationships between nyanzapithecids, afropithecids, and crown 1563 hominoids are not well resolved. The less basal divergence hypothesis for hylobatids (Fig. 8a) 1564 suggests that they are a dwarfed lineage (Pilbeam, 1996; Reichard et al., 2016) evolved from 1565 an afropithecid-like ancestor, which is compatible with molecular estimates and with either 1566 one or two dispersal events of crown hominoids into Eurasia during the Early and/or Middle 1567 Miocene. In contrast, the basal divergence hypothesis for hylobatids (Fig. 8b) favored by 1568 craniodental data suggests that they originated from a nyanzapithecid-like (or proconsulid- 1569 like) ancestor, is in better agreement with average divergence times estimated from 1570 molecular evidence, and requires two dispersal events of crown hominoids into Eurasia. The 1571 craniodental similarities shared between nyanzapithecids and hylobatids (e.g., relatively 1572 short face, broad interorbital distance, and projecting orbits; Nengo et al., 2017) may be 1573 interpreted as the plesiomorphic condition for crown hominoids and, hence, neither favor 1574 nor contradict a possible origin of hylobatids from more or less derived nyanzapithecids 66 1575 (which would imply that the latter are paraphyletic). Dendropithecids and proconsulids seem 1576 more unlikely ancestors for hylobatids given the results of cladistic analyses, but the 1577 plausibility of an afropithecid-like—as opposed to nyanzapithecid-like—ancestor, or a 1578 Eurasian divergence between hylobatids and hominids following a single dispersal event 1579 from Africa, largely depends on the extent to what most parsimonious cladistic analyses are 1580 biased by postcranial homoplasy between hylobatids and hominids. 1581 1582 4.5. The elusive origin of the great ape and human clade 1583 Equatorines The uncertain monophyly and phylogenetic relationships of afropithecids are 1584 illustrated by the fact that they have been hypothesized as stem hominids by various 1585 researchers (Andrews, 1992, 1996; Ward and Duren, 2002) whereas others have considered 1586 that at least afropithecines would precede the hylobatid–hominid divergence (e.g., Moyà- 1587 Solà et al., 2009b; Casanovas-Vilar et al., 2011; Alba, 2012; Begun, 2015; Andrews, 2020). As 1588 discussed in the preceding subsection, deciding whether afropithecids are stem hominoids, 1589 as favored by most parsimonious cladograms (Nengo et al., 2017; Gilbert et al., 2020a; Pugh, 1590 2022), instead of stem hominids, as supported by craniodental evidence (Pugh, 2022), 1591 depends on their branching sequence relative to hylobatids (Fig. 8). Afropithecids are 1592 recorded in the Early Miocene before 17 Ma, and probably at least by 20 Ma if the older 1593 dating of Morotopithecus is correct (see Section 3.4), which roughly coincides with the 1594 average divergence time between hylobatids and hominids estimated on molecular grounds, 1595 and is thus compatible with both possibilities. 1596 The phylogenetic relationships of the equatorine Nacholapithecus are particularly 1597 controversial. Some authors have considered it as a stem hominid (Alba, 2012; Kunimatsu et 1598 al., 2019) largely based on the possession of an elongated and procumbent premaxilla that 67 1599 slightly overlaps the maxillary palatine process (Ishida et al., 2004; Kunimatsu et al., 2004; 1600 Nakatsukasa and Kunimatsu, 2009). This condition is more derived than that of hylobatids, 1601 which differ from both pongines and hominines in the lack of premaxillary-maxillary overlap 1602 (Ward and Kimbel, 1983; McCollum et al., 1983; McCollum and Ward, 1997). Except 1603 pongines, other Miocene apes also lack a clear overlap, although dryopithecines (and maybe 1604 afropithecines to a lesser extent) display a more derived condition than proconsulids and 1605 hylobatids, with the posterior pole of the premaxilla closely approaching or minimally 1606 overlapping the palatine process of the maxilla (Ward and Kimbel, 1983; Begun, 1994; Moyà- 1607 Solà and Köhler, 1995; McCollum and Ward, 1997; Brown et al., 2005; Moyà-Solà et al., 1608 2009a; Pérez de los Ríos et al., 2012; Pugh, 2022). Based on current fossil evidence, it is not 1609 possible to determine whether the divergent subnasal configuration of orangutans and 1610 African apes independently evolved from a dryopithecine-like condition, or whether gorillas 1611 retain the plesiomorphic configuration for crown hominids. This makes it difficult to 1612 interpret whether the derived condition of Nacholapithecus is homologous with that of 1613 crown hominids (Kunimatsu et al., 2019) or merely homoplastic (Pugh, 2022)—although the 1614 more plesiomorphic subnasal configuration of dryopithecines supports the latter view as 1615 long as they are interpreted as stem hominids less basal than equatorines. 1616 The potential stem hominid status of Nacholapithecus has been reinforced by the report 1617 that this taxon has an obliterated subarcuate fossa (Kunimatsu et al., 2019), which is 1618 similarly absent or very shallow in dryopithecines (Hispanopithecus and Rudapithecus; 1619 Moyà-Solà and Köhler, 1993, 1995; Kordos and Begun, 1997, 2001). However, as admitted 1620 by Kunimatsu et al. (2019), a well-developed subarcuate fossa is variably absent in 1621 cercopithecoids and hylobatids (Spoor and Leakey, 1996), indicating that its phylogenetic 1622 value must be interpreted with care, given its variability and the possibility that its 68 1623 obliteration could be homoplastic in various stem hominoids (Pugh, 2022). The latter 1624 contention is further reinforced by the lack of the subarcuate fossa in Oreopithecus (Harrison 1625 and Rook, 1997; Moyà-Solà and Köhler, 1997), currently most parsimoniously interpreted as 1626 a stem hominoid (see Section 4.3). 1627 Current morphological support for the stem hominid status of Nacholapithecus is thus 1628 somewhat ambiguous, although craniodental data support equatorines as stem hominids 1629 when postcranial features are ignored (Pugh, 2022) and this systematic position appears 1630 most likely under a basal divergence hypothesis for hylobatids (Fig. 8b). As remarked by 1631 Kunitmatsu et al. (2019), a hominid status for Nacholapithecus would provide additional 1632 support to the contention that not only suspensory behaviors (Moyà-Solà et al., 2004; Pugh, 1633 2022) but also an orthograde body plan (Alba, 2012) would have been independently 1634 acquired by hylobatids and hominids. Given that equatorines roughly coincide in age (~16– 1635 15 Ma) with crown hominid divergence average dates estimated from molecular data—16.5 1636 (19.7–13.5) Ma (Perelman et al., 2011) and 15.1 (20.8-11.0) Ma (Springer et al., 2012)— 1637 these neither favor nor preclude a crown hominid (e.g., hominine) status for 1638 Nacholapithecus. However, this possibility is not favored by most parimonious cladograms 1639 even if based on craniodental evidence alone (Pugh, 2022), and hence it is much more 1640 unlikely that a stem hominid status. 1641 Kenyapithecines and dryopithecines Unlike in the case of Nacholapithecus, the hominid 1642 status of Kenyapithecus is much better supported by cladistic analyses, albeit with some 1643 discordant results. Begun et al. (1997) recovered Kenyapithecus s.l. (including Equatorius) as 1644 a stem hominoid, while Begun et al. (2012) recovered Griphopithecus as a stem hominoid 1645 and Kenyapithecus as a stem hominid. More recently, Kenyapithecus was supported as a 1646 pongine (Nengo et al., 2017; Gilbert et al., 2020a) or as a stem hominid together with 69 1647 Griphopithecus (Pugh, 2022). The latter view had been favored by Alba (2012), in agreement 1648 with many previous authors (see Section 3.6) and the interpretation that the craniodental 1649 similarities between Kenyapithecus and pongines are symplesiomorphic (Harrison, 1992, 1650 2010a). 1651 Kenyapithecus stands out from a paleobiogeographic viewpoint because it is the only 1652 Miocene ape genus thus far recorded both in Africa and Eurasia, but a clear-cut 1653 interpretation of this fact is obscured by its uncertain phylogenetic relationships with 1654 dryopithecines and crown hominids (Almécija et al., 2021). Based on the record of a possible 1655 kenyapithecine in Europe before the Langhian transgression (Heizmann and Begun, 2001; 1656 Casanovas-Vilar et al., 2011), Gilbert et al. (2020b) suggested that apes might have followed 1657 a more westward dispersal route from Africa into Eurasia than the ancestors of 1658 pliopithecoids, subsequently dispersing into Asia sometime between 16 and 13 Ma. 1659 However, the larger samples of Griphopithecus and Kenyapithecus (Alpagut et al., 1990; 1660 Begun, 1992c, 2002a; Begun et al., 2003b; Kelley et al., 2008) are not recorded until 1661 somewhat later (~14.5–14.0 Ma) in Turkey (see discussion in Casanovas-Vilar et al., 2011 and 1662 Gilbert et al., 2020b, regarding the older dates favored for these sites by some other 1663 authors). It is generally assumed that Kenyapithecus dispersed from Africa into Eurasia 1664 (Andrews and Kelley, 2007; Alba, 2012) rather than the other way around, as an excessively 1665 literal reading of the fossil record might suggest (Begun, 2000). However, it is uncertain 1666 whether Eurasian kenyapithecines originated from a single dispersal event (as suggested by 1667 the close phylogenetic link between Kenyapithecus and Griphopithecus recovered by Pugh, 1668 2022) or two (pre- and post-Langhian) dispersals from Africa. 1669 Sometime after kenyapithecines are first recorded in Turkey, both pongines and 1670 dryopithecines are recorded in Asia and Europe, respectively, ~13.0–12.5 Ma. Not 70 1671 surprisingly, thus, transcontinental dispersal events followed by vicariant divergence have 1672 played a major role in the interpretation of the Eurasian hominoid radiation (Agustí et al., 1673 1996; Andrews and Bernor, 1999; Begun, 2005). However, the recognition that at least some 1674 species formerly included in Lufengpithecus lack pongine synapomorphies (Kelley and Gao, 1675 2012; Ji et al., 2013; Pugh, 2022) requires a more complicated paleobiogeograpic 1676 interpretation (Begun and Kelley, 2016), which is nevertheless obscured by phylogenetic 1677 uncertainties regarding dryopithecines and graecopithecins (see Section 3.7). Agustí et al. 1678 (1996) and Köhler et al. (2001), following Moyà-Solà and Köhler (1993, 1995), hypothesized 1679 that, after the hominoid dispersal into Eurasia, hominines evolved in Africa, with 1680 dryopithecines and graecopithecins being successive members of the pongine stem lineage 1681 less closely related to orangutans than Sivapithecus. This scenario is very reasonable from a 1682 paleobiogeographic perspective (Andrews and Bernor, 1999), but not supported by most 1683 parsimonious cladograms, which favor a stem hominid status for dryopithecines (Alba et al., 1684 2015; Pugh, 2022). In contrast, based on the hominine hypothesis for both dryopithecines 1685 and graecopithecins, Begun and coauthors initially discussed different paleobiogeograpic 1686 scenarios (Begun, 1994, 1995; Begun et al., 1997) but ultimately more actively promoted a 1687 Eurasian origin and early divergence of crown hominids, followed by a subsequent back-to- 1688 Africa dispersal of the latter in the Late Miocene (Begun, 2001, 2002a, 2005, 2009, 2010, 1689 2013, 2015, 2016; Begun et al., 2003a, 2012; Begun and Nargolwalla, 2004). 1690 Moyà-Solà et al. (2009b) advocated the view that dryopithecines are closely related to 1691 kenyapithecines and considered likely a Eurasian origin of crown hominids, but did not 1692 discount an independent evolution of hominines and pongines in Africa and Eurasia, 1693 respectively, from similar kenyapithecine ancestors—which is essentially the same 1694 hypothesis proposed by Agustí et al. (1996). Alba (2012), despite classifying dryopithecines 71 1695 as stem hominids, further commented on the possibility that pongines and hominines 1696 evolved separately in different continents from similar kenyapithecine ancestors. According 1697 to this hypothesis, dryopithecines would be more closely related to pongines—in which 1698 case, as noted by Alba (2012) and Almécija et al. (2021), orthogrady would have had to 1699 independently evolve not only between hylobatids and hominids, but also between 1700 hominines, pongines (as currently conceived), and dryopithecines. While the latter is 1701 plausible in the light of the postcranial evidence available for taxa such as Pierolapithecus 1702 and Sivapithecus (see Section 3.7), the cranial similarities noted by some previous authors 1703 between dryopithecines and pongines (Moyà-Solà and Köhler, 1993, 1995; Pérez de los Ríos 1704 et al., 2012) do not provide sufficient support to this hypothesis according to most recent 1705 cladistic analyses (Pugh, 2022) and support instead that even hispanopithecins (which 1706 appear less basal than kenyapithecines and dryopithecins) are stem hominids. The inclusion 1707 of Danuvius—chronologically intermediate between dryopithecins and hispanopithecins 1708 (Böhme et al., 2019)—in a formal cladistic analysis might help disentangle the phylogenetic 1709 relationships between dryopithecins and hispanopithecins, given that this genus also 1710 appears morphologically intermediate between them (Almécija et al., 2021). However, 1711 clarifying the phylogenetic relationships of dryopithecines would probably require as well 1712 more complete fossils around the Middle/Late Miocene from Europe (Alba et al., 2022) and 1713 China (Begun and Kelley, 2016). 1714 Although Pugh’s (2022) analyses generally support hispanopithecin monophyly, the 1715 support of H. laietanus as a stem pongine and of R. hungaricus as a stem hominine by her 1716 craniodental analyses parallels the previous cladistic results by Cameron (1997a), which led 1717 this author to resurrect Hispanopithecus. If supported by additional evidence, this topology 1718 would imply a Eurasian divergence of crown hominids—albeit not in the usual sense in 72 1719 which hispanopithecines are all considered hominines (e.g., Begun, 2005). Pugh’s (2022) 1720 support for the stem hominine status of graecopithecins is also consistent with a Eurasian 1721 divergence of crown hominids followed by a Late Miocene dispersal of hominines into Africa. 1722 However, as explained in Section 3.7 (see also SOM S4), we do not consider this 1723 phylogenetic link to be sufficiently supported at present. In any case, its paleobiogeographic 1724 implications would be less straightforward than the purported hominine status for some 1725 dryopithecines, given the stem hominine status further supported by Pugh (2022) for 1726 Nakalipithecus from the Late Miocene of Africa. 1727 Before the finds of Nakalipithecus and Chororapithecus (Kunimatsu et al., 2007; Suwa et 1728 al., 2007), the lack of unambiguous great apes during the late Middle Miocene and early Late 1729 Miocene of Africa was used to support the view that hominines originated in Europe (e.g., 1730 Begun, 2001)—even though, as pointed by Cote (2004), the absence of ape fossils was 1731 largely attributable to low sampling effort or inadequate habitat sampling. The existence of 1732 such a gap in the African hominid record is no longer tenable for the Late Miocene, although 1733 the support for the hominine status of Nakalipithecus and Chororapithecus is tenuous at 1734 best. Both the oldest graecopithecin (O. macedoniensis) and Nakalipithecus predate the 1735 average molecular dates between gorillins and other hominines—8.3 (10.1–6.6) Ma 1736 (Perelman et al., 2011) or 8.0 (11.7–5.5) Ma (Springer et al., 2012). This fact suggests that 1737 these taxa, unlike Chororapithecus, are unlikely to be crown hominines based on their 1738 chronology, although this remains a possibility given the uncertainty range of molecular 1739 divergence dates. 1740 While the phylogenetic relationships of Chororapithecus cannot be properly evaluated 1741 (see Section 3.7), the potential link between Nakalipithecus and graecopithecins, if 1742 confirmed, would be suggestive of an intercontinental connection during the Late Miocene. 73 1743 Nevertheless, the less derived morphology of Nakalipithecus (Kunimatsu et al., 2007), in 1744 agreement with its slightly older dating, might be interpreted to support instead an African 1745 origin and subsequent dispersal into Eurasia of graecopithecins. A European origin of 1746 graecopithecins from derived dryopithecines such as hispanopithecins would be consistent 1747 with their known fossil record in chronological terms. However, according to Pugh’s (2022) 1748 results—which recover hispanopithecins as advanced stem hominids and graecopithecines 1749 as stem hominines—this would imply that hispanopithecins gave rise to both pongines and 1750 hominines, which is at odds with the older (>2 Myr) record of pongines as compared with 1751 hispanopithecins. 1752 Other possibilities would be: (1) graecopithecins represent a dispersal event of stem 1753 hominines from Africa into Eurasia (also consistent with the most parsimonious results of 1754 Pugh, 2022); (2) graecopithecins are stem hominids evolved from hispanopithecins that 1755 represent an evolutionary dead-end (not supported by the results of Pugh, 2022); or (3) 1756 ouranopithecins and at least some hispanopithecins are stem hominines (only supported by 1757 the craniodental analysis of Pugh, 2022). Only the latter possibility necessarily entails a 1758 hominine dispersal back to Africa, whereas Pugh’s (2022) best supported alternative implies 1759 a dispersal event that might have been in either direction. Therefore, the place of origin of 1760 hominines seems difficult to determine until more complete Late Miocene hominids from 1761 Africa are discovered, although the various alternatives discussed above illustrate why 1762 clarifying the phylogenetic relationships of dryopithecines is also key for clarifying this 1763 question. 1764 1765 5. Discussion 1766 5.1. Something is rotten in Miocene ape phylogeny 74 1767 Despite the progress made during the last decades in terms of Miocene ape systematics, 1768 many phylogenetic and paleobiogeographic uncertainties persist. Some of them, such as the 1769 systematic status of equatorines and the potential paraphyly of dryopithecines, might seem 1770 of relatively minor relevance for understanding the big picture of hominoid evolution, even 1771 if they have potentially deeper implications for the origin of crown hominids. In contrast, the 1772 enduring uncertainties about origin of hylobatids appear of utmost importance, given the 1773 implications of its branching topology to determine the systematic (stem vs. crown 1774 hominoid) status of Early Miocene apes. 1775 The parsimony analyses published during the last decade support a less basal divergence 1776 of hylobatids that implies a stem hominoid status for all Early and Middle Miocene apes 1777 from Africa except Kenyapithecus. This is the view that—coupled with some uncertainties 1778 regarding some Eurasian taxa (such as oreopithecins and graecopithecins)—has been 1779 reflected it in the systematic classification used in this work. Nevertheless, several lines of 1780 evidence strongly suggest that such an advanced branching of hylobatids is probably amiss: 1781 (1) Molecular estimates for the hylobatid–hominid divergence predate by several million 1782 years the oldest record of stem hominids and hylobatids; (2) A separate analysis of 1783 craniodental and postcranial data yields strikingly different topologies for hylobatids—with 1784 postcranial data excluding most Miocene apes from the crown hominoid clade, and 1785 craniodental evidence supporting the crown hominoid status of equatorines; and (3) Bona 1786 fide Miocene great apes (especially Pierolapithecus and Sivapithecus) support the 1787 independent evolution of suspensory and orthogrady-related features along various crown 1788 hominoid lineages. 1789 Discrepancies between two subsets of data taken from a single taxon-character matrix 1790 are inevitable, as the resulting most parsimonious cladograms will be but sampling estimates 75 1791 of the true (parametric) phylogeny (Rodrigo et al., 1993). However, highly discrepant results 1792 may also highlight the presence of insufficient (or misleading) phylogenetic signal in one or 1793 all of the analyzed subsets. In general, combining different morphological datasets in a total 1794 evidence analysis is preferable, as this procedure may reveal hidden support that is not 1795 evident from each dataset separately (de Queiroz and Gatesy, 2007). Nevertheless, if one 1796 dataset is systematically affected by homoplasy in a particular given direction (as in the 1797 assumption of postcranial paralellism between hylobatids and hominids), then the results of 1798 the total evidence analysis might be less accurate than those yielded by the other dataset 1799 (i.e., craniodental data). This suggests that the relative branching order among hylobatids 1800 and putative stem hominoids from Africa other than proconsulids might be less reliable 1801 (potentially less accurate) than other phylogenetic relationships recovered by the analysis. 1802 Pugh’s (2022) results based on postcranial data evoke Pilbeam’s (1997) contention that 1803 most Miocene apes except Oreopithecus likely belong to an ‘archaic’ radiation not directly 1804 related to modern hominoids. Paradoxically, Oreopithecus is the taxon that most clearly 1805 evinces the contradictory signal provided by craniodental and postcranial data. The detailed 1806 phylogenetic affinities of Oreopithecus deserve further investigation but multiple lines of 1807 craniodental evidence (from the facial morphology to the inner ear anatomy and tooth 1808 endostructural shape) support that it is a late descendant of an ancient lineage more basal 1809 than crown hominoids. The incongruent signal displayed by Oreopithecus is the opposite of 1810 that of Sivapithecus but highlights the same problem. 1811 Pilbeam (1997) and Harrison and Rook (1997) argued that the traditional tendency to 1812 invoke postcranial rather than cranial homoplasy was a historical bias owing to the scarcity 1813 of hominoid postcranial remains. However, these views were soon challenged by Larson 1814 (1998), who advocated parallelism between lesser and great apes. Subsequent discoveries 76 1815 have largely supported Larson’s (1998) views that there are good reasons to mistrust the 1816 purported postcranial synapomorphies of crown hominoids (Moyà-Solà et al., 2004; Alba, 1817 2012; Ward, 2015): the fact that Pierolapithecus and Sivapithecus are recovered as a stem 1818 hominid and as a pongine, respectively, by most parsimonious cladograms (Pugh, 2022), 1819 despite lacking some of the derived postcranial features shared by crown hominids, supports 1820 the view that these features are homoplasies rather than true synapomorphies. Under this 1821 view, the hominoid postcranium would be prone to evolve the same features again and 1822 again as long as it is subject to similar positional selection pressures. The fact that cladistic 1823 analyses can resolve the Sivapithecus dilemma could be taken as an indication that 1824 postcranial homoplasy can be readily overcome by parsimony analysis. However, in reality 1825 we cannot know how often and to what extent the potentially misleading evidence provided 1826 by the postcranial remains may override the true phylogenetic signal. 1827 The problem of abundant homoplasy has long been recognized in paleoanthropology 1828 (e.g., Fleagle, 1997; Larson, 1998; Lockwood and Fleagle, 1999; Young, 2003; Begun, 2007). 1829 Although homoplasies may result from character misscoring, in most cases they reflect a real 1830 biological phenomenon resulting from evolutionary constraints and/or recurrent selective 1831 pressures during evolution (Felsenstein, 1978; Lockwood and Fleagle, 1999; Begun, 2007). 1832 The fact that, on epistemological grounds, homoplasies cannot be determined a priori but 1833 must be identified a posteriori based on most parsimonious cladograms (see SOM S1.3) does 1834 not mean that they can be discarded as uninteresting ad hoc hypotheses or phylogenetic 1835 ‘noise’—rather the contrary, they should be scrutinized on the basis of morphofunctional 1836 and developmental considerations (Fleagle, 1997; Lockwood and Fleagle, 1999; Begun, 1837 2007). Nevertheless, this is more easily said than done. In practice, there is no other option 1838 but to rely on most parsimonious cladograms based on all available evidence, even if it 77 1839 suspected (as in this case) that homoplasy is more prevalent in a particular anatomical area. 1840 This does not prevent the results from being potentially biased by differential degrees of 1841 homoplasy between the cranium and the postcranium because the result will be “heavily 1842 dependent on which morphological area can muster the greater number of characters” 1843 (Andrews and Pilbeam, 1996: 124). 1844 The fact that most cladistic studies show a large minimum level of homoplasy (Fleagle, 1845 1997; Begun, 2007) represents a caveat for retrieving phylogenetic signal because 1846 morphological characters, in practice, are potentially exhaustible (Felsenstein, 1978) so that 1847 data matrices can become saturated by excessive homoplasy as clades age (Wagner, 2000). 1848 This saturation erodes and can even override true phylogenetic signal, leading to a long- 1849 branch attraction problem (Wagner, 2000; Bergsten, 2005). The latter occurs when lineages 1850 that have undergone many changes artifactually appear too closely related (Bergsten, 2005), 1851 either because phylogenetic signal has been saturated or because the long branches 1852 converge on similar morphologies. Cladistic analyses are expected to be more prone to long- 1853 branch attraction than other methods of phylogenetic inference because parsimony does 1854 not take into account the lengths of the branches and hence tends to underestimate the 1855 amount of change in long branches (Pagel, 1999). 1856 To conclude that postcranial homoplasies are causing a long-branch attraction problem 1857 between hylobatids and hominids we would need independent evidence that most 1858 parsimonious cladograms are yielding inaccurate results (Brower, 2017). Of course, we do 1859 not have such evidence, but the highly contradictory results yielded by craniodental and 1860 postcranial datasets (Pugh, 2022) point to this direction. Hylobatids, hominids, and 1861 Oreopithecus are the longest branches of hominoid phylogeny from the viewpoint of their 1862 duration and they all display postcranial adaptations for antipronograde behaviors. 78 1863 Therefore, the conditions for a long-branch attraction problem are undoubtedly met, being 1864 further aggravated by the high number of missing data. According to this, parsimony 1865 analyses might potentially recover hylobatids and hominids as more closely related than 1866 they actually are (relative to Early and Middle Miocene hominoids) simply because they have 1867 had more time to accumulate morphological changes. 1868 1869 5.2. Future directions of work 1870 The possibility that Miocene ape most parsimonious cladograms are biased by a long- 1871 branch attraction problem that systematically recovers hylobatids as too closely related to 1872 hominids has far-reaching implications: even if Pilbeam’s (1996, 1997) views about the 1873 synapomorphic nature of extant hominoid postcranial similarities have been progressively 1874 abandoned during the last two decades, they would still exert a powerful—even if largely 1875 unconscious—influence simply because of the way cladistics works (see SOM S1.3) and the 1876 fragmentary nature of the ape fossil record. Abandoning postcranial characters in Miocene 1877 ape phylogenetic inference is not an option, both from an epistemological perspective and 1878 on more practical grounds. We agree with previous authors (e.g., Ward, 2015; Pugh, 2022) 1879 that the incongruence between craniodental and postcranial features does not imply that 1880 the latter are not useful in hominoid phylogenetics. So, how can we make progress? In the 1881 following, we outline future directions of research, with emphasis on the possibilities offered 1882 by recent developments in phylogenetic inference. 1883 More and better fossils and analyses From the viewpoint of fieldwork, finding additional 1884 remains of Miocene apes will always be most helpful, not only to increase taxon sampling 1885 but especially to reduce the proportion of missing data. Too many Miocene ape genera are 1886 still known mainly from fragmentary dentognathic remains, so that partial skeletons of 79 1887 Miocene apes with associated craniodental remains would be particularly welcome. This is 1888 not an easy task because fossil hominoids tend to be comparatively less abundant than 1889 many other mammalian taxa, and thus their finding generally requires a large sampling 1890 effort (Cote, 2004; Alba et al., 2017). 1891 More detailed morphological analyses could also increase the number of phylogenetically 1892 informative characters analyzed. These efforts should ideally be focused on craniodental 1893 features that embed strong phylogenetic signal (sensu Blomberg and Garland, 2002)—such 1894 as the semicircular canals of the inner ear (Urciuoli et al., 2020, 2021a, 2021b) or enamel– 1895 dentine shape (Zanolli et al., 2022b)—because a priori they appear less prone to (albeit not 1896 entirely devoid of) homoplasy than other features with a greater functional signal (such as 1897 postcranial remains; e.g., Arias-Martorell et al., 2021). Automated quantitative 1898 morphometric methods would further reduce the inherent subjectivity of cladistic analyses. 1899 One of the main merits of cladistics—besides the computer-assisted analysis of large 1900 amounts of data—is the obligation to be transparent about the data upon which most 1901 parsimonious cladograms are based, by elaborating taxon-character matrices that can be 1902 scrutinized by other scholars. Nevertheless, in morphology-based phylogenetic inference 1903 there is still a subjective component in the selection, definition, and scoring of the characters 1904 employed (Pilbeam and Young, 2001; Cartmill, 2018). This subjectivity is much greater than 1905 in molecular phylogenetics (Scotland et al., 2003), to the extent that the “often subjective 1906 nature of discrete character coding can generate discordant results that are rooted in 1907 individual researchers’ subjective interpretations” (Parins-Fukichi, 2017: 328). This makes 1908 obtaining different results unavoidable, largely depending on the researchers’ unconscious 1909 preconceptions, informed preferences, and anatomical expertise. The Ramapithecus debate 1910 illustrates “the power of preconceptions” as we as paleoanthropologists—like all scientists— 80 1911 are all “guided to some degree by a set of assumptions, usually implicit rather than explicit” 1912 (Lewin, 1987: 126). 1913 Given that morphology can be discretized ad infinitum, there is no objective solution 1914 regarding how many characters or character states a particular anatomical area requires, 1915 further resulting in the loss of potentially informative data. Therefore, it is always desirable 1916 to develop more objective ways to quantitatively analyze shape from a phylogenetic 1917 viewpoint. Additional efforts would be thus required to determine the most suitable 1918 methods to code continuous data (Goloboff et al., 2006; Pugh, 2022) as well as to more fully 1919 integrate 3D geometric morphometric data with cladistics (Almécija et al., 2021). This 1920 synthesis is already underway thanks to recent methodological developments based on 1921 landmarks (Catalano et al., 2010; Goloboff and Catalano, 2011, 2016). Nevertheless, 1922 landmark-free methods (e.g., Urciuoli et al., 2020) should ideally be employed, as landmark 1923 protocols may introduce some biases of their own. If bone morphology could be analyzed 1924 using parsimony by entirely relying on automatic coding methods based on geometric 1925 morphometric techniques, the subjectivity of character and character state definition would 1926 be drastically reduced. 1927 In the midterm, paleoproteomics (based on the retrieval of phylogenetically informative 1928 amino acid sequences from fossil remains) might also provide invaluable data for deciding 1929 among some of the most controversial hypotheses about Miocene ape phylogeny and 1930 paleobiogeography (Almécija et al., 2021). Thus far, paleoproteomic data have confirmed 1931 that Early Pleistocene (1.9 Ma) Gigantopithecus is a pongine distantly related to orangutans 1932 (Welker et al., 2019), with an estimated divergence date of 12–10 Ma, compatible with 1933 being more closely related to the sugrivapithecins Sivapithecus and Indopithecus. If 1934 technological advances eventually enable the retrieval of phylogenetically informative 81 1935 paleoproteomic data from key Miocene apes (such as Oreopithecus or dryopithecines), this 1936 might be enough to confidently anchor them in the phylogeny of extant species and use it as 1937 an extended molecular backbone in cladistic analyses—hopefully resulting in a more 1938 accurate phylogeny of Miocene apes as a whole. 1939 Beyond parsimony Besides the various aspects mentioned in the paragraphs above, it is 1940 worth emphasizing that further progress in Miocene ape phylogeny could potentially be 1941 made based on currently available paleontological data. The most obvious next step would 1942 be to increase taxon sampling and replicate Pugh’s (2022) joint and separate analyses of 1943 craniodental and postcranial characters—including taxa such as afropithecines, more 1944 proconsulids, putative stem hylobatids, and dendropithecids—to better assess the potential 1945 long-branch attraction problem hypothesized above. The application of character weighting 1946 methods (e.g., Goloboff, 1993, 1997) to Pugh (2022) and others’ cladistic matrices might 1947 potentially help cope with differential homoplasy between anatomical regions as well. 1948 Finally, and most importantly, Miocene ape phylogenetics could take advantage of recent 1949 methodological advances that have yet to be applied to these taxa. 1950 The inclusion of extinct taxa in morphology-based cladistic analysis has a very positive 1951 effect because they are closer in time to ancestral nodes and display combinations of 1952 primitive and derived features not found among extant taxa, thus helping determine the 1953 polarity of change and better discriminate between homoplasy and homology (Gauthier et 1954 al., 1988; Donoghue et al., 1989; Huelsenbeck, 1991; Smith, 1998; Smith and Turner, 2005; 1955 Hunt and Slater, 2016; Mongiardino Koch et al., 2021). Methods of phylogenetic inference 1956 other than morphology-based cladistics are not immune either to the problems of pervasive 1957 homoplasy, subjectivity in character selection and definition, and abundant missing data. 1958 However, parsimony arguably lags behind because of its inability to incorporate one of the 82 1959 main contributions of paleontology to evolutionary biology: deep time. To solve this 1960 problem, several decades ago some paleontologists advocated a modified concept of 1961 parsimony incorporating stratigraphic data a priori, as in stratocladistics, which maximizes 1962 explanatory power based on a modified parsimony criterion (Fisher, 2008; see SOM S1.4 for 1963 further details). Although this method was criticized by some other paleontologists, if 1964 stratigraphic data are considered relevant for inferring phylogeny there is no reason why the 1965 former cannot overturn parsimony considerations based exclusively on morphological data 1966 (Grantham, 2004), in agreement with the principle of total evidence frequently advocated in 1967 cladistics (SOM S1.4). 1968 Despite criticisms, stratocladistic analyses performed well (Clyde and Fisher, 1997) and 1969 simulation studies supported that stratocladistics outperforms the accuracy of conventional 1970 cladistics (Fox et al., 1999), being eventually applied to Miocene apes (Finarelli and Clyde, 1971 2004). But despite the development of a computer program to perform automated 1972 stratocladistic searches (Marcot and Fox, 2008), this approach never became mainstream. 1973 This might be attributable to the development in the 2000s of Bayesian methods of 1974 phylogenetic inference using Markov chain Monte Carlo techniques applicable to discrete 1975 morphological data (e.g., Nylander et al., 2004; see SOM S1.5 for further details). According 1976 to simulations, Bayesian analyses yield more accurate (Puttick et al., 2019)—although less 1977 resolved (O’Reilly et al., 2016)—cladograms than parsimony analyses, even when extinct 1978 taxa are incompletely preserved and there are high levels of homoplasy, being less sensitive 1979 to long-branch attraction (see additional references in SOM S1.5). Instead of incorporating a 1980 molecular backbone in morphology-based cladistic studies or deriving molecular estimates 1981 of divergence times based on bounds taken from the fossil record (‘node dating’), total- 1982 evidence Bayesian analyses based on morphologic, chronostratigraphic, and molecular data 83 1983 produce time-calibrated phylogenies (‘tip dating’; e.g., Ronquist et al., 2016; Zhang et al., 1984 2016; Pozzi and Penna, 2022). 1985 Despite various criticisms and limitations (e.g., O’Reilly et al., 2016; Goloboff, 2018; see 1986 discussion in Pugh, 2022 and SOM S1.5), simulation studies support that tip-dated 1987 phylogenies generally have a better fit with stratigraphic data and a greater accuracy than 1988 other methods (King, 2021; Mongiardino Koch et al., 2021). Therefore, it is somewhat 1989 surprising that tip-dating methods have yet to be applied in Miocene ape phylogenetics. 1990 Thus far, the application of these methods to primates is still limited (see review in Pozzi and 1991 Penna, 2022). This is probably because it takes time to introduce methodological advances in 1992 phylogenetic inference into paleoanthropology (see Section 2.2. regarding the slow 1993 introduction of cladistics)—which, as a discipline, has played a much more important role in 1994 the development of geometric morphometrics (Slice, 2007). 1995 Only recently, Pugh (2022) compared the results of parsimony and Bayesian methods for 1996 inferring the phylogeny of Miocene apes, obtaining a better resolution using the former, in 1997 agreement with simulation results (O’Reilly et al., 2016). However, Pugh (2022) did not take 1998 advantage of Bayesian tip-dating methods to obtain a time-calibrated phylogeny of Miocene 1999 apes. Although various aspects of Bayesian analyses are not sufficiently understood yet 2000 (Almécija et al., 2021; Pugh, 2022), total-evidence (tip-dating) methods are very appealing in 2001 the light of recent simulation results, which support that “fossils help to extract true 2002 phylogenetic signals from morphology” not only because of “their distinctive morphology” 2003 but also because of “their temporal information” (Mongiardino Koch et al., 2021: 1). 2004 Therefore, performing a tip-dated Bayesian total evidence analysis and comparing the 2005 results with those of parsimony with implied character weighting should be a must for 2006 Miocene ape phylogenetics in the midterm. 84 2007 2008 6. Summary and conclusions 2009 Miocene apes are much more diverse than their extant counterparts, evincing a suite of 2010 mosaic morphologies that are essential to reconstruct the evolutionary history of the 2011 Hominoidea. Here we review Miocene ape evolution with emphasis on their phylogenetic 2012 relationships and the paleobiogeographic scenarios that derive from them. The oldest 2013 hominoids from the Oligocene, Miocene catarrhines of uncertain affinities, and Late 2014 Miocene purported hominins are excluded from this review. 2015 First, we provide a historical account of the progress made in hominoid phylogeny and 2016 paleobiogeography during the last one hundred and fifty years, with emphasis on the 2017 Ramapithecus debate, the molecular revolution, the spread of the cladistic paradigm, the 2018 Sivapithecus dilemma, the enigmatic ape Oreopithecus, and the synapomorphic vs. 2019 homoplastic nature of the postcranial similarities shared by extant apes. Second, based on 2020 our interpretation of the most parsimonious results yielded by recent cladistic analyses, we 2021 report an updated classification of Miocene apes. Our classification tentatively distinguishes 2022 three families of putative stem Hominoidea (Proconsulidae, Afropithecidae, and 2023 Nyanzapithecidae), a new subfamily Yuanmoupithecinae for stem Hylobatidae, and two 2024 subfamilies of stem Hominidae (Kenyapithecinae and Dryopithecinae), while the tribes 2025 Oreopithecini and Graecopithecini are provisionally left as subfamily incertae sedis. Third, 2026 we report a list of Miocene ape species-locality occurrences accompanied by an analysis of 2027 their paleobiodiversity dynamics and a discussion of the highly contradictory results yielded 2028 by parsimony analyses based on craniodental and postcranial features. On the basis of the 2029 latter, we argue that the less basal divergence of hylobatids relative to putative stem 2030 hominoids, as currently favored by most parsimonious cladograms, is far from being 85 2031 definitively settled and that a more basal divergence is likely given the abundant postcranial 2032 homoplasy between hylobatids and hominids. With these two competing hypotheses (more 2033 vs. less basal divergence for hylobatids) in mind, we discuss ongoing debates about the 2034 origin of hylobatids, Oreopithecus, and hominoids from a phylogenetic and 2035 paleobiogeographic perspective. 2036 Hylobatid origins remain uncertain because their branching topology relative to 2037 nyanzapithecids and afropithecids is unreliable, given the contrasting phylogenetic signal 2038 provided by craniodental and postcranial features. However, if the suspicion that recent 2039 cladistic analyses are biased toward a too advanced divergence of hylobatids were correct, 2040 an initial divergence of crown hominoids in Africa followed by two independent dispersal 2041 events into Eurasia would be most likely. It remains to be more conclusively determined 2042 whether equatorines are stem hominids rather than stem hominonids, as favored by most 2043 recent cladistic analysis, whereas kenyapithecines appear as the most likely stock from 2044 which crown hominids might have evolved. In contrast, current data favor the view that 2045 Oreopithecus is a stem hominoid rather than a stem hominid or hylobatid. The geographic 2046 origin of crown hominids remains unresolved due to phylogenetic uncertainties regarding 2047 dryopithecines and graecopithecins. The possibility that a dispersal event from 2048 kenyapithecines into Eurasia set the initial divergence between hominines and pongines 2049 remains a plausible interpretation but lacks cladistic support (unless European 2050 dryopithecines originated from a second dispersal event of stem hominids from Africa). 2051 Alternatively, the pongine–hominine divergence might have occurred in Eurasia, in 2052 agreement with the view that graecopithecins and at least some dryopithecines are 2053 hominines, which we consider insufficiently supported at present. 86 2054 The less basal divergence of hylobatids favored by most parsimonious cladograms might 2055 artifactually result from a long-branch attraction problem caused by the numerous 2056 postcranial similarities shared by extant hylobatids and hominids (which appear largely 2057 homoplastic), coupled with abundant missing data from Miocene apes. Although is it 2058 currently recognized that postcranial features functionally related to antipronograde 2059 behaviors likely evolved in parallel along various ape lineages, they still exert a strong 2060 influence in the outcome of parsimony analyses because homoplasy can only be identified a 2061 posteriori and branch lengths are not considered. Although it is not possible to determine to 2062 what extent this potential problem might affect the accuracy of most parsimonious 2063 cladograms, it could be ameliorated by the discovery of additional Miocene ape fossils 2064 (leading to increased taxon sampling and/or a decrease of missing data), the implementation 2065 of character weighting methods, the scoring of new characters from anatomical areas 2066 embedding high phylogenetic signal, and the integration of geometric morphometric 2067 continuous data in cladistic analyses. Paleoproteomic analyses offer even more promising 2068 prospects to more reliably reconstruct the evolutionary history of hominoids if future 2069 technological advances allow the retrieval of molecular data from Miocene apes of 2070 controversial systematic position. 2071 Finally, we further advocate incorporating chronostratigraphic information in 2072 phylogenetic inference by taking advantage of recent methodological advances. In 2073 particular, total-evidence (tip-dating) Bayesian methods of phylogenetic inference appear to 2074 outperform parsimony methods and provide time-calibrated phylogenies based on the 2075 simultaneous analysis of molecular, morphologic, and chronostratigraphic data. Performing 2076 these analyses and comparing their results with more conventional parsimony analyses 2077 would hopefully allow to ascertain if, as we suspect, our current concept of the hominoid 87 2078 stem lineage is artifactually inflated by a long-branch attraction problem between hylobatids 2079 and hominids. The data required to perform such analyses are already out there. 2080 2081 Acknowledgments 2082 This paper is part of R+D+I projects PID2020-116908GB-I00 and PID2020-117289GBI00, 2083 funded by the Agencia Estatal de Investigación of the Spanish Ministerio de Ciencia e 2084 Innovación (MCIN/AEI/10.13039/501100011033/). Our research has also been funded by 2085 CERCA Programme/Generalitat de Catalunya and a Margarita Salas postdoctoral fellowship 2086 funded by the European Union-NextGenerationEU to A.U. D.M.A. is member of the 2087 consolidated research group 2017 SGR 116 GR of the Agència de Gestió d’Ajuts Universitaris 2088 i de Recerca (Generalitat de Catalunya). This contribution is part of the special issue 'The 2089 50th Anniversary of Journal of Human Evolution: Current and Future Directions in 2090 Reconstructing Our Past'; we are indebted to the Editors of this special issue (Andrea Taylor 2091 and Clément Zanolli) for inviting us to contribute. We also thank Salvador Moyà-Solà and 2092 Sergio Almécija for plentiful discussions about fossil apes over the years; John Barry, Larry 2093 Flynn, Jay Kelley, Michèle Morgan, and David Pilbeam for allowing us to use their data about 2094 Sivapithecus-bearing localities from the Siwaliks; John Barry, Larry Flynn, Martin Pickford, 2095 David Pilbeam, and Lorenzo Rook for providing literature; Florian Bouchet for assistance 2096 regarding literature search; Terry Harrison for granting us access to Ji et al.'s (2022) 2097 manuscript while still in press; and the co-Editor-in-Chief (Clément Zanolli) and the three 2098 reviewers (Jay Kelley and two anonymous ones) for providing comments that helped us 2099 improve a previous draft of this paper. 2100 2101 References 88 2102 Abel, O., 1902. Zwei neue Menshenaffen aus den Leithakalkbildungen des Wiener Beckens. 2103 Sitz.-Ber. K. Akad. Wiss. Wien math.-naturwiss. 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An apomorphy-based 3469 definition would imply that the hominoid last common ancestor would depend on the crown 3470 hominoid synapomorphy selected as definitory of the group (character state 1 in the 3471 example of the figure). 3472 3473 Figure 2. Schematic cladograms summarizing the phylogenetic relationships of the hominoid 3474 families distinguished here relative to cercopithecoids and the most advanced stem 3475 catarrhines (pliopithecoids and dendropithecids) according to the contrasting cladistic 3476 results of various authors: a) based on Nengo et al. (2017) and Gilbert et al. (2020a); b) 3477 based on Rossie and Hill (2018). The hominoid stem lineage is denoted in light gray whereas 3478 the hominoid crown group is denoted in dark gray. Note that, according to Rossie and Hill 3479 (2018), Dendropithecidae s.s. (Dendropithecus and Simiolus) would be stem hominoids but 147 3480 Dendropithecidae s.l. (including Micropithecus) would be polyphyletic. Abbreviation: LCA = 3481 last common ancestor. 3482 3483 Figure 3. Geographic distribution of Early Miocene apes. The information reported mostly 3484 comes from Table 2 but further includes the following species-locality occurrences (see SOM 3485 Table S1 for further details and SOM Table S2 for references): Hominoidea indet. from 3486 Moruorot (Kenya; 17.5 Ma); Nyanzapithecidae indet. from Ryskop (South Africa; 16.0 Ma); 3487 and cf. Kenyapithecinae indet. from Engelswies (Germany; 16.5–16.0 Ma). Base map 3488 downloaded from ArcGIS Online (https://www.esri.com/it-it/arcgis/products/arcgis-online/); 3489 image sources: Esri, Maxar, GeoEye, Earthstar Geographics, CNES/Airbus DS, USDA, USGS, 3490 AeroGRID, IGN, and the GIS User Community, Sources: Esri, Airbus DS, USGS, NGA, NASA, 3491 CGIAR, N Robinson, NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, 3492 FEMA, Intermap and the GIS user community. 3493 3494 Figure 4. Geographic distribution of Middle Miocene apes. The information reported mostly 3495 comes from Table 2 but further includes the following species-locality occurrences (see SOM 3496 Table S1 for further details and SOM Table S2 for references): Nyanzapithecidae indet. from 3497 Fort Ternan and Kapsibor (Kenya; 13.7 Ma); Kenyapithecinae indet. from Thannhausen 3498 (Germany; 14.0 Ma); cf. Kenyapithecus sp. from Berg Aukas (Namibia; 13.0–12.0 Ma); 3499 Hominidae indet. from Ngorora (Kenya; 12.8–12.0 Ma). Base map downloaded from ArcGIS 3500 Online (https://www.esri.com/it-it/arcgis/products/arcgis-online/); image sources: Esri, 3501 Maxar, GeoEye, Earthstar Geographics, CNES/Airbus DS, USDA, USGS, AeroGRID, IGN, and 3502 the GIS User Community, Sources: Esri, Airbus DS, USGS, NGA, NASA, CGIAR, N Robinson, 148 3503 NCEAS, NLS, OS, NMA, Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, Intermap 3504 and the GIS user community. 3505 3506 Figure 5. Geographic distribution of Late Miocene apes. The information reported mostly 3507 comes from Table 2 but further includes the following species-locality occurrences (see SOM 3508 Table S1 for further details and SOM Table S2 for references): Hominoidea indet. from N 885 3509 (Niger; 11.0–5.0 Ma); Nyanzapithecinae indet. from Nakali (Kenya; 9.9–9.8 Ma); Hominidae 3510 indet. from Maragheh (Iran; 7.5 Ma); Dryopithecinae indet. from Wissberg (Germany; 13.7– 3511 7.5 Ma), Neuhausen, Egingen, and Trochtelfingen (Germany; 11.0–9.0 Ma), Melchingen 3512 (Germany; 11.2–7.5 Ma), and Salmendingen (Germany; 11.6–7.5 Ma); Graecopithecini indet. 3513 from Azmaka (Bulgaria; 7.2 Ma); and Hominidae indet. from Kapsomin and Cheboit, Lukeino 3514 (Kenya; 6.2–5.7 Ma). Base map downloaded from ArcGIS Online (https://www.esri.com/it- 3515 it/arcgis/products/arcgis-online/); image sources: Esri, Maxar, GeoEye, Earthstar 3516 Geographics, CNES/Airbus DS, USDA, USGS, AeroGRID, IGN, and the GIS User Community, 3517 Sources: Esri, Airbus DS, USGS, NGA, NASA, CGIAR, N Robinson, NCEAS, NLS, OS, NMA, 3518 Geodatastyrelsen, Rijkswaterstaat, GSA, Geoland, FEMA, Intermap and the GIS user 3519 community. 3520 3521 Figure 6. Chronostratigraphic ranges and broad geographic distribution of Miocene apes 3522 summarized at the genus level and organized according to the classification reported in 3523 Table 1; family names are colored as in Figures 3–5. Chronostratigraphic ranges are 3524 organized from oldest to youngest (left to right) for each (sub)family and colored based on 3525 geographic distribution at the continental level (gray denotes geographic uncertainty due to 3526 lack of record for taxa represented in more than a single continent). The depicted genus 149 3527 ranges are reported in Table 2. Also depicted are the ranges for some taxa not identified to 3528 the genus rank (see SOM Table S1 for further details and SOM Table S2 for references): cf. 3529 Kenyapithecinae from Engelswies (Germany; 16.5–16.0); and Nyanzapithecidae indet. from 3530 Fort Ternan, Kapsibor (Kenya; 13.7 Ma) and Nakali (Kenya; 9.9–9.8 Ma). 3531 3532 Figure 7. Paleobiodiversity curve of Miocene apes through time: range-through or total 3533 diversity (top) and estimated standing diversity (bottom). Diversity metrics are based on the 3534 data reported in Table 3, see SOM Table S3 for further details. 3535 3536 Figure 8. Alternative phylogenetic hypotheses discussed in this paper as depicted by 3537 schematic time-calibrated cladograms at the (sub)tribe rank: a) phylogenetic hypotheses 3538 favored in this paper based on our interpretation of the current state of knowledge 3539 according to most parsimonious cladograms; b) alternative hypothesis of a more basal 3540 divergence for hylobatids coupled with multiple polytomies to highlight current phylogenetic 3541 uncertainties. Colored rectangles represent the chronostratigraphic ranges of the depicted 3542 taxa (color-coded as in Figs. 3–6 based on geographical distribution; gray denotes geographic 3543 uncertainty due to lack of record for taxa represented in more than a single continent); 3544 colored dots at the tips of extant lineages denote their current geographic distribution. Gray 3545 semitransparent rectangles represent the maximum-minimum divergence age estimates for 3546 crown clades. Internal nodes have been depicted arbitrarily 0.5 Myr before the oldest record 3547 of the group or relative to the oldest node that immediately follows, with the exception of 3548 crown groups, for which average estimated divergence times (Perelman et al., 2011) have 3549 been used (except when they are too close or even younger than the oldest record of the 150 3550 group). Note that no Plio-Pleistocene ranges are depicted, whereas in contrast the Oligocene 3551 range of proconsulids and nyanzapithecines has been depicted. 3552 151 = extant clades 0 † CATARRHINI † = hypothetical extinct clades stem lineage ? = character states 0 † 2 CERCOPITHECOIDEA † HOMINOIDEA stem lineage 0 0 † HOMINOIDEA total group 0 † 0 † 1 † 1 Hylobatidae HOMINOIDEA crown group 1 Hominidae Last common ancestors: Definitions of the Hominoidea: Crown Hominoidea Apomorphy-based Hominoidea (total group) Crown-based Crown Catarrhini Stem-based a PLIOPITHECOIDEA† b PLIOPITHECOIDEA† Micropithecus† Dendropithecidae s.l.†* CERCOPITHECOIDEA CERCOPITHECOIDEA Dendropithecidae s.s.†* Proconsulidae† Nyanzapithecidae† Afropithecidae†* Proconsulidae†* Nyanzapithecidae† Afropithecidae† Hylobatidae Hylobatidae Hominidae Hominidae † Extinct taxa * Paraphyletic taxa Hominoidea (total group) LCA Crown Hominoidea LCA Proconsulidae Nyanzapithecidae Afropithecinae Equatorinae Afropithecidae indet. Hylobatidae Kenyapithecinae Dryopithecinae Ponginae Homininae Oreopithecini Graecopithecini Hominidae indet./incertae sedis Hominoidea indet./incertae sedis Proconsulidae Nyanzapithecidae Afropithecinae Equatorinae Afropithecidae indet. Hylobatidae Kenyapithecinae Dryopithecinae Ponginae Homininae Oreopithecini Graecopithecini Hominidae indet./incertae sedis Hominoidea indet./incertae sedis Proconsulidae Nyanzapithecidae Afropithecinae Equatorinae Afropithecidae indet. Hylobatidae Kenyapithecinae Dryopithecinae Ponginae Homininae Oreopithecini Graecopithecini Hominidae indet./incertae sedis Hominoidea indet./incertae sedis 20 15 10 5 Ma EARLY MIOCENE MIDDLE MIOCENE LATE MIOCENE Proconsul Kalepithecus Proconsulidae Ekembo Turkanapithecus Xenopithecus Rangwapithecus Nyanzapithecus Mabokopithecus Nyanzapithecidae Nyazapithecidae indet. Samburupithecus Morotopithecus Afropithecus Heliopithecus Equatorius Afropithecidae Nacholapithecus Otavipithecus ‘Dryopithecus’ wuduensis Oreopithecus Incertae sedis Yuanmoupithecus Hylobatidae cf. Kenyapithecinae Kenyapithecus Griphopithecus Dryopithecus Anoiapithecus Africa Pierolapithecus Danuvius Hispanopithecus Rudapithecus ?Udabnopithecus Sivapithecus Asia Khoratpithecus Hominidae Ankarapithecus Indopithecus Lufengpithecus Nakalipithecus Chororapithecus Europe Ouranopithecus Graecopithecus Sinopithecus 14 12 Range-through or total diversity 10 8 6 4 2 0 23-22 22-21 21-20 20-19 19-18 18-17 17-16 16-15 15-14 14-13 13-12 12-11 11-10 10-9 9-8 8-7 7-6 6-5 Stratigraphic bins (1 Myr) 6 5 4 Standing diversity 3 2 1 0 23-22 22-21 21-20 20-19 19-18 18-17 17-16 16-15 15-14 14-13 13-12 12-11 11-10 10-9 9-8 8-7 7-6 6-5 Stratigraphic bins (1 Myr) a Proconsulidae Nyanzapithecidae Oreopithecini Afropithecidae Afropithecinae Equatorinae Hyloba�dae Yuanmoupithecinae Hyloba�nae Kenyapithecinae Ponginae Dryopithecinae Dryopithecini Crown Hominoidea Hispanopithecini Hominidae Graecopithecini Lufengpithecini Sugrivapithecini Pongini Stem Homininae? Homininae Gorillini Panini Hominini EARLY MIOCENE MIDDLE MIOCENE LATE MIOCENE 25 20 15 10 5 0 Ma b Proconsulidae Nyanzapithecidae Oreopithecini Hyloba�dae Yuanmoupithecinae Hyloba�nae Afropithecidae Afropithecinae Equatorinae Kenyapithecinae Ponginae Dryopithecinae Dryopithecini Crown Hominoidea Hispanopithecini Hominidae Graecopithecini Lufengpithecini Sugrivapithecini Pongini Stem Homininae? Gorillini Homininae Panini Hominini EARLY MIOCENE MIDDLE MIOCENE LATE MIOCENE 25 20 15 10 5 0 Ma 1 Table 1 2 A summary of the systematic classification of the Hominoidea followed in this paper down to 3 tribe rank. A dagger denotes extinct taxa. See Table 2 for taxonomic authorities and a 4 classification of Miocene ape genera and species. Family Subfamily Tribe Proconsulidae† — — Nyanzapithecidae† — — Afropithecidae† Afropithecinae† — Equatorinae† — Hylobatidae Yuanmoupithecinae† — Hylobatinae — Hominidae Kenyapithecinae† — Dryopithecinae† Dryopithecini† Hispanopithecini† Ponginae Pongini Lufengpithecini† Sugrivapithecini† Homininae Gorillini Panini Hominini incertae sedis Graecopithecini† incertae sedis incertae sedis Oreopithecini† 5 6 Table 2 7 Systematics of Miocene apes including taxonomic rank, taxon name, and taxonomic authoritya. A dagger denotes extinct taxa. Small-bodied 8 catarrhines of uncertain systematic status (such as dendropithecids) are not included, whereas two species variously interpreted as hominoids 9 or pliopithecoids depending on the authors are included as incertae sedis at superfamily rank. 10 Order Primates Linnaeus, 1758 11 Semiorder Euprimates Hoffstetter, 1977 12 Suborder Haplorrhini Pocock, 1918 13 Infraorder Anthropoidea Mivart, 1864 14 Parvorder Catarrhini É. Geoffroy Saint-Hilaire, 1812 15 Superfamily Hominoidea Gray, 1825 16 Family Proconsulidae Leakey, 1963† 17 Genus Proconsul Hopwood, 1933† 18 Proconsul africanus Hopwood, 1933† 19 Proconsul major Le Gros Clark and Leakey, 1950† 20 Proconsul gitongai (Pickford and Kunimatsu, 2005)† 21 Proconsul meswae Harrison and Andrews, 2009† 22 Proconsul legetetensis (Pickford et al., 2009a)† 23 Genus Kalepithecus Harrison, 1988† 24 Kalepithecus songhorensis (Andrews, 1978)† 25 Kalepithecus kogolensis Pickford et al., 2017† 26 Genus Ekembo McNulty et al., 2015† 27 Ekembo heseloni (Walker et al., 1993)† 28 Ekembo nyanzae (Le Gros Clark and Leakey, 1950)† 29 Family Afropithecidae Andrews, 1992† 30 Subfamily Afropithecinae Andrews, 1992† 31 Genus Afropithecus Leakey and Leakey, 1986a† 32 Afropithecus turkanensis Leakey and Leakey, 1986a† 33 Genus Heliopithecus Andrews and Martin, 1987b† 34 Heliopithecus leakeyi Andrews and Martin, 1987b† 35 Genus Morotopithecus Gebo et al., 1997† 36 Morotopithecus bishopi Gebo et al., 1997† 37 Subfamily Equatorinae Cameron, 2004† 38 Genus Equatorius Ward et al., 1999† 39 Equatorius africanus (Le Gros Clark and Leakey, 1950)† 40 Genus Nacholapithecus Ishida et al., 1999† 41 Nacholapithecus kerioi Ishida et al., 1999† 42 Subfamily incertae sedis 43 Genus Otavipithecus Conroy et al., 1992† 44 Otavipithecus namibiensis Conroy et al., 1992† 45 Family Nyanzapithecidae Harrison, 2002† 46 Genus Xenopithecus Hopwood, 1933† 47 Xenopithecus koruensis Hopwood, 1933† 48 Genus Mabokopithecus von Koenigswald, 1969† 49 Mabokopithecus clarki von Koenigswald, 1969† 50 Genus Rangwapithecus Andrews, 1974† 51 Rangwapithecus gordoni (Andrews, 1974)† 52 Genus Nyanzapithecus Harrison, 1986† 53 Nyanzapithecus vancouveringorum (Andrews, 1974)† 54 Nyanzapithecus pickfordi Harrison, 1986† 55 Nyanzapithecus harrisoni Kunimatsu, 1997† 56 Nyanzapithecus alesi Nengo et al., 2017† 57 Genus Turkanapithecus Leakey and Leakey, 1986b† 58 Turkanapithecus kalakolensis Leakey and Leakey, 1986b† 59 Turkanapithecus rusingensis Pickford, 2010† 60 Genus Samburupithecus Ishida and Pickford, 1997† 61 Samburupithecus kiptalami Ishida and Pickford, 1997† 62 Family Hylobatidae Gray, 1870 63 Subfamily Yuanmoupithecinae subfam. nov.† 64 Genus Yuanmoupithecus Pan, 2006† 65 Yuanmoupithecus xiaoyuan Pan, 2006† 66 Family Hominidae Gray, 1825 67 Subfamily Kenyapithecinae Andrews, 1992† 68 Genus Griphopithecus Abel, 1902† 69 Griphopithecus suessi Abel 1902† 70 Griphopithecus alpani (Tekkaya, 1974)† 71 Genus Kenyapithecus Leakey, 1961† 72 Kenyapithecus wickeri Leakey, 1961† 73 Kenyapithecus kizili Kelley et al., 2008† 74 Subfamily Dryopithecinae Gregory and Hellman, 1939† 75 Tribe Dryopithecini Gregory and Hellman, 1939† 76 Genus Dryopithecus Lartet, 1856† 77 Dryopithecus fontani Lartet, 1856† 78 Genus Pierolapithecus Moyà-Solà et al., 2004† 79 Pierolapithecus catalaunicus Moyà-Solà et al., 2004† 80 Genus Anoiapithecus Moyà-Solà et al., 2009b† 81 Anoiapithecus brevirostris Moyà-Solà et al., 2009b† 82 Genus incertae sedis 83 ‘Sivapithecus’ occidentalis Villalta Comella and Crusafont Pairó, 1944 species inquirenda† 84 Tribe Hispanopithecini Cameron, 1997a† 85 Genus Hispanopithecus Villalta Comella and Crusafont Pairó, 1944† 86 Hispanopithecus laietanus Villalta Comella and Crusafont Pairó, 1944† 87 Hispanopithecus crusafonti (Begun, 1992b)† 88 Genus Rudapithecus Kretzoi, 1969† 89 Rudapithecus hungaricus Kretzoi, 1969† 90 Genus Danuvius Böhme et al., 2019† 91 Danuvius guggenmosi Böhme et al., 2019† 92 Tribe incertae sedis 93 Genus Neopithecus Abel, 1902 nomen dubium† 94 Neopithecus brancoi (Schlosser, 1901) nomen dubium† 95 Genus ?Udabnopithecus Burchak-Abramovich and Gabashvili, 1945† 96 ?Ubadnopithecus garedziensis Burchak-Abramovich and Gabashvili, 1945 97 Subfamily Ponginae Elliot, 1913 98 Tribe Pongini Elliot, 1913 99 Genus Khoratpithecus Chaimanee et al., 2004† 100 Khoratpithecus chiangmuanensis (Chaimanee et al., 2003)† 101 Khoratpithecus piriyai Chaimanee et al., 2004† 102 Khorapithecus ayeyarwadyensis Jaeger et al., 2011† 103 Khoratpithecus magnus Chaimanee et al., 2022† 104 Tribe Sugrivapithecini Simonetta, 1958† 105 Genus Sivapithecus Pilgrim, 1910† 106 Sivapithecus sivalensis (Lydekker, 1879)† 107 Sivapithecus indicus Pilgrim, 1910† 108 Sivapithecus parvada Kelley, 1988† 109 Genus Indopithecus von Koenigswald, 1949† 110 Indopithecus giganteus (Pilgrim, 1915)† 111 Tribe Lufengpithecini Alba, 2012† 112 Genus Ankarapithecus Ozansoy, 1957† 113 Ankarapithecus meteai Ozansoy, 1957† 114 Genus Lufengpithecus Wu, 1987† 115 Lufengpithecus lufengensis (Xu et al., 1978)† 116 Subfamily Homininae Gray, 1825 117 Tribe incertae sedis 118 Genus Chororapithecus Suwa et al., 2007† 119 Chororapithecus abyssinicus Suwa et al., 2007† 120 Genus Nakalipithecus Kunimatsu et al., 2007† 121 Nakalipithecus nakayamai Kunimatsu et al., 2007† 122 Subfamily incertae sedis 123 Tribe Graecopithecini Cameron, 1997b† 124 Genus Graecopithecus von Koenigswald, 1972† 125 Graecopithecus freybergi von Koenigswald, 1972† 126 Genus Ouranopithecus de Bonis and Melentis, 1977† 127 Ouranopithecus macedoniensis (de Bonis et al., 1974)† 128 Ouranopithecus turkae Güleç et al., 2007† 129 Tribe incertae sedis 130 Genus Sinopithecus Zhang et al., 1990 131 Sinopithecus keiyuanensis (Woo, 1957)† 132 Sinopithecus hudienensis (Zhang et al., 1987)† 133 Family incertae sedis 134 Subfamily incertae sedis 135 Tribe Oreopithecini Schwalbe, 1915† 136 Genus Oreopithecus Gervais, 1872† 137 Oreopithecus bambolii Gervais, 1872† 138 Tribe incertae sedis 139 Genus incertae sedis 140 ‘Dryopithecus’ wuduensis Xue and Delson, 1988† a 141 Taxonomic authority consists of the author(s) that originally described a given taxon followed by year of publication. Following the 142 requirements of the Code (ICZN, 1999), taxonomic authorities are placed within parentheses only when a species was originally described 143 within a different genus than the one in which it is included in this work. 144 Table 3 145 Chronostratigraphic range and geographic distribution of Miocene apes, synthesized from species-locality occurrences reported in SOM Table 146 S2. The details for each locality are provided in SOM Table S1. Species of doubtful taxonomic validity and citations indeterminate to the species 147 rank are excluded from this table; in contrast, tentative attributions to species (with ‘cf.’) are included. Species Family Subfamily Age Subepoch Distribution Proconsul africanus Proconsulidae — 20.3–18.5 Early Miocene Kenya, Uganda Proconsul major Proconsulidae — 20.5–19.0 Early Miocene Kenya, Uganda Proconsul gitongai Proconsulidae — 15.8a Early to Middle Miocene Kenya, Uganda Proconsul meswae Proconsulidae — 22.5 Early Miocene Kenya Proconsul legetetensis Proconsulidae — 20.5–19.0 Early Miocene Uganda Kalepithecus songhorensis Proconsulidae — 20.5–19.0 Early Miocene Kenya, Uganda Kalepithecus kogolensis Proconsulidae — 21.0–20.0 Early to Middle Miocene Uganda Ekembo heseloni Proconsulidae — 18.5–16.0 Early to Middle Miocene Kenya Ekembo nyanzae Proconsulidae — 18.5–16.0 Early to Middle Miocene Kenya Xenopithecus koruensis Nyanzapithecidae — 20.0–19.0 Early Miocene Kenya Mabokopithecus clarki Nyanzapithecidae — 15.3 Middle Miocene Kenya Rangwapithecus gordoni Nyanzapithecidae — 20.0–17.0 Early Miocene Kenya Nyanzapithecus pickfordi Nyanzapithecidae — 15.8–15.3 Middle Miocene Kenya Nyanzapithecus vancouveringorum Nyanzapithecidae — 18.3 Early Miocene Kenya Nyanzapithecus alesi Nyanzapithecidae — 13.3 Middle Miocene Kenya Nyanzapithecus harrisoni Nyanzapithecidae — 15.0 Middle Miocene Kenya Turkanapithecus kalakolensis Nyanzapithecidae — 17.5–17.0 Early Miocene Kenya, Ethiopia Turkanapithecus rusingensis Nyanzapithecidae — 20.5–18.3 Early Miocene Kenya, Uganda Samburupithecus kiptalami Nyanzapithecidae — 8.5 Late Miocene Kenya Afropithecus turkanensis Afropithecidae Afropithecinae 17.6–17.2a Early to Middle Miocene Kenya, Uganda Heliopithecus leakeyi Afropithecidae Afropithecinae 16.0 Early Miocene Saudi Arabia Morotopithecus bishopi Afropithecidae Afropithecinae 21.0–20.0b Early Miocene Uganda Equatorius africanus Afropithecidae Equatorinae 16.0–15.0 Middle Miocene Kenya Nacholapithecus kerioi Afropithecidae Equatorinae 15.0a Early to Middle Miocene Kenya, Uganda Otavipithecus namibiensis Afropithecidae Incertae sedis 12.0 Middle Miocene Namibia Yuanmoupithecus xiaoyuan Hylobatidae Yuanmoupithecinae 8.2–7.1 Late Miocene China Griphopithecus suessi Hominidae Kenyapithecinae 13.8–12.7 Late Miocene Austria, Slovakia Griphopithecus alpani Hominidae Kenyapithecinae 14.5–13.4 Middle Miocene Turkey Kenyapithecus wickeri Hominidae Kenyapithecinae 13.8 Middle Miocene Kenya Kenyapithecus kizili Hominidae Kenyapithecinae 14.5–14.0 Middle Miocene Turkey Dryopithecus fontani Hominidae Dryopithecinae 12.5–11.0 Middle to Late Miocene Austria, France, Spain Pierolapithecus catalaunicus Hominidae Dryopithecinae 12.0 Middle Miocene Spain Anoiapithecus brevirostris Hominidae Dryopithecinae 12.4–12.0 Middle Miocene Spain Hispanopithecus laietanus Hominidae Dryopithecinae 10.0–9.6 Late Miocene Spain Hispanopithecus crusafonti Hominidae Dryopithecinae 10.3–10.0 Late Miocene Spain Rudapithecus hungaricus Hominidae Dryopithecinae 10.0–9.8 Late Miocene Hungary Danuvius guggenmosi Hominidae Dryopithecinae 11.6 Late Miocene Germany ?Udabnopithecus garedziensis Hominidae Dryopithecinae 8.1–7.7 Late Miocene Georgia Khoratpithecus chiangmuanensis Hominidae Ponginae 12.4–12.2 Middle Miocene Thailand Khoratpithecus piriyai Hominidae Ponginae 9.0–6.0 Late Miocene Thailand Khorapithecus ayeyarwadyensis Hominidae Ponginae 10.4–8.0 Late Miocene Myanmar Khoratpithecus magnus Hominidae Ponginae 9.0–6.0 Late Miocene Thailand Ankarapithecus meteai Hominidae Ponginae 9.8 Late Miocene Turkey Lufengpithecus lufengensis Hominidae Ponginae 6.9–6.2 Late Miocene China Sivapithecus sivalensis Hominidae Ponginae 10.4–7.5 Late Miocene India, Pakistan Sivapithecus indicus Hominidae Ponginae 13.0–10.8c Middle to Late Miocene India, Pakistan Sivapithecus parvada Hominidae Ponginae 10.1–10.0 Late Miocene Pakistan Indopithecus giganteus Hominidae Ponginae 8.9–8.6 Late Miocene India, Pakistan Chororapithecus abyssinicus Hominidae Homininae 8.0 Late Miocene Ethiopia Nakalipithecus nakayamai Hominidae Homininae 9.9–9.8 Late Miocene Kenya Graecopithecus freybergi Hominidae incertae sedis 7.2 Late Miocene Greece Ouranopithecus macedoniensis Hominidae incertae sedis 9.7–8.8 Late Miocene Greece Ouranopithecus turkae Hominidae incertae sedis 8.1–7.6 Late Miocene Turkey Sinopithecus keiyuanensis Hominidae incertae sedis 12.5–11.6 Middle Miocene China Sinopithecus hudienensis Hominidae incertae sedis 8.2–7.1 Late Miocene China Oreopithecus bambolii incertae sedis incertae sedis 8.3–6.7 Late Miocene Italy ‘Dryopithecus’ wuduensis incertae sedis — 8.3 Late Miocene China a 148 Pickford et al. (2017) and Pickford (2021) identified A. turkanensis, P. gitongai, and N. kerioi at Moroto, whereas MacLatchy et al. (2019) 149 recognized ?M. bishopi and Proconsulidae indet. and Jansma and MacLatchy (2015) further reported a nyanzapithecid. We have tentatively 150 included cf. P. gitongai and cf. N. kerioi in SOM Table S1, but not considered them in the ranges of this table. If confirmed by subsequent 151 studies, they would modify the chronostratigraphic ranges of these taxa, depending on the age attributed to Moroto (see next footnote). b 152 The age of Moroto localities has been much debated. It is noteworthy that some researchers (e.g., MacLatchy et al., 2019) favor the 153 radiometric date of >20.6 Ma (Gebo et al., 1997) for Moroto, whereas Pickford and colleagues have favored younger ages of ~17.5–17.0 Ma 154 (Pickford et al., 1986, 2003; Pickford and Mein, 2006) or even ~16.5–15.5 (Pickford et al., 2017) on biostratigraphic grounds. Most recently, Van 155 Couvering and Delson’s (2020) provided a date of 19.3 Ma based on the redating of Bukwa at ~19 Ma (MacLatchy et al., 2006; Cote et al., 156 2018), which are somewhat older than Pickford’s (2017) radiometric dates for the same site (~18.0–17.5 Ma). Until new radiometric date for 157 Moroto are published, we follow Cote (2018) in considering that biostratigraphic data support an older age for Moroto, tentatively around 158 ~21.0–20.0 Ma. c 159 Kelley (2002) reported an approximate age range of ~12.5–10.5 Ma for S. indicus. The oldest citation from Pakistan would be 12.8–12.7 Ma 160 according to Barry et al. (pers. comm. in SOM Table S1). Remains from Ramnagar in India (see SOM Table S1) might be older (13.8–12.5 Ma), 161 probably close to ~13.0 Ma (Gilbert et al., 2020a), which is the maximum date considered here until the dates are clarified further. Supplementary Online Material (SOM): Systematics of Miocene apes: State of the art of a neverending controversy Alessandro Urciuoli1,2,3 & David M. Alba3,* 1 Universitat Autònoma de Barcelona, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain. 2 Division of Palaeoanthropology, Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt am Main, Germany. 3 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. *Corresponding author. E-mail address: david.alba@icp.cat (D.M. Alba). 1 SOM S1 The principles of systematics and phylogenetic inference 1.1. Biological classifications as human constructs based on a natural system Systematics aims to classify organisms according to a ‘natural’ (as opposed to ‘artificial’) system, which implies the existence of an external reality that needs to be discovered. Evolution, as the unifying paradigm for life sciences, provides such an objective basis for biological classification, and cladogenesis (beginning with speciation) further justifies the hierarchical structure of the system. Phylogenetic inference therefore plays a central role in the practice of systematics. However, besides cladogenesis, evolution further implies anagenesis, and there are different systematic approaches to reflect the patterns originated by these evolutionary processes—respectively, taxonomic diversity and morphological disparity (for a distinction between these concepts, see e.g., Briggs et al., 1992; Foote, 1994)—into classification, particularly given the double (both utilitarian and evolutionary) function of biological classifications (Benton, 2000). Despite the universal acceptance that systematics must reflect the evolutionary relationships among taxa, biological classifications are necessarily human constructs, so that there is no single true classification to be discovered in Nature (Benton, 2000). As a result, choosing among competing systematic classifications depends on both the systematic philosophy of each taxonomist and the phylogenetic hypotheses perceived by them as more accurate. 1.2. Monophyletic vs. paraphyletic groups With the advent of phylogenetic systematics (Hennig, 1966), emphasis shifted from disparity to phylogeny (determined on the basis of synapomorphies), but the treatment of paraphyletic groups (i.e., those including taxa that share a common ancestor but that do not include all of its descendants) has proven controversial ever since. Phylogenetic systematics considers that only clades (i.e., strictly monophyletic groups) must be considered natural, such that paraphyletic groups should not be treated as taxa because they are based on the crossing of an arbitrary morphological threshold (or ‘grade’). Of course, paraphyletic groups can be inadvertently erected as presumed monophyletic taxa within a cladistic framework when most parsimonious cladograms do not reflect the true phylogeny of the group. Nevertheless, from an evolutionary viewpoint, the possibility that an originally monophyletic taxon becomes paraphyletic is contingent on it giving rise to a morphologically distinct subclade instead of becoming extinct (Carroll, 1988). Paraphyly is thus inescapable at the species and genus ranks because new species necessarily evolve from a pre-existing one and—unless we abandon 2 Linnean nomenclature—every species must belong to a genus (Sarmiento et al., 2002). This problem is often overlooked due to the cladistic convention that phylogenetic relationships must be depicted as dichotomic branching patterns, as if all species were monophyletic. This may generally hold true when only extant species are analyzed, but when applied to extinct species, sister-taxon relationships do not reflect the difference between sister species originated from a common ancestor and ancestor–descendant species pairs. In summary, paraphyly is an expected result of evolution (Carroll 1988) except when a lineage goes extinct. Therefore, some degree of tolerance toward paraphyly is advisable at lower taxonomic ranks. As noted by Sarmiento et al. (2002: 54), “Although at higher levels (above genus) a paraphyletic taxon is unacceptable, at lower levels it is inevitable”. To solve the purported problem of paraphyly, some authors have advocated the abandonment of binomial nomenclature and Linnean ranks altogether (de Queiroz and Gauthier, 1992; Cantino et al., 1999; de Queiroz and Cantino, 2001; Bryant and Cantino, 2002; Lee and Skinner, 2007). However, this proposal that has not been exempt of criticism (Benton, 2000; Forey, 2002) and has not gained wide acceptance. Adherence to the so-called phylogenetic nomenclature largely reflects a fundamental confusion about the difference between a phylogeny (which is real) and biological classification (which must be useful). Biological classifications must always be somewhat arbitrary: first, because they are conceived as practical means to efficiently transmit scientific knowledge; and second, because they cannot faithfully depict all the details of the continuum represented by the tree of life. Therefore, this work relies on the use of Linnean ranks—not because they have any intrinsic biological meaning, but rather because of their practical utility to simultaneously reflect (albeit imperfectly) both disparity and relatedness. Moreover, the use of Linnean ranks is not at all incompatible with trying to avoid paraphyly at suprageneric ranks—while recognizing that its avoidance at the genus and species-group levels is futile. 1.3. The epistemological basis of cladistics Cladistics has been the predominant paradigm in morphology-based phylogenetic inference for several decades now. The word ‘cladistics’ may be employed as a synonym of ‘phylogenetic systematics’ or more specifically refer to a particular methodology of phylogenetic inference (i.e., cladistic analysis). Much has been written from the viewpoint of the philosophy of science about cladistic analysis, which is based on the main premise that only shared-derived features (synapomorphies) are phylogenetically informative (Hennig, 1966; Farris, 1983), as well as in the application of the principle of maximum parsimony. This 3 principle, first introduced in phylogenetics as the ‘principle of minimum evolution’ (Edwards, 1996), stems from the assumption that homology must be presumed in the lack of evidence to the contrary (Hennig, 1966; Wiley, 1975), which is equivalent to assume that homoplasy must not be postulated beyond necessity a priori (Farris, 1983). In this sense, maximum parsimony is but the application in phylogenetic inference of an old and well-known epistemological approach that restricts inferential reasoning to the simplest theoretical framework necessary to account for the data (Kluge, 1984; Brower, 2000). Following Wiley (1975) and Farris (1983), the scientific character of cladistic analysis based on maximum parsimony has generally been formulated as a refutationist (falsificationist) approach in terms of Popperian testability (Kluge, 1997, 1999, 2001). According to this interpretation, cladistic hypotheses (cladograms) would be composite explanations consisting of hypotheses of monophyly (cladogenetic events) and homology (character transformation events; Grant and Kluge, 2003), which are tested on the basis of the principle of parsimony. The latter emphasizes the degree of corroboration for deciding among competing hypotheses, so that the most parsimonious cladogram is the least disconfirmed and, hence, the most highly corroborated hypothesis (Kluge, 1999). Traditionally, it has been argued that parsimony maximizes explanatory power (presumed homologies) by minimizing the requirement for ad hoc hypotheses of homoplasy (Farris, 1983; Kluge, 1997). Subsequent reformulations have alluded to the ‘antisuperfluity principle,’ according to which explanatory power is maximized by minimizing the number of transformation events required as hypotheses of homology (Kluge and Grant, 2006). From an operational viewpoint, similarity is only relevant for proposing hypotheses of primary homology based on similar traits shared by several taxa (Kluge, 1997). Hypotheses of primary homology, formalized as character statements, must be tested on the basis of the most parsimonious hypothesis (de Pinna, 1991), thus being corroborated (secondary homology) or refuted (homoplasy) a posteriori. Despite the claims to the contrary (Brower, 2000), cladistics must assume evolution (descent with modification) as background knowledge so as to interpret secondary homologies as such, as well as to interpret most parsimonious cladograms in terms of phylogenetic hypotheses. However, unless the most parsimonious hypothesis coincides with the true phylogeny, homoplasy as measured by cladistic analyses will always underestimate the true degree of homoplasy (Archie, 1996). In the lack of a better solution for the problem of homoplasy, a priori character weighting (Neff, 1986) might seem appealing. However, it is contrary to the epistemological basis of cladistics, according to which hypotheses of homology can only be corroborated or refuted a posteriori. Methods of 4 character weighting a posteriori, such as successive weighting (Farris, 1969; Carpenter, 1988, 1994) or especially implied weighting (Goloboff, 1993, 1997) seem preferable, as the latter approach allows considering some character state transformations as more reliable than others as a consequence of the analysis, not as an assumption. Other alternative approaches include using a modified concept of parsimony (as in stratocladistics; see SOM S1.4) or probabilistic methods (such as Bayesian analyses; SOM S1.5). Cladistics has been criticized by some researchers by arguing that it does not adhere to Popper’s (1959, 1962) approach to the philosophy of science (e.g., Cartmill, 1981; Hull, 1988). Other authors have supported the scientific character of cladistics but questioned the interpretation of parsimony as a falsificationist method (Rieppel, 2003; Vogt, 2008; de Queiroz and Poe, 2001, 2003; de Santis, 2021). Some of the latter authors have argued that parsimony methods are only valid in Popperian terms if they incorporate probabilistic assumptions, with likelihood methods being considered the basis of Popper’s degree of corroboration (de Queiroz and Poe, 2001, 2003; de Queiroz, 2004). This interpretation, which has been used to favor a unified and inclusive philosophy of phylogenetic inference (de Queiroz, 2004), has been criticized by others based on the contention that Popper’s explanatory power is maximized deductively instead of inductively (Kluge and Grant, 2006). However, the latter position seems difficult to maintain and it is more reasonable to accept that testing among competing cladogram hypotheses cannot be based on Popperian falsificationism (Rieppel, 2003; Vogt, 2008, 2014). We therefore concur with the views of Rieppel (2003) and Vogt (2008, 2014), according to whom: (1) cladistics is better interpreted as an inductive approach that is not based on Popperian falsificationism; (2) the hypothetico-deductive setting that constitutes the basis of Popperian falsificationism, developed for experimental sciences, must not necessarily be the only valid scientific approach, particularly for historical sciences such as phylogenetics; (3) hypothesis testing is not unique to Popperian falsificationism and plays a central role in phylogenetic inference by means of the application of general concepts that allow for hypothesis testing. Whether this means that cladograms are unfalsifiable (Vogt, 2008, 2014) or falsifiable (Crother and Murray, 2015) outside a Popperian philosophical framework is debatable—because a philosophy of phylogenetic inference independent from Popper’s falsificationist approach would be necessary but is still a work in progress (Rieppel, 2003; de Queiroz, 2004; Helfenbein and DeSalle, 2005; Vogt, 2008; de Santis, 2021). However, this represents a rather semantical issue, as long as all methods of phylogenetic inference allow for testing among competing cladograms and potentially refuting them based on some general 5 principle (such as maximum parsimony, among others). In any case, there can be little doubt that parsimony analysis differs epistemologically from probabilistic (likelihood and Bayesian) methods because the latter depend on specific evolutionary models (Huelsenbeck and Rannala, 1997; Goloboff et al., 2018), which arguably have some advantages and associated problems simultaneously, as explained in SOM S1.5. 1.4. Stratocladistics Stratocladistics is a method of phylogenetic inference alternative to conventional cladistics that was developed during the 1990s to simultaneously consider morphological and chronostratigraphic data (Fisher, 1991, 1994, 2008). This method adheres to a similar epistemology to conventional cladistics (SOM S1.3) and is thus based on parsimony, but differs from conventional cladistics by minimizing ad hoc hypotheses of both character homoplasy and non-preservation in the fossil record (ghost lineages). Other differences of stratocladistics from conventional cladistics include taking into account autapomorphies and performing the analyses at the level of phylogenetic tree. This allows stratocladistic methods to infer ancestor– descendant relationships (Fisher, 2008), which are considered untestable under a strict cladistic paradigm. When stratocladistics was developed, several researchers warned against approaches incorporating stratigraphic data a priori because they would prevent using phylogeny as an independent test other temporal trends (Smith, 2000; Sumrall and Brochu, 2003), favoring instead the use of metrics measuring the congruence between cladograms and the stratigraphic record (e.g., Huelsenbeck, 1994; Hitchin and Benton, 1997). Nevertheless, if it is conceded that stratigraphic data are relevant for inferring phylogeny, there is no reason to insist that they cannot in principle overturn parsimony considerations based on morphological data (Grantham, 2004). This contention fulfills the principle of total evidence—a basic maxim for non-deductive inference frequently advocated in cladistics— according to which “evidence must be considered if it has relevance to an inference” (Fitzhugh, 2006: 309). Stratocladistics thus minimizes the number of homoplasies and the number of ghost lineages simultaneously based on the concept of total parsimony debt (the sum of morphologic parsimony and stratigraphic parsimony debts) by assuming that an ad hoc hypothesis of preservation probability represents as much loss in explanatory power as an ad hoc hypothesis of homoplasy (Clyde and Fisher, 1997; Fisher, 1999, 2008). Despite criticisms, stratocladistic analyses with real data performed reasonably well, yielding cladograms with much lower stratigraphic parsimony debts than conventional cladistics at the expense only of slightly higher 6 parsimony debts (Clyde and Fisher, 1997). Simulation studies further supported that stratocladistics outperforms the accuracy of conventional cladistics (Fox et al., 1999). 1.5. Bayesian analyses Bayesian methods of phylogenetic inference using Markov chain Monte Carlo techniques developed two decades ago (Huelsenbeck et al., 2001; Holder and Lewis, 2003) can be applied to discrete morphological data and thus represent an alternative to parsimony analysis (Lewis, 2001a, 2001b; Nylander et al., 2004). Multiple simulation studies have shown that Bayesian methods outperform parsimony in terms of accuracy (Wiens, 2005; Wright and Hillis, 2014; O’Reilly et al., 2016, 2018; Puttick et al., 2019; Keating et al., 2020), particularly when homoplasy is high (Puttick et al., 2019), even if the former perform less efficiently in terms of node resolution (O’Reilly et al., 2016; Smith, 2019). Simulations further indicate that Bayesian analyses are less sensitive to long-branch attraction problems and that even very incompletely preserved extinct taxa can improve accuracy, unlike in parsimony analyses (Wright and Hillis, 2014). Moreover, Bayesian analyses allow for the simultaneous analysis of morphologic and molecular data, resulting (like stratocladistics) in time-calibrated phylogenies. Molecular evidence is customarily incorporated in parsimony analyses using a molecular backbone (e.g., Pugh, 2022) that does not inform about divergence times. In turn, fossil data are often included in molecular analyses to constrain the estimates of divergence times (e.g., Roos et al., 2019), an approach known as ‘node dating’. In contrast, total-evidence analyses (combining morphologic and molecular data) based on Bayesian methods date divergence times by directly incorporating the age of the all the analyzed fossils, a procedure known as ‘tip dating’ or ‘total- evidence dating’ (e.g., Pyron, 2011; Ronquist et al., 2012, 2016; Donoghue and Yang, 2016). This approach has become more popular since the introduction of the fossilized birth–death process that models extant and fossil data as outcomes of the same macroevolutionary process (Heath et al., 2014; Zhang et al., 2016; King, 2021; Mongiardino Koch et al., 2021). Tip dating differs from stratocladistics (see SOM S1.4) because topologies implying a higher stratigraphic debt are not explicitly penalized, although it has a similar result by making it more unlikely the grouping of morphologically similar but stratigraphically distant taxa (Hunt and Slater, 2016). Simulation studies have shown that tip-dated Bayesian analyses yield different topologies than both undated Bayesian and parsimony analyses, with the former having a better fit with stratigraphic data (King, 2021) and a greater accuracy (Mongiardino Koch et al., 2021), at least as long as fossil age uncertainties (Barido-Sottani et al., 2020) as well as fossilization and sampling biases (Zhang et al., 2016) are adequately modeled. Nevertheless, Bayesian 7 methods had not been exempt from criticism, particularly regarding simulation results indicating that they have a lower resolution than parsimony methods (O’Reilly et al., 2016; Smith, 2019) or suggesting that the latter yield comparable, if not better, results as long as implied weighting is used (Goloboff, 2018; Smith, 2019; but see Keating et al., 2020). One of the caveats that currently apply to tip dating is that, unlike parsimony (Goloboff et al., 2006; Goloboff and Catalano, 2016), Bayesian methods do not permit the treatment of continuous morphological data as such, which is beneficial for reducing the subjectivity and potential loss of information implicit in the use of discrete characters (Parins-Fukichi, 2017), even if discretized using the gap-weighted method (Thiele, 1993). Nevertheless, recent developments of Bayesian methods are most promising in this regard (Álvarez-Carretero et al., 2017; Parins- Fukuchi, 2018). Even more important from an epistemological viewpoint is the fact that, unlike parsimony, Bayesian methods necessitate the a priori assumption of a particular evolutionary model of uncertain applicability to morphological data (Goloboff, 2018). On the other hand, this allows for further improvement in the future (as more realistic methods are developed) as well as the possibility to tackle potential problems of long-branch attraction (which parsimony cannot directly address because it does not take branch length or evolutionary rates into account; Pagel, 1999). 1.6. Apomorphy vs. clade-based definitions of taxa Even if paraphyletic groups are avoided above the genus rank, not all clades can be erected as taxa. The advent of cladistics originally favored the use of apomorphy-based definitions of taxa, but subsequently phylogenetic nomenclature promoted instead clade-based definitions based on common ancestry (de Queiroz and Gauthier, 1990), which refer to a common ancestor of two taxa included in the taxon being defined. Two types of clade-based definitions of taxa are possible (de Queiroz and Gauthier 1992; Benton 2000): node-based definitions, which include all the descendants of the last common ancestor; and stem-based definitions, which further include those taxa more closely related to this last common ancestor than that of its closer formally-defined sister-taxon. The Code (ICZN, 1999) does not specify how taxa should be defined, so that Linnean ranks are not restricted to taxa defined on the basis of characters (Benton, 2000). Although both apomorphy-based and clade-based definitions may be useful, apomorphy-based definitions are the most problematic because they rely on an arbitrary selection of one or more synapomorphies that ultimately constitute a grade. Such a selection is particularly problematic when dealing with extinct taxa, due to the mosaic nature of evolution— the more basal is an extinct member of a clade, the more likely it will lack the synapomorphies 8 of more advanced members of the same clade. Furthermore, some of the purported synapomorphies employed to define a particular group might eventually prove to be homoplastic. In contrast, clade-based definitions are more stable, and stem-based definitions are further favored here for groups with extant representatives. 1.7. Stem lineage vs. crown group For an extant monophyletic group formally erected as a taxon, we can distinguish the crown group from the stem lineage (Jefferies, 1979; Ax, 1985; Smith, 1994; Benton, 2000), which together constitute the total group. The crown group is the clade that includes all the extant representatives of the taxon plus extinct representatives more closely related to some (but not all) of them. In contrast, the stem lineage is a paraphyletic assemblage of basal taxa that are more or less distantly (but equally) related to all crown group members, but more closely related to the latter than to its sister-group. Extinct representatives of the taxon’s total group will be included into its crown group or stem lineage, depending on their particular phylogenetic relationships. Although supporters of node-based definitions of taxa have tended to restrict taxon names to the crown group (de Queiroz and Gauthier, 1992; Rowe and Gauthier, 1992), we prefer stem-based definitions, which apply to the whole community of descent (total group), because it may be difficult to determine whether a particular extinct species is a stem or a crown member of a particular clade. With a stem-based definition, the inclusion of this particular species into the taxon remains stable under both phylogenetic hypotheses. The crown group constitutes a clade and can be therefore formally erected as a taxon, whereas the stem lineage will rarely constitute a clade and, hence, it would constitute a paraphyletic taxon if formally designated. To avoid paraphyly, the use of several subtaxa (even if monotypic) for various subclades of the stem lineage (with the same rank as the crown group) is favored here—unless the current state of knowledge does not enable to discern whether the known members of the stem lineage constitute a monophyletic or paraphyletic assemblage. 9 SOM S2 Nomenclatural remarks 2.1. On the valid name for the family-group taxa including Kenyapithecus and Griphopithecus Begun (2001, 2002) distinguished the family Griphopithecidae to include either Griphopithecus s.l. (including Equatorius) + Afropithecus (Begun, 2001) or Griphopithecus s.l. + Kenyapithecus (Begun, 2002), while Kelley (2002) distinguished a subfamily Griphopithecinae for Griphopithecus within the Afropithecidae. More recently, Begun (2010) distinguished a subfamily Griphopithecinae for these three genera plus Nacholapithecus within the Hominidae, while Andrews (2020) restricted the Griphopithecidae to Griphopithecus. Nevertheless, as remarked by previous authors (Moyà-Solà et al., 2009b; Harrison, 2010; Casanovas-Vilar et al., 2011), the tribes Afropithecini and Kenyapithecini erected by Andrews (1992) would have priority over family-group names derived from Griphopithecus if Afropithecus and/or Kenyapithecus are included. Furthermore, as noted by Casanovas-Vilar et al. (2011), Begun (2001, 2002) and Kelley (2002) did not fulfill the conditions of the Code (ICZN: Art. 16.1 and 16.2), according to which every new family- group name published after 1999 must be explicitly indicated as intentionally new and accompanied by citation of the name of the type genus; the same applies to Andrews (2020). None of these authors stated taxonomic authorities, so it is unclear if they attributed the taxon names to a previous author. Only Begun (2002) denoted both Griphopithecinae (for Griphopithecus s.l.) and Griphopithecidae (including Griphopithecinae + Kenyapithecinae) as ‘new rank’ but, to our knowledge, previous usage of this family-group taxon is restricted to Begun’s (2001) Griphopithecidae. In any event, no family-group name derived from Griphopithecus is nomenclaturally available from these publications, irrespective of whether Afropithecus and Kenyapithecus are excluded, as in Kelley (2002) and Andrews (2020). 2.2. On the valid name for the European species of Griphopithecus There are two species names available for the Griphopithecus species recorded in Germany and Slovakia, which were erected in the same publication based on material from the same locality: Griphopithecus suessi Abel, 1902 (the type species of the genus) and Dryopithecus darwini Abel, 2012. As noted by Simonetta (1958), Güleç and Begun (2003), and Casanovas-Vilar et al. (2011), either Remane (1921a) or Remane (1921b)—not Glaessner (1931), as argued by Holec and Emry (2003)—acted as the First Reviser (ICZN, 1999: Art. 24.2) by choosing D. darwini as the senior synonym. Following the resurrection of 10 Griphopithecus by Andrews et al. (1996), the combination Griphopithecus darwini has been frequently used (e.g., Begun, 2002, 2009; Kelley, 2002; Güleç and Begun, 2003). Nevertheless, G. suessi is the type species of Griphopithecus by monotypy (Holec and Emry, 2003; Casanovas-Vilar et al., 2011). A determination of precedence based on the Principle of the First Reviser is nullified if it can be subsequently shown that the precedence can be determined objectively. If it is interpreted that the designation of a type species fulfills the latter condition (Holec and Emry, 2003; Casanovas-Vilar et al., 2011), then the Principle of the First Reviser does not apply and the valid name for the species is G. suessi, with G. darwini being its junior subjective synonym. 2.3. On the valid name of the tribe including Sivapithecus Although the intuitive name for a tribe including Sivapithecus would be Sivapithecini Pilbeam et al., 1977 (e.g., Andrews, 1992; Kelley, 2002), Bramapithecini Simonetta, 1958, Ramapithecini Simonetta, 1958, and Sugrivapithecini Simonetta, 1958 take precedence. Note that, although the publication year of these nominal taxa has generally been attributed to 1957 (e.g., Szalay and Delson, 1979), as noted by Ceccolini (2022) the publication was not issued until 1958—the date of the proofs stated in the last contribution of the same volume is February 4th, 1958, indicating that it could not have been issued before this date. Simonetta (1958) included these tribes within the Dryopithecinae, being based on genera erected by Lewis (1934), whereas Sivapithecus was included in the nominotypical tribe (Dryopithecini). Simonetta’s (1958) spelling of Bramapithecini was Brahmapithecini, because he intentionally emended the spelling of Bramapithecus Lewis, 1934 into Brahmapithecus. However, although Lewis (1934) explicitly dedicated the genus to ‘Brahma’, the genus name was consistently written in Lewis (1934). Incorrect transliteration or latinization cannot be taken as an inadvertent error (ICZN, 1999: Art. 32.5), so that Simonetta’s (1958) emendation of the genus name must be considered unjustified (ICZN, 1999: Art. 33.2). This means that Brahmapithecus Simonetta, 1958 is a junior objective synonym of Bramapithecus Lewis, 1934 and that the name of the tribe must be corrected as well (ICZN, 1999: Art. 35.4). Neither Simons and Pilbeam (1965) nor Szalay and Delson (1979) followed Simonetta’s (1958) emendation of Bramapithecus, and Szalay and Delson (1979) apparently considered Brahmapithecini a lapsus, which was not. Pilbeam et al. (1977) distinguished the subfamilies Sivapithecinae and Ramapithecinae, but Szalay and Delson (1979) considered that Sivapithecini was a junior synonym of Sugrivapithecini. Following Simons and Pilbeam (1965), Szalay and Delson (1979) 11 considered Bramapithecus a subjective synonym of Ramapithecus Lewis, 1934 (then included in the Hominidae), and hence listed both Ramapithecini and Bramapithecini as synonyms of this family. In contrast, following Simons and Pilbeam (1965), Szalay and Delson (1979) considered that Sugrivapithecus Lewis, 1934 was a junior subjective synonym of Sivapithecus Pilgrim, 1910, and hence considered that Sugrivapithecini had priority over Sivapithecini. Given that Lewis’ (1934) genera are all currently considered junior subjective synonyms of Sivapithecus, the three tribes erected by Simonetta (1958) must be considered subjective synonym as well, but their priority cannot be objectively determined. Following Szalay and Delson (1979), Alba (2012) used Sugrivapithecini and noted it had priority over Sivapithecini, but failed to comment as to why Sugrivapithecini would have priority over Ramapithecini or Bramapithecini. According to the Principle of the First Reviser (ICZN, 1999: Art. 24.2), when the precedence between names cannot be objectively determined, the precedence is fixed by the action of the first author citing in a published work those names and selecting from them. Given that Szalay and Delson (1979) did not synonymize the three tribes erected by Simonetta’s (1958) and that Alba (2012) did not cite two of the three available names, neither of them qualifies as First Reviser. As we have been unable to find another work where these family-group names are mentioned, the present paper should fulfill the requirements of the Principle of the First Reviewer by stating that, to preserve current usage (Szalay and Delson, 1977; Alba, 2012), we choose Sugrivapithecini over Ramapithecini and Bramapithecini as the valid name for the tribe including Sivapithecus and its junior subjective synonyms Ramapithecus, Bramapithecus, and Sugrivapithecus. It is noteworthy that the name Gigantopithecinae von Koenigswald, 1958 might be available for any family-group taxon including Gigantopithecus von Koenigswald, 1935. Although von Koenigswald (1958) did not explicitly note his will to establish a new taxon or designate a type genus, these requirements were not mandatory at the time (ICZN, 1999: Art. 16). As we include Gigantopithecus in the same tribe as Sivapithecus, Gigantopithecini von Koenigswald becomes a synonym of Sugrivapithecini as used here. Although we have been unable to determine the exact publication date of von Koenigswald's (1958) contribution in a conference proceedings volume, it could not have appeared before May 23th, 1958, when the conference has held, while Simonetta (1958) likely appeared shortly after February 4th the same year (see above). As such, there is currently no reason to support the precedence of Gigantopithecini over Sugrivapithecini. 12 2.4. On the valid name of the tribe including Ouranopithecus and Graecopithecus The tribe Graecopithecini was first used in two articles published in the same year (Cameron, 1997a, 1997b). Only Cameron (1997b) explicitly noted the intention to erect a new taxon but, as both articles were published before 2000, this requirement of the Code (ICZN, 1999: Art. 16.1) does not apply for the name to be valid. In any event, based on publication date, the authorship of the taxon must be attributed to Cameron (1997b)—unlike for Hispanopithecini, which was only mentioned by Cameron (1997a) and must be considered available from that publication despite the lack of explicit mention of the erection of a new taxon, for the reason outlined above. Both Graecopithecus freybergi and Ouranopithecus macedoniensis were included in the Graecopithecini by Cameron (1997b), although the latter species was included in Graecopithecus by this author. The different opinions about the synonymy between Ouranopithecus and Graecopithecus and the taxonomic validity of G. freybergi have caused some confusion as to the correct name for the tribe including these taxa, particularly since another family-group taxon, the subtribe Ouranopithecina Begun, 2009, was subsequently described to include both genera. Alba (2012) elevated the latter taxon to tribe rank (i.e., Ouranopithecini Begun, 2009) to refer exclusively for Ouranopithecus, implicitly considering that Graecopithecus and its type species were nomina vana (Casanovas-Vilar et al., 2011). The latter authors remarked that Graecopithecini Cameron, 1997b would take precedence to include Ouranopithecus if Graecopithecus was considered a taxonomically valid genus included in the same tribe. As this is the opinion followed in the present paper, Ouranopithecini Begun, 2009 must be considered a junior subjective synonym of Graecopithecini Cameron, 1997b, although the former remains available if Ouranopithecus and Graecopithecus were eventually classified in different family-group taxa of the same rank. 13 SOM S3 On the deformation of the Pierolapithecus cranium Based on the distortion of the Pierolapithecus cranium, Begun (2009, 2010) casted doubts on Moyà-Solà et al.’s (2004) interpretation of its facial profile as more plesiomorphic than that of crown hominids. Begun (2015: 1302–1303) further considered that the face of Anoiapithecus is “distorted” and that of Pierolapithecus is “seriously damaged,” but provided no further details in this regard. According to the more detailed account of the Pierolapithecus cranium preservation published by Pérez de los Ríos et al. (2012), the most serious issue relates to the missing contact between the premaxilla and the maxilla. Nevertheless, the study of Pérez de los Ríos et al. (2012), which was focused on internal cranial anatomy, failed to notice the difficulties in adequately orienting the lower face relative to the upper face. Based on our own evaluation of the original specimen, the distortion that affects the fossil in multiple areas is not caused by plastic deformation of the bone but by multiple cracks filled with matrix and the consequent displacement of some fragments from their original position, which can be reasonably corrected by means of virtual 3D reconstruction. The latter is currently underway—see preliminary results by Pugh et al. (2022), which are consistent with the stem hominid status originally favored by Moyà-Solà et al. (2004) and later supported by cladistic analyses (Alba et al., 2015; Pugh, 2022). Even if the reconstructed facial profile of Pierolapithecus is ultimately demonstrated to be more orthognathous than preserved, it will be difficult to reconcile with the much more orthognathous facial profile of Anoiapithecus, which further displays other important craniodental differences that justify their generic distinction (Moyà-Solà et al., 2009b; Pérez de los Ríos et al., 2012). 14 SOM S4 The phylogenetic relationships between Ouranopithecus, Nakalipithecus, and hominines The cladistic analyses of Pugh (2022) supported a stem hominine status for Ouranopithecus and Graecopithecus, and further lent some support to the phylogenetic link between Ouranopithecus and Nakalipithecus (as previously hypothesized by Kunimatsu et al., 2007). We consider such phylogenetic links plausible but insufficiently supported. For this reason, we refrain from including Nakalipithecus into the tribe Graecopithecini and also from classifying the latter taxon within the Homininae. We provide some discussion below as to why we consider this link insufficiently supported. First, as recognized by Pugh (2022), most of the purported hominine synapomorphies of graecopithecins are ambiguous because they are present in many other taxa (e.g., relatively broader P3, inflated glabella, or square orbits). Second, other purported hominine synapomorphies can be easily explained by the overall cranial robusticity of Ouranopithecus (e.g., robust supraorbital 'torus' and broad nasal aperture). A geometric morphometric analysis of the Ouranopithecus macedoniensis face found closer similarities with gorillas among extant great apes (Ioannidou et al., 2019). This confirmed the previous morphometric results by McNulty (2005), which were based exclusively on the supraorbital region, thus apparently supporting phenetic affinities between Ouranopithecus and hominines. However, as recognized by Ioannidou et al. (2019), about half of the variance of the axis distinguishing gorillas from chimpanzees and orangutans was correlated with size, while a reconstruction of the Hispanopithecus laietanus face was also classified as gorilla despite occupying an intermediate position on the morphospace among the three great ape genera (Ioannidou et al., 2019). This suggests that the Ouranopithecus might resemble gorillas in facial morphology more closely than dryopithecines simply owing to size-scaling (allometric) effects. Third, the definition and/or scoring of some of the characters mentioned above is debatable. For example, Pugh’s (2022) scored the orbits of Sivapithecus and Pongo with the same state (‘rounded’) as those of Ekembo and hylobatids (among many other taxa). This arguably obscures the derived condition of the former in this regard and, in any event, according to Pugh (2022) the purportedly derived ‘squared’ state of Ouranopithecus is also present in the dryopithecine Rudapithecus. Regarding the supraorbital ‘torus,’ given previous controversies about the presence of supraorbital costae (Moyà-Solà and Köhler, 1995) vs. a supraorbital torus (Begun, 1994) in Hispanopithecus, Rudapithecus, and Ouranopithecus, Pugh (2022) explicitly refrained from coding this structure as such. Instead, she split it into five different characters to better capture the morphological variation in this area. Pugh (2022) 15 noted similarities between Ouranopithecus and Hispanopithecus in the configuration of the temporal lines and supraglabellar region, but interpreted them as primitive retentions. Nevertheless, of the five characters used by Pugh (2022) to code the morphology this region, only three (orientation of temporal lines, presence of transverse supratoral sulcus, and glabellar development) show differences between orangutans and African apes and could potentially be phylogenetically informative for resolving the closer phylogenetic relationships of graecopithecins among crown hominids. For the first two features, Ouranopithecus shows the same condition as dryopithecines and pongines, so that Ouranopithecus only more closely resembles African apes regarding the development of the glabella. Nevertheless, the same character state is displayed by Rudapithecus, while the pronounced supraglabellar depression of Ouranopithecus is shared with Hispanopithecus and seems autapomorphically derived for these taxa (as opposed to the apparently plesiomorphic condition retained by extant hominids and other dryopithecines). Finally, the interpretation of similarities in P3 morphology between Ouranopithecus, Nakalipithecus, and gorillas (Pugh, 2022) is also debatable. Kunimatsu et al. (2007) noted similarities between Nakalipithecus and Ouranopithecus in the morphology of the P3 (broader and with a more lingually oriented transverse cristid originating from the protoconid than in other Eurasian Miocene apes). Pugh (2022) elaborated further on these differences by noting that Nakalipithecus and Ouranopithecus uniquely share with gorillas (Delezene and Kimbel, 2011) a distal curvature of the P3 transverse cristid. Nevertheless, according to Pugh’s (2022) scorings, this cristid is more distally oriented in gorillas than in Nakalipithecus and Ouranopithecus; the latter genera rather resemble, in this regard, Australopithecus, Indopithecus, and Gigantopithecus—which might be related to the independent molarization of this tooth (Pugh, 2022, and references therein). In contrast, the distal curvature of the cristid, according to Pugh (2022), would only be present in gorillas, Nakalipithecus, and some specimens of O. macedoniensis (but not in other specimens of the latter or in Ouranopithecus turkae). However, we do not consider this configuration to be fundamentally different from that variably displayed by some dryopithecines (except by the more sectorial and buccolingually compressed crown of the latter), including the holotypes of Dryopithecus fontani and H. laietanus (SOM Fig. S1). Pugh (2022) argued that some chimpanzees display a distally curved cristid but considered this configuration to be different from that of gorillas, Nakalipithecus, and some specimens of O. macedoniensis in lacking a pronounced protrusion into the distal fovea. However, such a description might be applicable to the aforementioned dryopithecines and, even if the morphology of the cristid in considered most similar between 16 the only available specimen of Nakalipithecus and some specimens of Ouranopithecus (something questionable in the light of the variability displayed by gorillas; SOM Fig. S1), the fact that this feature is variable in Ouranopithecus (Pugh, 2022) despite being dentally more derived than Nakalipithecus (Kunimatsu et al., 2007) renders its phylogenetic significance very tenuous. An alternative scoring of the P3 transverse cristid in apes and humans based on enamel-dentine junction shape (Davies et al., 2019) reported that all the gorillas and chimpanzees investigated displayed a protoconid cristid that is either distally deflected or that flattens to the level of the surrounding fovea—these taxa only differing by the more distal origin of the cristid (relative to the protoconid dentine horn) in most chimpanzees. The configuration of African apes regarding the orientation of the cristid is shared with hylobatids but not with orangutans, which like most hominins display a more transversely oriented cristid originating at or near the apex of the protoconid dentine horn (Davies et al., 2019). On this basis, the distal deflection of the cristid in Nakalipithecus, Ouranopithecus, dryopithecines, and hominines, with its multiple variants, might be more plausibly interpreted as a hominoid symplesiomorphy. 17 SOM S5 Paleobiodiversity dynamics To estimate Miocene ape paleobiodiversity through time, we divided the Miocene into equal-duration stratigraphic bins of 1 Myr (e.g., the bin 19–18 corresponds to 19.0–18.1 Ma), except for the most recent bin, which was restricted to 6.0–5.3 Ma (in agreement with the Miocene/Pliocene boundary). The maximum and minimum age for the ranges of each species were attributed to their respective bins, and a range-through approach was followed to compute diversity. Ranges that include a single dating ending in .0 Ma (e.g., 19.0 Ma) were thus assigned to the 19–18 bin, but ranges that include two datings ending in .0 Ma (e.g., 19.0–18.0 Ma) were assigned to a single bin (the 19–18 bin, as in the previous example), so as not to artifactually increase diversity counts. The sample is too small to undertake a rigorous study of diversity dynamics (including the computation of origination and extinction rates), but two different metrics of diversity were computed: range-through or total diversity (Ntot) and standing diversity (Nst). The former is the most standard measure of diversity and is computed as follows (Foote, 2000): Ntot = Nbt + NbL + NFt + NFL, where Nbt is the number of range-through taxa (those found before and after the interval considered), NbL is the number of bottom-only boundary crossers (those found before but not after the interval considered), NFt is the number of top-only boundary crossers (those found after but not before the interval considered), and NFL is the number of singletons (single-interval taxa). Given that Ntot includes singletons, this metric is particularly sensitive to variation in preservation rates (Foote, 2000). In contrast, Nst ignores single-interval taxa because it is computed as the average of two successive (bottom and top) boundary crossers, thereby being a more robust estimate of standing diversity at a point in time within the interval (Foote, 2000); it is computed as follows: Nst = (NbL + NFt + 2Nbt) / 2 = Nrt – NFL – ½ (NbL + NFt). The ranges for Miocene apes and per-bin metrics are shown in SOM Table S3. It should be taken into account that such metrics are seriously affected by multiple biases of the fossil record and can lead to misleading conclusions when interpreted literally (e.g., see discussion in Casanovas-Vilar et al., 2014). Nevertheless, they provide a rough idea of ape diversity changes throughout the Miocene. 18 SOM Figure S1. P3 morphology in extant great apes and a selected sample of Miocene apes, in occlusal view. The approximate course of the transverse cristid originating from the protoconid is outlined by a dotted blue line. All specimens are depicted as from the left side (mirrored when necessary), with the mesial and distal cristids oriented to define an arbitrary mesiodistal axis (with mesial on top), and not to scale. a) Nakalipithecus nakayamai, KNM- NA 46423. b) Ouranopithecus macedoniensis, KNM-NA 46423. c) IPS1803, Hispanopithecus laietanus (holotype, mirrored). d) IPS1764, H. laietanus (mirrored). e) HGP2, Dryopithecus fontani (holotype, mirrored). f) AMNH-M 80008, Pongo sp. g) AMNH- M 28253, Pongo pygmaeus (mirrored). h) Gorilla gorilla, AMNH-M 167340. i) Pan troglodytes, AMNH-M 90292 (mirrored). j) P. troglodytes, AMNH-M 89406. k) P. troglodytes, AMNH-M 89351 (mirrored). l) Pan paniscus, AMNH-M 86857 (mirrored). Panels a–b, h–j, and k reproduced from Pugh (2022). 19 SOM Table S1 Miocene ape species-locality occurrences, including classification to family and subfamily ranks (see Table 2), subepoch, locality and site, country, age range, and citations of primary literature.a For further details on each locality, see SOM Table S2. (Provided separately as an Excel file.) a When indicated as "Barry et al. (pers. comm.)", identifications of Siwalik hominoid remains checked or updated based on data kindly provided by John Barry, Larry Flynn, Jay Kelley, Michèle Morgan, and David Pilbeam on September 2022. SOM Table S2 Miocene ape-bearing localities (in alphabetical order), including site/geographic area, country, age range, subepoch, and citations for the age.a (Provided separately as an Excel file.) a When indicated as "Barry et al. 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(Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Hopwood (1933a); MacInnes (1943); Le Gros Clark and Leakey (1950, 1951); Le Gros Clark (1952); Pilbeam (1969a); Greenfield (1972)f; Andrews (1978); Bosler (1981); Martin (1981); Pickford and Andrews (1981); Harrison (2010); Proconsulidae - Proconsul africanus Legetet Fm. (Koru-Muhoroni area) Kenya Early Miocene 20,0 19,0 Martin (1981); Pickford and Andrews (1981); Pickford et al. (2009a); Harrison (2010) Proconsulidae - Proconsul africanus Songhor (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Le Gros Clark (1952); Andrews (1978); Bosler (1981); Pickford and Andrews (1981); Harrison (2010) Proconsulidae - Proconsul major Chamtwara Member (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Martin (1981); Harrison (2010) Proconsulidae - Proconsul major Koru Fm. (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Le Gros Clark and Leakey (1951); Bosler (1981); Martin (1981); Pickford and Andrews (1981); Harrison (2010) Proconsulidae - Proconsul major Legetet Fm. (Koru-Muhoroni area) Kenya Early Miocene 20,0 19,0 Martin (1981); Pickford and Andrews (1981); Harrison (2010) Proconsulidae - Proconsul major Locality 14 - Maize Crib (Koru-Muhoroni area) Kenya Early Miocene 20,0 19,0 Hopwood (1933a); Martin (1981) Proconsulidae - Proconsul major Locality 34 - Maize Crib (Koru-Muhoroni area) Kenya Early Miocene 20,0 19,0 Hopwood (1933a); Martin (1981) Proconsulidae - Proconsul major Napak I (Akisim) Uganda Early Miocene 20,5 19,0 Rafferty et al. (1995); Senut et al. (2000); Pickford et al. (2009a, 2020); Harrison (2010) Proconsulidae - Proconsul major Napak IV (Akisim) Uganda Early Miocene 20,5 19,0 Bishop (1964); Harrison (2010) Proconsulidae - Proconsul major Napak V (Akisim) Uganda Early Miocene 20,5 19,0 Bishop (1964); Pickford et al. (2009a, 2020); Harrison (2010) Proconsulidae - Proconsul major Napak IX (Akisim) Uganda Early Miocene 20,5 19,0 Gommery et al. (1998); Senut et al. (2000); Pickford et al. (2009a, 2020); Harrison (2010) Proconsulidae - Proconsul major Napak XIII (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul major Napak XV (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2009a, 2020, 2021) Proconsulidae - Proconsul major Napak XXVI (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2021) Proconsulidae - Proconsul major Napak XXX (Akisim) Uganda Early Miocene 20,5 20,5 Pickford et al. (2020) Proconsulidae - Proconsul major Napak XXXI (Akisim) Uganda Early Miocene 20,5 20,5 Pickford et al. (2020) Proconsulidae - Proconsul major Napak CC (Akisim) Uganda Early Miocene 20,5 19,0 MacLatchy and Rossie (2005) Proconsulidae - Proconsul major Songhor (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Le Gros Clark and Leakey (1950); Le Gros Clark (1952); Pilbeam (1969a); Andrews (1978); Pickford and Andrews (1981); Pickford (1985b); Rafferty et al. (1995); Harrison (2010); McNulty et al. (2015); Patel et al. (2017); Foecke et al. (2022) Proconsulidae - Proconsul gitongai BPRP K089A (Baringo) Kenya Middle Miocene 15,8 15,8 Hill et al. (1991); Harrison (2010) Proconsulidae - Proconsul gitongai BPRP K091 (Baringo) Kenya Middle Miocene 15,8 15,8 Pickford (1988); Harrison (2010) Proconsulidae - Proconsul gitongai Muruyur Fm. (Cheparawa) Kenya Middle Miocene 15,8 15,8 Pickford and Kunimatsu (2005); Harrison (2010) Proconsulidae - cf. Proconsul gitongai Moroto II (Karamoja) Uganda Early Miocene 21,0 20,0 Senut et al. (2000); Pickford et al. (2003, 2009a, 2017a) but see MacLatchy (2019) Proconsulidae - Proconsul meswae Locality 36 - Meswa Bridge (Koru-Muhoroni area) Kenya Early Miocene 22,5 22,5 Andrews et al. (1981); Pickford and Andrews (1981); Harrison and Andrews (2009); Harrison (2010) Proconsulidae - Proconsul legetetensis Napak CC (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2009a, 2020); Harrison (2010) Proconsulidae - Proconsul legetetensis Napak I (Akisim) Uganda Early Miocene 20,5 19,0 Harrison (2010); Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak IV (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak V (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2009a, 2020); Harrison (2010) Proconsulidae - Proconsul legetetensis Napak IX (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak V (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak XII (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak XIII (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak XV (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak XVIII (Akisim) Uganda Early Miocene 20,5 20,5 Pickford et al. (2020) Proconsulidae - Proconsul legetetensis Napak XXX (Akisim) Uganda Early Miocene 20,5 20,5 Pickford et al. (2021) Proconsulidae - Proconsul legetetensis Napak XXXI (Akisim) Uganda Early Miocene 20,5 20,5 Pickford et al. (2020) Proconsulidae - Proconsul sp. Fort Ternan (Fort Ternan) Kenya Middle Miocene 13,7 13,7 Andrews and Walker (1976); Andrews (1978); Harrison (1992, 2010) Proconsulidae - Kalepithecus songhorensis Chamtwara Member (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Harrison (1988, 2010) Proconsulidae - Kalepithecus songhorensis Koru Fm. (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Harrison (1988, 2010) Proconsulidae - Kalepithecus songhorensis Napak IV (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2010, 2021) Proconsulidae - Kalepithecus songhorensis Napak V (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2010, 2021) Proconsulidae - Kalepithecus songhorensis Napak XV (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2021) Proconsulidae - Kalepithecus songhorensis Napak XXXI (Akisim) Uganda Early Miocene 20,5 20,5 Pickford et al. (2021) Proconsulidae - Kalepithecus songhorensis Songhor (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Le Gros Clark and Leakey (1950); Andrews (1978); Harrison (1982, 1988, 2010); Nengo and Rae (1992) Proconsulidae - Kalepithecus kogolensis Moroto II (Karamoja) Uganda Early Miocene 21,0 20,0 Pickford et al. (2017a) Proconsulidae - Ekembo heseloni Hiwegi Fm. (Rusinga Island) Kenya Early Miocene 18,3 18,3 Leakey (1943); MacInnes (1943); Le Gros Clark (1950, 1952); Le Gros Clark and Leakey (1951); Napier and Davis (1959); Davis and Napier (1963); Simons and Pilbeam (1965); Andrews (1978); Harrison (1982, 2010); Walker and Pickford (1983); Walker et al. (1983, 1986, 1993); Beard et al. (1986); Langdon (1986); Teaford et al. (1988); Walker and Teaford (1988); Ruff et al. (1989); Ward et al. (1991); Begun et al. (1994); Beynon et al. (1998); Nakatsukasa et al. (2004); Dunsworth (2006); Michel et al. (2013); McNulty et al. (2015); Daver and Nakatsukasa (2015); Muteti et al. (2017) Proconsulidae - Ekembo heseloni Kulu Fm. (Rusinga Island) Kenya Early Miocene 16,0 16,0 Le Gros Clark (1952); Harrison (2010) Proconsulidae - Ekembo heseloni Mfangano Island (Mfangano Island) Kenya Early Miocene 18,3 18,3 Keith (1932); Le Gros Clark and Leakey (1951); Andrews (1978); Ruff et al. (1988); Harrison (2010); McNulty et al. (2015) Proconsulidae - Ekembo heseloni Kiahera Fm. (Rusinga Island) Kenya Early Miocene 18,5 18,5 Le Gros Clark (1952); Harrison (2010); McNulty et al. (2015) Proconsulidae - Ekembo nyanzae Kiahera Fm. (Rusinga Island) Kenya Early Miocene 18,5 18,5 Le Gros Clark and Leakey (1951); Le Gros Clark (1952); Harrison (2010); McNulty et al. (2015) Proconsulidae - Ekembo nyanzae Hiwegi Fm. (Rusinga Island) Kenya Early Miocene 18,3 18,3 MacInnes (1943); Le Gros Clark (1950, 1952); Le Gros Clark and Leakey (1950, 1951); Greenfield (1972, 1973); Whybrow and Andrews (1978); Harrison (1982; 2010); Walker and Pickford (1983); Beard et al. (1986); Teaford et al. (1988); Ruff et al. (1989); Ward (1991); Begun et al. (1994); Ward et al. (1995); Beynon et al. (1998); McNulty et al. (2015) Proconsulidae - Ekembo nyanzae Kulu Fm. (Rusinga Island) Kenya Early Miocene 16,0 16,0 Le Gros Clark (1952); Harrison (2010); McNulty et al. (2015) Proconsulidae - Ekembo nyanzae Mfangano Island (Mfangano Island) Kenya Early Miocene 18,3 18,3 Ruff et al. (1988); Ward (1993); Ward et al. (1993); Harrison (2010); McNulty et al. (2015) Proconsulidae - Proconsulidae indet. Bukwa II (Bukwa) Uganda Early Miocene 19,5 19,1 MacLatchy and Cote (2017); Pickford et al. (2017b) Nyanzapithecidae - Xenopithecus koruensis Locality 14 - Maize Crib (Koru-Muhoroni area) Kenya Early Miocene 20,0 19,0 Hopwood (1933a,b); Harrison (2010) Nyanzapithecidae - Mabokopithecus clarki Bed 3 (Maboko Island) Kenya Middle Miocene 15,3 15,3 von Koenigswald (1969); Pickford (1985a); Harrison (1986); Harrison (2010) Nyanzapithecidae - Rangwapithecus gordoni Redbed Member Unit 5 (Koru-Muhoroni area - Songhor) Kenya Early Miocene 20,0 18,5 Andrews (1970, 1978); Greenfield (1972); Harrison (1986, 2010); Kelley (1986); Nengo and Rae (1992); Gebo et al. (2009); Wuthrich et al. (2019); Patel et al. (2017) Nyanzapithecidae - Rangwapithecus gordoni Redbed Member Unit 6 (Koru-Muhoroni area - Songhor) Kenya Early Miocene 20,0 18,5 Harrison (2010); Cote et al. (2014); Hill et al. (2013) Nyanzapithecidae - Rangwapithecus gordoni Calcified Tuff Member (Lower Kapurtay) Kenya Early Miocene 20,0 18,5 Andrews (1970, 1974, 1978); Gebo et al. (2009); Harrison (2010) Nyanzapithecidae - Rangwapithecus gordoni LpM4 (Loperot) Kenya Early Miocene 17,0 17,0 Grossman et al. (2014) Nyanzapithecidae - Nyanzapithecus pickfordi Bed 3 (Maboko Island) Kenya Middle Miocene 15,3 15,3 Harrison (1986, 2010); McCrossin (1992) Nyanzapithecidae - Nyanzapithecus cf. pickfordi Muruyur - Kipsaraman (Baringo) Kenya Middle Miocene 15,8 15,8 Hill et al. (1986); Kelley et al. (2002); Pickford and Kunimatsu (2005); Harrison (2010) Nyanzapithecidae - Nyanzapithecus vancouveringorum Hiwegi Fm. (Rusinga Island) Kenya Early Miocene 18,3 18,3 Le Gros Clark and Leakey (1951); Andrews (1974, 1978); Harrison (2010) Nyanzapithecidae - Nyanzapithecus vancouveringorum Mfangano Island (Mfangano Island) Kenya Early Miocene 18,3 18,3 Andrews (1974, 1978); Kunimatsu (1992a); Harrison (2010) Nyanzapithecidae - Nyanzapithecus vancouveringorum Magare (Uyoma Peninsula) Kenya Early Miocene 18,3 18,3 McNulty et al. (2017) Nyanzapithecidae - Nyanzapithecus alesi North Napudet (Napudet) Kenya Middle Miocene 13,3 13,3 Nengo et al. (2017) Nyanzapithecidae - Nyanzapithecus harrisoni BG-X (Nachola) Kenya Middle Miocene 15,0 15,0 Pickford et al. (1987); Kunimatsu (1992b, 1997); Harrison (2010) Nyanzapithecidae - Turkanapithecus kalakolensis FJ-18 (Fejej) Ethiopia Early Miocene 17,5 17,0 Richmond et al. (1998); Harrison (2010) Nyanzapithecidae - Turkanapithecus kalakolensis Kalodirr Member (Losodok) Kenya Early Miocene 17,5 17,5 Leakey and Leakey (1986a); Leakey et al. (1988a); Harrison (2002, 2010); Rossie and Cote (2022) Nyanzapithecidae - Turkanapithecus rusingensis Hiwegi Fm. (Rusinga Island) Kenya Early Miocene 18,3 18,3 Pickford et al. (2010) Nyanzapithecidae - Turkanapithecus rusingensis Mfangano Island (Mfangano Island) Kenya Early Miocene 18,3 18,3 Pickford et al. (2010) Nyanzapithecidae - Turkanapithecus rusingensis Napak V (Akisim) Uganda Early Miocene 20,5 19,0 Pickford et al. (2010) Nyanzapithecidae - Turkanapithecus rusingensis Songhor (Koru-Muhoroni area) Kenya Early Miocene 20,3 20,3 Pickford et al. (2010) Nyanzapithecidae - Samburupithecus kiptalami Namurungule Fm. (Samburu Hills) Kenya Late Miocene 8,5 8,5 Ishida (1984); Ishida et al. (1984); Ishida and Pickford (1997); Harrison (2010) Nyanzapithecidae - Nyanzapithecidae indet. Ryskop (Ryskop) South Africa Early Miocene 16,0 16,0 Pickford and Senut (1997); Senut et al. (1997); Harrison (2010) Nyanzapithecidae - Nyanzapithecidae indet. Nakali (Nakali) Kenya Late Miocene 9,9 9,8 Kunimatsu et al. (2016, 2017) Nyanzapithecidae - Nyanzapithecidae indet. Fort Ternan (Fort Ternan) Kenya Middle Miocene 13,7 13,7 Leakey (1968a); Simons (1968); Harrison (1986, 1992, 2010); Pickford (1985b); Kelley (1986); Foecke et al. (2022) Nyanzapithecidae - Nyanzapithecidae indet. Kapsibor (Kapsibor) Kenya Middle Miocene 13,7 13,7 Harrison (1986a, 1992, 2010) Nyanzapithecidae - Nyanzapithecidae indet. Moroto II (Karamoja) Uganda Early Miocene 21,0 20,0 Jansma et al. (2015) Afropithecidae Afropithecinae Afropithecus turkanensis Buluk Member (Bakate Fm.) Kenya Early Miocene 17,2 17,2 Leakey and Walker (1985); Harrison (2010) Afropithecidae Afropithecinae Afropithecus turkanensis Kalodirr Member (Losodok) Kenya Early Miocene 17,5 17,5 Leakey and Walker (1985); Leakey and Leakey (1986b); Leakey et al. (1988b); Leakey and Walker (1997); Patel and Grossman (2006); Harrison (2010); Rossie and MacLatchy (2013); Rossie and Cote (2022) Afropithecidae Afropithecinae Afropithecus turkanensis Locherangan (Locherangan) Kenya Early Miocene 17,6 17,6 Andrews (1978); Anyonge (1991); Harrison (2010) Afropithecidae Afropithecinae Afropithecus turkanensis Moruorot (Losodok) Kenya Early Miocene 17,5 17,5 Andrews (1978); Leakey et al. (1988b); Harrison (2010) Afropithecidae Afropithecinae Heliopithecus leakeyi Ad Dabtiyah (Ad Dabtiyah) Saudi Arabia Early Miocene 16,0 16,0 Andrews et al. (1978); Andrews and Martin (1987); Harrison (2010) Afropithecidae Afropithecinae Morotopithecus bishopi Moroto I (Karamoja) Uganda Early Miocene 21,0 20,0 Bishop (1964); Gebo et al. (1997); MacLatchy and Pilbeam (1999); MacLatchy et al. (2000); Pickford et al. (1999, 2003); MacLatchy (2004); Young and MacLatchy (2004); Nakatsukasa (2008); Harrison (2010) Afropithecidae Afropithecinae Morotopithecus bishopi Moroto II (Karamoja) Uganda Early Miocene 21,0 20,0 Allbrook and Bishop (1963); Bishop (1964); Walker and Rose (1968); Pilbeam (1969a); Greenfield (1978); Ward and Kimbel (1983); Gebo et al. (1997); MacLatchy and Pilbeam (1999); MacLatchy et al. (2000, 2019); Senut et al. (2000); Pickford et al. (1999, 2003, 2017a, 2020); MacLatchy (2004); Young and MacLatchy (2004); Patel and Grossman (2006); Harrison (2010) Afropithecidae Afropithecinae cf. Afropithecinae Napak XXXII (Akisim) Uganda Early Miocene 20,1 20,1 Pickford et al. (2020) Afropithecidae Equatorinae Equatorius africanus Ombo (Ombo) Kenya Early Miocene 16,0 16,0 Le Gros Clark and Leakey (1950) Afropithecidae Equatorinae Equatorius africanus Bed 3 (Maboko Island) Kenya Middle Miocene 15,3 15,3 MacInnes (1943); Le Gros Clark and Leakey (1950, 1951); Leakey (1967, 1968b); Pilbeam (1969a); Greenfield (1972, 1973); Andrews (1978); Pickford (1985b); Kelley (1986); Allen and McCrossin (2008); Harrison (2010); Foecke et al. (2022) Afropithecidae Equatorinae Equatorius africanus BPRP 122 (Baringo) Kenya Middle Miocene 15,8 15,8 Hill et al. (1991); Brown et al. (1995); Ward et al. (1999); McCrossin and Benefit (1997); Kelley et al. (2002); Sherwood et al. (2002); Pickford and Kunimatsu (2005); Harrison (2010) Afropithecidae Equatorinae Equatorius africanus Kaloma (Kaloma) Kenya Middle Miocene 15,3 15,3 Pickford (1982); Harrison (2010) Afropithecidae Equatorinae Equatorius africanus Majiwa (Majiwa) Kenya Middle Miocene 15,3 15,3 Pickford (1982, 1985b); Harrison (2010) Afropithecidae Equatorinae Equatorius africanus Muruyur Fm. (Cheparawa) Kenya Middle Miocene 15,8 15,8 Pickford (1983); Pickford and Kunimatsu (2005); Harrison (2010) Afropithecidae Equatorinae Equatorius africanus Nyakach (Nyakach) Kenya Middle Miocene 15,0 15,0 Pickford (1985b); Harrison (2010) Afropithecidae Equatorinae Nacholapithecus kerioi BG-I (Nachola) Kenya Middle Miocene 15,0 15,0 Kunimatsu et al. (2019); Urciuoli et al. (2021) Afropithecidae Equatorinae Nacholapithecus kerioi BG-K (Nachola) Kenya Middle Miocene 15,0 15,0 Ishida (1984); Ishida et al. (1984, 1999, 2004); Pickford et al. (1987); Rose et al. (1996); Nakatsukasa et al. (1998; 2003a,b, 2007a,b, 2012); Takano et al. (2018, 2020); Kunimatsu et al. (2004a); Senut et al. (2004); Nakatsukasa and Kunimatsu (2009); Harrison (2010); Kikuchi et al. (2012, 2015, 2016, 2018); Ogihara et al. (2016); Pina et al. (2021) Afropithecidae Equatorinae Nacholapithecus kerioi BG-X (Nachola) Kenya Middle Miocene 15,0 15,0 Ishida et al. (1984); Harrison (2010) Afropithecidae Equatorinae cf. Nacholapithecus kerioi Moroto I (Karamoja) Uganda Early Miocene 21,0 20,0 Pickford et al. (2017a) but see MacLatchy (2019) Afropithecidae Equatorinae cf. Nacholapithecus kerioi Moroto II (Karamoja) Uganda Early Miocene 21,0 20,0 Pickford et al. (2017a) but see MacLatchy (2019) Afropithecidae Incertae sedis Otavipithecus namibiensis Berg Aukas (Berg Aukas) Namibia Middle Miocene 12,0 12,0 Conroy et al. (1992, 1993a,b, 1996); Conroy (1996); Pickford et al. (1997); Gommery (2000); Harrison (2010) Hylobatidae Yuanmoupithecinae Yuanmoupithecus xiaoyuan Leilao (Xiaohe Fm.) China Late Miocene 8,2 7,1 Pan (2006); Harrison et al. (2008); Ji et al. (2022) Hominidae Kenyapithecinae Griphopithecus suessi Kleinshadersdorf (Vienna Basin) Austria Middle Miocene 13,8 12,7 Ehrenberg (1938) Hominidae Kenyapithecinae Griphopithecus suessi Devínska Nová Ves – Sandberg (Devínska Nová Ves –Sandberg (= Neudorf SlovakiaSandberg)) Middle Miocene 13,6 13,1 Abel (1902); Remane (1921a,b); Glaessner (1931); Steininger (1967); Holec and Emry (2003) Hominidae Kenyapithecinae Griphopithecus alpani Çandir (Çandir) Turkey Middle Miocene 14,1 13,4 Tekkaya (1974); Greenfield (1979); Begun et al. (2003b); Güleç and Begun (2003) Hominidae Kenyapithecinae Griphopithecus alpani Paşalar (Paşalar) Turkey Middle Miocene 14,5 14,0 Andrews and Tobien (1977); Alpagut et al. (1990); Martin and Andrews (1993); Waddle et al. (1995); Kelley and Alpagut (1999); Ersoy et al. (2008); Gençturk et al. (2008); Humphrey and Andrews (2008); Mortzou and Andrews (2008) Hominidae Kenyapithecinae Kenyapithecus wickeri Fort Ternan (Fort Ternan) Kenya Middle Miocene 13,8 13,8 Leakey (1961, 1967, 1968a); Remane (1965); Walker and Andrews (1973); Andrews and Walker (1976); Andrews (1971; 1978); Greenfield (1974, 1978); Pickford (1985b); Harrison (1992, 2010); McCrossin and Benefit (1993) Hominidae Kenyapithecinae Kenyapithecus kizili Paşalar (Paşalar) Turkey Middle Miocene 14,5 14,0 Andrews and Tobien (1977); Alpagut et al. (1990); Martin and Andrews (1993); Waddle et al. (1995); Kelley and Alpagut (1999); Gençturk et al. (2008); Humphrey and Andrews (2008); Kelley (2008); Kelley et al. (2008) Hominidae Kenyapithecinae cf. Kenyapithecus sp. Berg Aukas (Berg Aukas) Namibia Middle miocene 12,0 12,0 Pickford and Senut (2010); Mocke et al. (2022) Hominidae Kenyapithecinae Kenyapithecinae indet. Thannhausen (Thannhausen) Germany Middle Miocene 14,0 14,0 Pickford (2013) Hominidae Kenyapithecinae cf. Kenyapithecinae Engelwiess (Engelswies) Germany Early Miocene 16,5 16,0 Heizmann (1992); Heizmann and Begun (2001) Hominidae Dryopithecinae Dryopithecus fontani ACM/C3-Ae (ACM) Spain Middle Miocene 11,9 11,9 Moyà-Solà et al. (2009a); Alba (2012) Hominidae Dryopithecinae Dryopithecus fontani ACM/C4-Ap (ACM) Spain Middle Miocene 11,9 11,9 Alba and Moyà-Solà (2012); Alba et al. (2017) Hominidae Dryopithecinae Dryopithecus fontani La Grive Saint-Alban M (Isère) France Middle Miocene 12,5 11,9 Depéret (1911); Pérez de los Ríos et al. (2013) Hominidae Dryopithecinae Dryopithecus fontani St. Stefan im Lavanttal (Gratkonr Basin) Austria Middle Miocene 12,2 12,0 Mottl (1957) Hominidae Dryopithecinae Dryopithecus fontani Can Mata s.l. (Can Mata) Spain Middle-Late Miocene 12,0 11,0 Crusafont Pairó and Hürzeler (1961); Van der Made and Ribot (1999); Alba et al. (2013, 2020, 2022); Marigó et al. (2014); Fortuny et al. (2021). Hominidae Dryopithecinae Dryopithecus fontani La Grive Saint-Alban L3/La Grive Saint-Alban L5? (Isère) France Middle-Late Miocene 11,9 11,2 Depéret (1911); Pérez de los Ríos et al. (2013) Hominidae Dryopithecinae Dryopithecus fontani St. Gaudens (Haute-Garonne) France Middle-Late Miocene 12,5 11,2 Lartet (1856); Gaudry (1890); Harlé (1898, 1899); Remane (1921a,b); Simons and Pilbeam (1965); Pilbeam and Simons (1971) Hominidae Dryopithecinae Dryopithecus cf. fontani ACM/C3-Az (ACM) Spain Middle Miocene 11,9 11,9 Moyà-Solà et al. (2009a); Alba (2012); Pina et al. (2019) Hominidae Dryopithecinae Pierolapithecus catalaunicus ACM/BCV1 (ACM) Spain Middle Miocene 12,0 12,0 Moyà-Solà et al. (2004, 2005b); Almécija et al. (2009); Alba et al. (2010a,b); Pérez de los Ríos et al. (2010, 2012a,b); Susanna et al. (2010); Alba (2012); Hammond et al. (2013); Pina et al. (2014, 2020); Fortuny et al. (2021) Hominidae Dryopithecinae Anoiapithecus brevirostris ACM/C1-E* (ACM) Spain Middle Miocene 12,4 12,3 Alba et al. (2013); Fortuny et al. (2021) Hominidae Dryopithecinae Anoiapithecus brevirostris ACM/C3-Aj (ACM) Spain Middle Miocene 12,0 12,0 Moyà-Solà et al. (2009b); Alba et al. (2010b, 2013); Pérez de los Ríos et al. (2010, 2012a); Alba (2012); Fortuny et al. (2021) Hominidae Dryopithecinae Anoiapithecus brevirostris ACM/C4-Cp (ACM) Spain Middle Miocene 12,0 12,0 Alba et al. (2017) Hominidae Dryopithecinae Hispanopithecus laietanus Can Llobateres 1 (Can Llobateres) Spain Late Miocene 9,8 9,8 Crusafont Pairó (1958, 1965); Crusafont Pairó and Hürzeler (1961, 1969); Crusafont Pairó and Golpe-Posse (1974); Golpe-Posse (1982, 1993); Begun et al. (1990); Harrison (1991a); Ribot et al. (1996); McNulty (2005); Alba et al. (2010a, 2012a,b); Pina et al. (2012); Fortuny et al. (2021); Urciuoli et al. (2021) Hominidae Dryopithecinae Hispanopithecus laietanus Can Llobateres 2 (Can Llobateres) Spain Late Miocene 9,6 9,6 Moyà-Solà and Köhler (1993, 1995, 1996); Köhler et al. (2001, 2002); Almécija et al. (2007); Tallman et al. (2013); Susanna et al. (2014); Fortuny et al. (2021) Hominidae Dryopithecinae Hispanopithecus laietanus La Tarumba 1 (La Tarumba) Spain Late Miocene 9,6 9,6 Villalta Comella and Crusafont Pairó (1944); Crusafont Pairó (1965); Crusafont Pairó and Hürzeler (1969); Crusafont Pairó and Golpe-Posse (1974); Golpe-Posse (1982, 1993); Harrison (1991a); Fortuny et al. (2021) Hominidae Dryopithecinae Hispanopithecus laietanus Can Feu 1 (Can Feu) Spain Late Miocene 10,0 9,7 Alba et al. (2012b) Hominidae Dryopithecinae Hispanopithecus cf. laietanus Can Pallars i Llobateres (Can Pallars i Llobateres) Spain Late Miocene 10,0 9,7 Golpe-Posse (1982); Alba et al. (2018); Fortuny et al. (2021) Hominidae Dryopithecinae Hispanopithecus crusafonti Can Poncic 1 (Can Poncic) Spain Late Miocene 10,3 10,0 Crusafont Pairó and Hürzeler (1961, 1969); Crusafont Pairó (1965); Crusafont-Pairò and Golpe-Posse (1973, 1974); Golpe-Posse (1982); Begun (1992b); Harrison (1992); Ribot et al. (1996) Hominidae Dryopithecinae Hispanopithecus crusafonti Teuleria del Firal (Teuleria del Firal) Spain Late Miocene 10,3 10,0 Vidal (1913a,b, 1914); Woodward (1914); Crusafont Pairó and Hürzeler (1969); Crusafont Pairó and Golpe-Posse (1974); Begun (1992b); Ribot et al. (1996); Fortuny et al. (2021) Hominidae Dryopithecinae Hispanopithecus sp. EDAR13 (EDAR) Spain Late Miocene 10,3 10,0 Checa Soler and Rius Font (2003); Alba et al. (2018) Hominidae Dryopithecinae Hispanopithecus sp. Polinyà 2 (Polinyà) Spain Late Miocene 11,6 9,7 Crusafont-Pairò and Golpe-Posse (1973, 1974); Golpe-Posse (1982, 1993); Harrison (1991a); Ribot et al. (1996) Hominidae Dryopithecinae Rudapithecus hungaricus Rudabánya 2 (Pannonian Basin) Hungary Late Miocene 10,0 9,8 Schlosser (1901); Kretzoi (1969, 1975); Morbeck (1983); Kordos (1987, 1988); Begun (1988, 1992a, 1993, 1994, 2002, 2009, 2015); Begun and Kordos (1993); Greenfield (1979); Kordos and Begun (1997, 2001a,b); McNulty (2005); Kivell and Begun (2009); Begun et al. (2012); Ryan et al. (2012); Kullmer et al. (2013); Shapiro and Begun (2016); Smith et al. (2019); Ward et al. (2019); Urciuoli et al. (2021); Gunz et al. (2020) Hominidae Dryopithecinae Danuvius guggenmosi Hammerschmiede 5 (Pforzen) Germany Late Miocene 11,6 11,6 Böhme et al. (2019) Hominidae Dryopithecinae ?Udabnopithecus garedziensis Udabno 1 (Udabno) Georgia Late Miocene 8,1 7,7 Burchak-Abramovich and Gabachvili (1945); Szalay and Delson (1979); Andrews et al. (1996); Gabunia et al. (2001); Agustí et al. (2020) Hominidae Dryopithecinae Dryopithecinae indet. Castell de Barberà (Castell de Barberà) Spain Late Miocene 11,2 11,1 Crusafont Pairó and Hürzeler (1969); Crusafont Pairó and Golpe-Posse (1974); Moyà Solà et al. (1990); Harrison (1991a); Ribot et al. (1996); Alba et al. (2011); Almécija et al. (2012) Hominidae Dryopithecinae Dryopithecinae indet. Can Mata 1 (Can Mata) Spain Late Miocene 11,6 11,2 Crusafont-Pairò and Golpe-Posse (1973); Golpe-Posse (1982, 1993); Harrison (1991a); Alba (2012) Hominidae Dryopithecinae Dryopithecinae indet. Ebingen (Swabian Alb) Germany Late Miocene 11,0 9,0 Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. Melchingen (Swabian Alb) Germany Late Miocene 11,2 7,5 Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. Melchingen (Swabian Alb) Germany Late Miocene 11,2 7,5 Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. Neuhausen (Swabian Alb) Germany Late Miocene 11,0 9,0 Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. Neuhausen (Swabian Alb) Germany Late Miocene 11,0 9,0 Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. Neuhausen (Swabian Alb) Germany Late Miocene 11,0 9,0 Jäger (1839); Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. Trochtelfingen (Swabian Alb) Germany Late Miocene 11,0 9,0 Pickford (2012) Hominidae Dryopithecinae Dryopithecinae indet. ACM/C5-C3 (ACM) Spain Middle Miocene 11,9 11,9 Alba et al. (2017) Hominidae Dryopithecinae Dryopithecinae indet. ACM/C5-D1 (ACM) Spain Middle Miocene 11,7 11,7 Alba et al. (2017) Hominidae Dryopithecinae Dryopithecinae indet. ACM/C8-B* (ACM) Spain Middle Miocene 11,9 11,9 Alba et al. (2017) Hominidae Dryopithecinae Dryopithecinae indet. Wissberg (Gau-Weinheim) Germany Middle-Late Miocene 13,7 7,5 Pickford (2012) Hominidae Dryopithecinae Neopithecus brancoi nomem dubium Salmendingen (Swabian Alb) Germany Late Miocene 11,6 7,5 Abel (1902); Remane (1921a,b); Begun and Kordos (1993); Moyà-Solà et al. (2009a); Pickford (2012); Begun (2015) Hominidae Dryopithecinae 'Sivapithecus ' occidentalis species inquirenda ACM/BCV4 (ACM) Spain Middle Miocene 11,9 11,9 Alba et al. (2020) Hominidae Dryopithecinae 'Sivapithecus ' occidentalis species inquirenda Can Vila (Can Vila) Spain Middle Miocene 12,0 12,0 Villalta Comella and Crusafont Pairó (1941); Crusafont Pairó (1965); Crusafont Pairó and Hürzeler (1969); Crusafont Pairó and Golpe-Posse (1974); Golpe-Posse (1982); Harrison (1991); Alba et al. (2020); Fortuny et al. (2021) Hominidae Ponginae Khoratpithecus chiangmuanensis Chiang Muan - Upper Lignite (Nakhon Ratchasima Province, Khorat) Thailand Middle Miocene 12,4 12,2 Chaimanee et al. (2003); Kunimatsu et al. (2004b, 2005a,b); Pickford et al. (2004) Hominidae Ponginae Khoratpithecus piriyai Somsak (Nakhon Ratchasima Province, Khorat) Thailand Late Miocene 9,0 6,0 Chaimanee et al. (2003, 2004, 2006) Hominidae Ponginae Khoratpithecus piriyai Lower Sand Unit (Nakhon Ratchasima Province, Khorat) Thailand Late Miocene 9,0 6,0 Chaimanee et al. (2019, 2022) Hominidae Ponginae Khoratpithecus cf. piriayi YS1 (Irrawaddy Fm.) Myanmar Late Miocene 9,0 6,0 Jaeger et al. (2011) Hominidae Ponginae Khoratpithecus ayeyarwadyensis YS1 ? (Irrawaddy Fm.) Myanmar Late Miocene 10,4 8,8 Jaeger et al. (2011) Hominidae Ponginae Khoratpithecus cf. ayeyarwadyensis SMG1 (Irrawaddy Fm.) Myanmar Late Miocene 10,0 8,0 Takai et al. (2021) Hominidae Ponginae Khoratpithecus magnus Lower Sand Unit (Nakhon Ratchasima Province, Khorat) Thailand Late Miocene 9,0 6,0 Chaimanee et al. (2022) Hominidae Ponginae Ankarapithecus meteai Sinap Locality 12 (Sinap Formation) Turkey Late Miocene 9,8 9,8 Ozansoy (1957, 1965); Andrews and Tekkaya (1980); McHenry et al. (1980); Alpagut et al. (1996); Begun and Güleç (1998 ); Kappelman et al. (2003) Hominidae Ponginae Lufengpithecus lufengensis Shihuiba 75033 D2-6 (Lufeng) China Late Miocene 6,9 6,2 Xu et al. (1978); Xu and Lu (1979, 1980, 2008); Lu et al. (1981); Xiao (1981); Wu et al. (1981, 1982, 1983, 1984, 1985, 1986); Wu and Oxnard (1983a,b); Wu (1984, 1987); Wu and Xu (1985); Lin et al. (1987); Wood and Xu (1991); Zhang et al. (1993); Harrison et al. (2002); Harrison (2006); Zhao et al. (2008) Hominidae Ponginae Lufengpithecus cf. lufengensis Shuitangba (Shuitangba) China Late Miocene 6,2 6,2 Ji et al. (2013) Hominidae Ponginae Sivapithecus sivalensis Jabbi (Potwar Plateau) Pakistan Late Miocene 10,4 8,5 Lydekker (1879); Remane (1921a,b); Kay (1982); Ward and Brown (1986); Kelley (2005a) Hominidae Ponginae Sivapithecus sivalensis Potwar Plateau [Dhok Pathan] (Potwar Plateau) Pakistan Late Miocene 9,8 8,5 Brown et al. (1924); Pandey and Sastri (1968); Greenfield (1979); Kay (1982); Ward and Brown (1986); Kelley (2005b) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0182 (Gandakas) Pakistan Late Miocene 9,3 9,2 Pilbeam et al. (1980); Kay (1982); Ward and Pilbeam (1983); Ward and Brown (1986); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0224 (Dinga Kas) Pakistan Late Miocene 9,4 9,4 Lewis (1934); Pilbeam et al. (1980); Kay (1982); Ward and Brown (1986); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0226 (Dinga Kas) Pakistan Late Miocene 9,4 9,4 Pilbeam et al. (1980); Kay (1982); Ward and Brown (1986); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0260 (Kaulial Kas) Pakistan Late Miocene 9,2 9,2 Pilbeam et al. (1980); Kay (1982); Ward and Brown (1986); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0317 (Choutriwali Kas) Pakistan Late Miocene 9,3 9,2 Pilbeam et al. (1980); Kay (1982); Ward and Pilbeam (1983); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0350 (269 Section) Pakistan Late Miocene 9,4 9,4 Pilbeam et al. (1980); Kay (1982); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0409 (Kaulial Kas) Pakistan Late Miocene 9,3 9,2 Pilbeam et al. (1980); Kay (1982); Ward and Brown (1986); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0410 (Kaulial Kas) Pakistan Late Miocene 9,4 9,4 Pilbeam (1982); Preuss (1982); Ward and Kimbel (1983); Ward and Pilbeam (1983); Ward and Brown (1986); McNulty (2005); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0463 (Kaulial Kas) Pakistan Late Miocene 9,3 9,2 Pilbeam et al. (1980); Kay (1982); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0137 (Gandakas Road) Pakistan Late Miocene 9,3 9,2 Pilbeam et al. (1980); Ward and Brown (1986); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0191 (Gandakas) Pakistan Late Miocene 9,4 9,2 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0207 (Malhuwala Kas) Pakistan Late Miocene 9,4 9,4 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0211 (Dinga Kas) Pakistan Late Miocene 9,3 9,3 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0221 (Dinga Kas) Pakistan Late Miocene 9,4 9,3 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0227 (227 Section) Pakistan Late Miocene 9,4 9,4 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0230 (227 Section) Pakistan Late Miocene 9,4 9,4 Pilbeam et al. (1980); Kay (1982); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0309 (Dhok Mila, Khaur Kas) Pakistan Late Miocene 9,4 9,4 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0310 (Dhok Mila, Khaur Kas) Pakistan Late Miocene 9,4 9,3 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0314 (Dhok Mila, Khaur Kas) Pakistan Late Miocene 9,3 9,3 Pilbeam et al. (1980); Ward and Pilbeam (1983); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0327 (Kot Maliaran) Pakistan Late Miocene 9,3 9,2 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0414 (Kaulial Kas) Pakistan Late Miocene 9,3 9,3 Pilbeam et al. (1980); Ward and Brown (1986); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0416 (Kaulial Kas) Pakistan Late Miocene 9,4 9,3 Pilbeam et al. (1980); Ward and Brown (1986); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0442 (Kaulial Kas) Pakistan Late Miocene 8,6 8,5 Pilbeam et al. (1980); Ward and Brown (1986); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0604 (Kaulial Kas) Pakistan Late Miocene 8,6 8,5 Barry (1986); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0328 (Hasal Kas, Bora Kas) Pakistan Late Miocene 9,4 9,4 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0885 (?) Pakistan - ? ? Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0891 (Kaulial Kas) Pakistan Late Miocene 8,8 8,7 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0990 (?) Pakistan - ? ? Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0997 (Kaulial Kas) Pakistan Late Miocene 9,3 9,2 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y1002 (Kaulial Kas) Pakistan Late Miocene 9,4 9,4 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y1004 (Kaulial Kas) Pakistan Late Miocene 9,4 9,3 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0261 (Kaulial Kas) Pakistan Late Miocene 9,6 9,6 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0251 (Ratha Kas) Pakistan Late Miocene 10,0 10,0 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks L0094 (Dhala Nala) Pakistan Late Miocene 10,3 10,1 Lewis (1934, 1937); Jukar and Brinkman (2021); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks KL0001 (Kaulial Kas) Pakistan Late Miocene 9,3 9,2 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sivalensis Siwaliks Y0158 (=Y0388) (Malhuwala Kas) Pakistan Late Miocene 8,8 8,7 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus cf. sivalensis Haritalyangar (Haritalyangar) India Late Miocene 9,2 8,1 Pilgrim (1927); Gregory et al. (1938); Wadia and Aiyengar (1938); Prasad (1962, 1964, 1969); Khatri (1964, 1975); Simons and Chopra (1969b); Simons and Pilbeam (1971); Greenfield (1974); Chopra (1983); Johnson et al. (1983); Kay (1982); Ward and Brown (1986); Kelley (2005b); Pickford (2010); Pickford and Tiwari (2010) Hominidae Ponginae Sivapithecus cf. sivalensis Chakrana (Haritalyangar) India Late Miocene 9,2 8,1 Simons (1961); Khatri (1975); Kay (1982); Ward and Brown (1986); Kelley (2005b) Hominidae Ponginae Sivapithecus cf. sivalensis Bandal [Nagri Beds] (Bandal) India Late Miocene 10,1 8,6 Pandey and Sastri (1968); Kay (1982); Kelley (2005b); Pickford (2010) Hominidae Ponginae Sivapithecus cf. sivalensis Bharari (Haritalyangar) India Late Miocene 8,1 7,5 Sankhyan (1985); Pickford and Tiwari (2010) Hominidae Ponginae Sivapithecus cf. sivalensis VPL-1 (Hari Mandar Dhar) India Late Miocene 9,0 9,0 Ward and Brown (1986); Cameron et al. (1997); Patnaik and Cameron (1997); Patnaik et al. (2005) Hominidae Ponginae Sivapithecus cf. sivalensis Hari Devi 1 (HD1) (Haritalyangar) India Late Miocene 8,9 8,9 Patnaik et al. (2005); Pillans et al. (2005) Hominidae Ponginae Sivapithecus indicus Rashole 3 (Ramnagar) India Middle Miocene 13,8 12,5 Gilbert et al. (2019); Gilbert et al. (2020) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0083 (Hutch Nala) Pakistan Late Miocene 11,6 11,6 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0499 (Kanatti Upper) Pakistan Middle Miocene 12,1 12,1 Gregory et al. (1938); Raza et al. (1983); Rose (1989); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0500 (Kanatti Upper) Pakistan Late Miocene 12,1 12,1 Raza et al. (1983); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0663 (Parrawali) Pakistan Late Miocene 11,6 11,6 Morgan et al. (2015); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0076 (Gambhir Section) Pakistan Late Miocene 11,5 11,4 Raza et al. (1983); Pilbeam et al. (1990); Kappelman et al. (1991); DeSilva et al. (2010); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Potwar Plateau [Chinji] (Potwar Plateau) Pakistan Middle-Late Miocene 12,7 11,2 Pilgrim (1910, 1915); Gregory et al. (1938); Wadia and Aiyengar (1938); Simons and Pilbeam (1965); Kay (1982); Ward and Brown (1986); Ward (1997); Kelley (2002, 2005a,b) Hominidae Ponginae Sivapithecus indicus Ramnagar (Ramnagar) India Middle Miocene 13,8 12,5 Brown et al. (1924); Pilgrim (1927); Gregory et al. (1938); Wadia and Aiyengar (1938); Khatri (1975); Greenfield (1978, 1979); Kay (1982); Chopra (1983); Ward and Brown (1986); Kelley (2005b) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0494 (Gambhir Section) Pakistan Middle Miocene 12,4 12,4 Kay (1982); Raza et al. (1983); Ward and Pilbeam (1983); Kappelman et al. (1991); Kelley (2005b); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0495 (Gambhir Section) Pakistan Late Miocene 11,6 11,6 Raza et al. (1983); Rose (1984); Kelley (2005b); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0496 (Gambhir Section) Pakistan Middle Miocene 12,4 12,4 Raza et al. (1983); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0498 (Hutch Nala) Pakistan Middle Miocene 11,7 11,7 Kay (1982); Raza et al. (1983); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0647 (Gambhir Section) Pakistan Middle Miocene 12,4 12,4 Raza et al. (1983); Kappelman et al. (1991); Morgan et al. (2015); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0750 (Samriala Nala) Pakistan Middle Miocene 12,8 12,7 Raza et al. (1983); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0767 (Gambhir Section) Pakistan Middle Miocene 12,4 12,3 Raza et al. (1983); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0775 (Rata Dala Nala) Pakistan Middle Miocene 12,4 12,2 Raza et al. (1983); Kappelman et al. (1991); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Dalsar (Ramnagar) India Middle Miocene 13,8 12,5 Sehgal and Patnaik (2012) Hominidae Ponginae Sivapithecus indicus Kundal Nala [Chinji] (Potwar Plateau) Pakistan Late Miocene 11,5 11,4 Pilgrim (1915); Remane (1921a,b); Gregory et al. (1938); Kay (1982); Kelley (1988; 2002; 2005a) Hominidae Ponginae Sivapithecus indicus Siwaliks L0021 (Hutch Nala) Pakistan Late Miocene 11,4 11,4 Lewis (1934); Barry (1986); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0829 (Samriala Nala) Pakistan Middle-Late Miocene 11,9 11,1 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks L0024 (Hutch Nala) Pakistan Middle Miocene 12,4 12,2 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks Y0502 (Hutch Nala) Pakistan Middle Miocene 12,0 12,0 Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus indicus Siwaliks B0145 (?) Pakistan - ? ? Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus cf. indicus Siwaliks L0081 (Andar Kas) Pakistan Late Miocene 11,2 10,8 Lewis (1934, 1937); Barry (1986); Jukar and Brinkman (2021) [Sivapithecus sp. according to Barry et al., pers. comm., this citation corresponds to the holotype of Bramapithecus thorpei Lewis, 1934] Hominidae Ponginae Sivapithecus parvada Siwaliks Y0311 (Gambhir Section) Pakistan Late Miocene 10,1 10,0 Lydekker (1879); Pilgrim (1910); Pilbeam et al. (1977; 1980, 1990); Dehm (1983); von Koenigswald (1983); Kelley and Pilbeam (1986a,b); Rose (1986); Kelley (1988, 2002, 2005a,b); Spoor et al. (1991); Kelley et al. (1995); Ward (1997); Madar et al. (2002); Pickford (2010); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sp. Siwaliks B0009 (Hasnot) Pakistan Late Miocene 11,2 10,8 Brown et al. (1924); Gregory and Hellman (1926); Simons (1964); Greenfield (1974, 1978, 1979); Kay (1982); Ward and Brown (1986); Kelley (2005a); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sp. Dhara Reserve (Uttar Pradesh) India Middle-Late Miocene 12,7 11,2 Sahni et al. (1983); Tiwari and Kumar (1984); Cameron et al. (1999); Pickford and Tiwari (2010) Hominidae Ponginae Sivapithecus sp. Siwaliks L0035 (Haritalyangar) India Late Miocene ? ? Lewis (1934, 1937); Simons (1964); Greenfield (1974, 1979); Khatri (1975); Chopra (1978); Kay (1982); Barry (1986); Kelley (1988); Jukar and Brinkman (2021); Barry et al. (pers. comm.) [this citation includes the holotypes of Dryopithecus sivalensis Lewis, 1934 and Ramapithecus hariensis Lewis, 1934] Hominidae Ponginae Sivapithecus sp. Tinau Khola (Dang Valley) Nepal Late Miocene 9,5 9,0 Munthe et al. (1983) Hominidae Ponginae Sivapithecus sp. Tapar (Kutch) India Late Miocene 10,8 10,0 Bhandari et al. (2018) Hominidae Ponginae Sivapithecus sp. Siwaliks Y0259 (Kaulial Kas) Pakistan Late Miocene 10,5 10,4 Pilbeam et al. (1980); Barry et al. (pers. comm.) Hominidae Ponginae Sivapithecus sp. Dera Gopipur (Himachal Pradesh) India - ? ? Kaul and Vasishat (1981); Gupta et al. (1982); Kay (1982); Pickford and Tiwari (2010) Hominidae Ponginae Sivapithecus sp. Domeli (Potwar Plateau) Pakistan Late Miocene 11,5 9,0 Pilbeam (1969b); Greenfield (1974); Kay (1982); Ward and Brown (1986); Kelley (1988; 2002; 2005a) Hominidae Ponginae Sivapithecus sp. Siwaliks L0039 (Haritalyangar) India - ? ? Lewis (1934); Simons (1964); Greenfield (1974, 1979); Kay (1982); Chopra (1978, 1983); Ward and Kimbel (1983); Ward and Pilbeam (1983); Bhandari et al. (2018); Jukar and Brinkman (2021); Barry et al. (pers. comm.) [this citation corresponds to the holotype of Ramapithecus brevirostris Lewis, 1934] Hominidae Ponginae Sivapithecus sp. Siwaliks L0040 (Haritalyangar) India - ? ? Lewis (1936, 1937); Greenfield (1974, 1979); Kelley (1988); Jukar and Brinkman (2021); Barry et al. (pers. comm.) [this citation corresponds to the holotype of Sugrivapithecus gregoryi Lewis, 1936] Hominidae Ponginae Sivapithecus sp. Siwaliks S0008 (?) Pakistan - ? ? Barry et al. (pers. comm.) Hominidae Ponginae Indopithecus giganteus Chandigarh (Haritalyangar) India Late Miocene 8,6 8,6 Simons and Chopra (1969a,b); Khatri (1975); Chopra (1978, 1983) Hominidae Ponginae Indopithecus giganteus HD1 (Haritalyangar) India Late Miocene 8,9 8,9 Patnaik et al. (2005); Pillans et al. (2005) Hominidae Ponginae Indopithecus giganteus Alipur (Alipur) India Late Miocene 8,6 8,6 Pilgrim (1915); Remane (1921a,b); von Koenigswald (1949); Prasad (1969); Szalay and Delson (1979); Cameron (2001) Hominidae Homininae Chororapithecus abyssinicus Chorora Fm. (Beticha) Ethiopia Late Miocene 8,0 8,0 Suwa et al. (2007); Harrison (2010) Hominidae Homininae Nakalipithecus nakayamai Nakali (Nakali) Kenya Late Miocene 9,9 9,8 Kunimatsu et al. (2007); Nakatsukasa (2009); Harrison (2010); Morita et al. (2017) Hominidae Homininae Homininae indet. Kapsomin (Lukeino) Kenya Late Miocene 6,2 5,7 Hill et al. (1986); Senut et al. (2001); Pickford and Senut (2005); Harrison (2010) Hominidae Homininae Homininae indet. Cheboit (Lukeino) Kenya Late Miocene 6,2 5,7 Pickford (1975); Hill et al. (1986); Hill and Ward (1988); Senut et al. (2001); Pickford and Senut (2005); Harrison (2010) Hominidae Incertae sedis Graecopithecus freybergi Pikermi Fm. - Red Conglomerate Member (Pyrgos) Greece Late Miocene 7,2 7,2 Freyberg (1951); von Koenigswald (1972); Martin and Andrews (1984); McNulty (2005); Fuss et al. (2017) Hominidae Incertae sedis Ouranopithecus macedoniensis Nikiti 1 (Chalkidiki Peninsula) Greece Late Miocene 9,7 8,8 Koufos and de Bonis (2006) Hominidae Incertae sedis Ouranopithecus macedoniensis Ravin de la Pluie (Axios Valley) Greece Late Miocene 9,4 9,3 Arambourg and Piveteau (1929); de Bonis (1974); de Bonis et al. (1975, 1992, 1998); de Bonis and Melentis (1977a,b, 1978); de Bonis and Koufos (1993, 2014); Koufos and de Bonis (2004, 2006, 2017); Smith et al. (2004); Macchiarelli et al. (2009); Koufos et al. (2016) Hominidae Incertae sedis Ouranopithecus macedoniensis Xirochori 1 (Axios Valley) Greece Late Miocene 9,7 9,7 de Bonis et al. (1990, 1992); de Bonis and Koufos (1993); Koufos and de Bonis (2006) Hominidae Incertae sedis Ouranopithecus turkae Çorakyerle (Çorakyerle) Turkey Late Miocene 8,1 7,6 de Bonis (1974); de Bonis and Melentis (1977a); Begun et al. (2003a); Güleç et al. (2007); Begun (2009) Hominidae Incertae sedis Graecopithecini indet. Azmaka 6 (Azmaka) Bulgaria Late Miocene 7,2 7,2 Spassov et al. (2012) Hominidae Incertae sedis Sinopithecus keiyuanensis Xiaolungtan (Kaiyuan) China Middle-Late Miocene 12,5 11,6 Woo (1957, 1958a,b); Xu et al. (1978); Zhang et al. (1983, 1987, 1993); Zhang (1987); Wu et al. (1989); Pan (1994); Wu and Poirier (1995); Harrison et al. (2002); Harrison (2006) Hominidae Incertae sedis Sinopithecus hudienensis Baozidongqing (Xiaohe Fm.) China Late Miocene 8,2 8,1 Jiang et al. (1987); Zhang et al. (1993); Pan (1996); Zheng and Zhang (1997); Liu et al. (1999, 2000); Qi et al. (2006); Zheng (2006) Hominidae Incertae sedis Sinopithecus hudienensis Fangbeiliangzi (Xiaohe Fm.) China Late Miocene 8,2 8,1 Jiang et al. (1987); Zhang et al. (1993); Pan (1996); Zheng and Zhang (1997); Liu et al. (1999, 2000); Qi et al. (2006); Zheng (2006) Hominidae Incertae sedis Sinopithecus hudienensis Hudieliangzi (Xiaohe Fm.) China Late Miocene 8,2 8,1 Jiang et al. (1987); Zhang et al. (1987, 1988, 1993); Ho (1988, 1990); Wu and Poirier (1995); Pan (1996); Zheng and Zhang (1997); Liu et al. (1999, 2000); Harrison et al. (2002); Harrison (2006); Qi et al. (2006); Zheng (2006); Kelley and Gao (2012) Hominidae Incertae sedis Sinopithecus hudienensis Leilao (Xiaohe Fm.) China Late Miocene 8,2 7,1 Jiang et al. (1987); Ho (1988, 1990); Zhang et al. (1993); Wu and Poirier (1995); Pan (1996); Zheng and Zhang (1997); Liu et al. (1999, 2000); Harrison et al. (2002); Harrison (2006); Qi et al. (2006); Zheng (2006); Zhang et al. (2020) Hominidae Incertae sedis Sinopithecus hudienensis Zhupeng (Xiaohe Fm.) China Late Miocene 7,4 7,2 Jiang et al. (1987); Qi et al. (2006) Hominidae - Hominidae indet. Maragheh - Middle Pumice (Maragheh) Iran Late Miocene 7,5 7,5 Suwa et al. (2016) Hominidae - Hominidae indet. BPRP 38 (Ngorora) Kenya Middle Miocene 12,8 12,8 Hill et al. (2002); Harrison (2010) Hominidae - Hominidae indet. BPRP 60 (Ngorora) Kenya Middle Miocene 12,0 12,0 Bishop and Chapman (1970); Hill and Ward (1988); Hill et al. (2002); Harrison (2010) Hominidae - Hominidae indet. BPRP 65 (Ngorora) Kenya Middle Miocene 12,8 12,8 Bishop and Chapman (1970); Bishop and Pickford (1975); Hill et al. (1986, 2002); Hill and Ward (1988); Pickford and Ishida (1998); Pickford and Senut (2005); Harrison (2010) Incertae sedis - Oreopithecus bambolii Fiume Santo (Fiume Santo) Italy Late Miocene 8,1 6,7 Cordy and Ginesu (1994); Rook et al. (2006); Abbazzi et al. (2008) Incertae sedis - Oreopithecus bambolii V1 (Baccinello) Italy Late Miocene 8,3 8,1 Gervais (1872a,b); Forsyth Major (1880); Schlosser (1887); Ristori (1890); Ottolenghi (1898); Merciai (1907); Schwalbe (1915); Hürzeler (1949, 1951, 1954, 1958, 1968); Butler and Mills (1950); Heberer (1952); Schultz (1960); Simons (1960); Straus (1960, 1963); Knussmann (1967); Berzi (1972); Szalay and Langdon (1986); Harrison (1987; 1991b); Jungers (1987); Sarmiento (1987); Clarke (1997); Harrison and Rook (1997); Köhler and Moyà-Solà (1997); Moyà-Solà and Köhler (1997); Moyà-Solà et al. (1999, 2005a); Rook et al. (1999, 2004); Susman (2004); Almécija et al. (2014); Hammond et al. (2020); Urciuoli et al. (2020) Incertae sedis - Oreopithecus bambolii V2 (Baccinello) Italy Late Miocene 8,1 6,7 Rook (1993); Rook et al. (1996) Incertae sedis - 'Dryopithecus' wuduensis Wudu (Gansu) China Late Miocene 8,3 8,3 Xue and Delson (1979) - - Hominoidea indet. Moruorot (Losodok) Kenya Early Miocene 17,5 17,5 Andrews (1978); Anyonge (1991) - - Hominoidea indet. N 885 (N 885) Niger Late Miocene 11,0 8,0 Pickford et al. (2008, 2009b); Harrison (2010); Mocke et al. (2022) a When indicated as 'Barry et al. (pers. comm.)", identifications of Siwalik hominoid remains checked or updated based on data kindly provided by John Barry, Larry Flynn, Jay Kelley, Michèle Morgan, and David Pilbeam on September 2022. SOM Table S2 Miocene ape-bearing localities (in alphabetical order), including site/geographic area, country, age range, subepoch, and citations for the age. a Locality Site Continent Country Maximum age (Ma) Minimum age (Ma) Subepoch Citations ACM/BCV1 ACM Europe Spain 12,0 12,0 Middle Miocene Alba et al. (2017) ACM/BCV4 ACM Europe Spain 11,9 11,9 Middle Miocene Alba et al. (2017) ACM/C1-E* ACM Europe Spain 12,3 12,4 Middle Miocene Alba et al. (2017) ACM/C3-Ae ACM Europe Spain 11,9 11,9 Middle Miocene Alba et al. (2017) ACM/C3-Aj ACM Europe Spain 12,0 12,0 Middle Miocene Alba et al. (2017) ACM/C3-Az ACM Europe Spain 11,9 11,9 Middle Miocene Alba et al. (2017) ACM/C4-Ap ACM Europe Spain 11,9 11,9 Middle Miocene Alba et al. (2017) ACM/C4-Cp ACM Europe Spain 12,0 12,0 Middle Miocene Alba et al. (2017) ACM/C5-C3 ACM Europe Spain 11,9 11,9 Middle Miocene Alba et al. (2017) ACM/C5-D1 ACM Europe Spain 11,6 11,6 Late Miocene Alba et al. (2017) ACM/C8-B* ACM Europe Spain 11,7 11,7 Middle Miocene Alba et al. (2017) Ad Dabtiyah Ad Dabtiyah Africa Saudi Arabia 16,0 16,0 Early Miocene Van Couvering and Delson (2020) Alipur Alipur Asia India 8,6 8,6 Late Miocene Pillans et al. (2005) Azmaka 6 Azmaka Europe Bulgaria 7,2 7,2 Late Miocene Böhme et al. (2017) Bandal Bandal Asia India 10,1 8,6 Late Miocene Pickford (2010) Baozidongqing Xiaohe Fm. Asia China 8,2 8,1 Late Miocene Woodburne et al. (2013) Bed 3 Maboko Island Africa Kenya 15,3 15,3 Middle Miocene Van Couvering and Delson (2020) Berg Aukas Berg Aukas Africa Namibia 12,0 12,0 Middle Miocene Conroy et al. (1992) BG-I Nachola Africa Kenya 15,0 15,0 Middle Miocene Van Couvering and Delson (2020) BG-K Nachola Africa Kenya 15,0 15,0 Middle Miocene Van Couvering and Delson (2020) BG-X Nachola Africa Kenya 15,0 15,0 Middle Miocene Van Couvering and Delson (2020) Bharari Haritalyangar Asia India 8,1 7,5 Late Miocene Pillans et al. (2005); Pickford and Tiwari (2010) BPRP 122 Baringo Africa Kenya 15,8 15,8 Middle Miocene Van Couvering and Delson (2020) BPRP 38 Ngorora Africa Kenya 12,8 12,8 Middle Miocene Van Couvering and Delson (2020) BPRP 60 Ngorora Africa Kenya 12,0 12,0 Middle Miocene Van Couvering and Delson (2020) BPRP 65 Ngorora Africa Kenya 12,8 12,8 Middle Miocene Van Couvering and Delson (2020) BPRP K089A Baringo Africa Kenya 15,8 15,8 Middle Miocene Van Couvering and Delson (2020) BPRP K091 Baringo Africa Kenya 15,8 15,8 Middle Miocene Van Couvering and Delson (2020) Bukwa II Bukwa Africa Uganda 19,5 19,1 Early Miocene MacLatchy et al. (2006); Cote et al. (2018) [contra Pickford (2017)] Buluk Member Bakate Fm. Africa Kenya 17,2 17,2 Early Miocene McDougall and Watkins (1985) Calcified Tuff Member Lower Kapurtay Africa Kenya 20,0 18,5 Early Miocene Bishop et al. (1969) Can Feu 1 Can Feu Europe Spain 10,0 9,7 Late Miocene Casanovas-Vilar et al. (2016) Can Llobateres 1 Can Llobateres Europe Spain 9,8 9,8 Late Miocene Casanovas-Vilar et al. (2016) Can Llobateres 2 Can Llobateres Europe Spain 9,6 9,6 Late Miocene Casanovas-Vilar et al. (2016) Can Mata 1 Can Mata Europe Spain 11,6 11,2 Late Miocene Casanovas-Vilar et al. (2011) Can Mata s.l. Can Mata Europe Spain 11,3 11,0 Late Miocene Alba et al. (2022) Can Pallars i Llobateres Can Pallars i Llobateres Europe Spain 10,0 9,7 Late Miocene Alba et al. (2018) Can Poncic 1 Can Poncic Europe Spain 10,3 10,0 Late Miocene Casanovas-Vilar et al. (2011) Can Vila Can Vila Europe Spain 12,0 12,0 Middle Miocene Casanovas-Vilar et al. (2011) Çandir Çandir Asia Turkey 14,1 13,4 Middle Miocene Casanovas-Vilar et al. (2011) Castell de Barberà Castell de Barberà Europe Spain 11,2 11,2 Late Miocene Casanovas-Vilar et al. (2011) Chakrana Haritalyangar Asia India 9,2 8,1 Late Miocene Patnaik (2013) Chamtwara member Koru-Muhoroni area Africa Kenya 20,3 20,3 Early Miocene Van Couvering and Delson (2020) Cheboit Lukeino Africa Kenya 6,2 5,7 Late Miocene Pickford and Senut (2001) Chorora Fm. Beticha Africa Ethiopia 8,0 8,0 Late Miocene Katoh et al. (2016) Çorakyerle Çorakyerle Asia Turkey 8,1 7,6 Late Miocene Kaya et al. (2015) Dalsar Ramnagar Asia India 13,8 12,5 Middle Miocene Sehgal and Patnaik (2012); Gilbert et al. (2020) Dera Gopipur Himachal Pradesh Asia India ? ? - Pickford and Tiwari (2010) Devínska Nová Ves –Sandberg (= Neudorf Sandberg) Devínska Nová Ves –Sandberg (= Neudorf Sandberg) Europe Slovakia 13,6 13,1 Middle Miocene Březina et al. (2021); Sabol et al. (2021) Dhara Reserve Uttar Pradesh Asia India 11,0 10,0 Late Miocene Johnson et al. (1983); Cameron et al. (1999) Domeli Potwar Plateau Asia Pakistan 11,5 9,0 Late Miocene Barry et al. (2013) Ebingen Swabian Alb Europe Germany 11,0 9,0 Late Miocene Pickford (2012) EDAR13 EDAR Europe Spain 10,3 10,0 Late Miocene Casanovas-Vilar et al. (2011) Engelwies Engelswies Europe Germany 16,5 16,0 Early Miocene Casanovas-Vilar et al. (2011) Fangbeiliangzi Xiaohe Fm. Asia China 8,2 8,1 Late Miocene Woodburne et al. (2013) Fiume Santo Fiume Santo Europe Italy 8,1 6,7 Late Miocene Rook et al. (2011) FJ-18 Fejej Africa Ethiopia 16,2 16,2 Early Miocene Richmond et al. (1998) Fort Ternan Fort Ternan Africa Kenya 13,8 13,8 Middle Miocene Van Couvering and Delson (2020) Hammerschmiede 5 Pforzen Europe Germany 11,6 11,6 Late Miocene Böhme et al. (2019) Hari Devi 1 (HD1) Haritalyangar Asia India 8,9 8,9 Late Miocene Pillans et al. (2005) Haritalyangar Haritalyangar Asia India 9,2 8,1 Late Miocene Patnaik (2013) Hiwegi Fm. Rusinga Island Africa Kenya 18,3 18,3 Early Miocene Van Couvering and Delson (2020) Hudieliangzi Xiaohe Fm. Asia China 8,2 8,1 Late Miocene Woodburne et al. (2013) Jabbi Potwar Plateau Asia Pakistan 10,4 8,5 Late Miocene Lydekker (1879); Kelley (2005a) Kalodirr Member Losodok (=Lothidok, Losidok) Africa Kenya 17,5 17,5 Early Miocene Van Couvering and Delson (2020) Kaloma Kaloma Africa Kenya 15,3 15,3 Middle Miocene Van Couvering and Delson (2020) Kapsibor Kapsibor Africa Kenya 13,7 13,7 Middle Miocene Harrison (1986) Kapsomin Lukeino Africa Kenya 6,2 5,7 Late Miocene Pickford and Senut (2001) Kapurtay Kapurtay Africa Kenya 20,0 18,5 Early Miocene Bishop et al. (1969) Kiahera Fm. Rusinga Island Africa Kenya 18,5 18,5 Early Miocene Van Couvering and Delson (2020) Kleinshadersdorf (= Klein Hadersdorf) Vienna Basin Europe Austria 13,8 12,7 Middle Miocene Steininger (1986), Kováč et al. (2018) y Březina et al. (2021) Koru Fm. Koru-Muhoroni area Africa Kenya 20,3 20,3 Early Miocene Van Couvering and Delson (2020) Kulu Fm. Rusinga Island Africa Kenya 16,0 16,0 Early Miocene Van Couvering and Delson (2020) Kundal Nala [Chinji] Potwar Plateau Asia Pakistan 11,5 11,4 Late Miocene Barry et al. (2013) La Grive Saint-Alban L3/La Grive Saint-Alban L5? Isère Europe France 11,9 11,2 Middle-Late Miocene Casanovas-Vilar et al. (2011) La Grive Saint-Alban M Isère Europe France 13,0 11,9 Middle Miocene Casanovas-Vilar et al. (2011) La Tarumba 1 La Tarumba Europe Spain 9,6 9,6 Late Miocene Casanovas-Vilar et al. (2016) Legetet Fm. Koru-Muhoroni area Africa Kenya 20,0 19,0 Early Miocene McCollum et al. (2013) Leilao Xiaohe Fm. Asia China 8,2 7,1 Late Miocene Woodburne et al. (2013) Locality 14 (Maize Crib) Koru-Muhoroni area (Koru) Africa Kenya 20,0 19,0 Early Miocene Pickford (1986) Locality 34 (Maize Crib) Koru-Muhoroni area (Koru) Africa Kenya 20,0 19,0 Early Miocene Pickford (1986) Locherangan Locherangan Africa Kenya 17,6 17,6 Early Miocene Van Couvering and Delson (2020) Lower Sand Unit Nakhon Ratchasima Province (Khorat) Asia Thailand 9,0 6,0 Late Miocene Chaimanee et al. (2006) LpM4 Loperot Africa Kenya 17,0 17,0 Early Miocene Liutkus-Pierce et al. (2019) Magare Uyoma Peninsula Africa Kenya 18,3 18,3 Early Miocene Drake et al. (1988); Van Couvering and Delson (2020) Majiwa Majiwa Africa Kenya 15,3 15,3 Middle Miocene Van Couvering and Delson (2020) Maragheh (Middle Pumice) Maragheh Asia Iran 7,5 7,5 Late Miocene Suwa et al. (2016) Melchingen Swabian Alb Europe Germany 11,2 7,5 Late Miocene Casanovas-Vilar et al. (2011) Meswa Bridge (Locality 36) Koru-Muhoroni area Africa Kenya 22,5 22,5 Early Miocene Van Couvering and Delson (2020) Mfangano Island Mfangano Island Africa Kenya 18,3 18,3 Early Miocene Van Couvering and Delson (2020) Moroto I Karamoja Africa Uganda 21,0 20,0 Early Miocene Cote et al. (2018) Moroto II Karamoja Africa Uganda 21,0 20,0 Early Miocene Cote et al. (2018) Moruorot Losodok (=Lothidok, Losidok) Africa Kenya 17,5 17,5 Early Miocene Boschetto et al. (1992) Muruyur Cheparawa Africa Kenya 15,8 15,8 Middle Miocene Van Couvering and Delson (2020) Muruyur - Kipsaraman Baringo Africa Kenya 15,8 15,8 Middle Miocene Van Couvering and Delson (2020) N 885 N 885 Africa Niger 11,0 8,0 Late Miocene Mocke et al. (2022) Nakali Nakali Africa Kenya 9,9 9,8 Late Miocene Kunimatsu et al. (2007) Namurungule Fm. Samburu Hills Africa Kenya 8,5 8,5 Late Miocene Van Couvering and Delson (2020) Napak CC Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak I Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak IV Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak IX Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak V Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak XII Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak XIII Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak XV Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak XVIII Akisim Africa Uganda 20,5 20,5 Early Miocene Pickford et al. (2020) Napak XXVI Akisim Africa Uganda 20,5 19,0 Early Miocene Pickford et al. (2020) Napak XXX Akisim Africa Uganda 20,5 20,5 Early Miocene Pickford et al. (2020) Napak XXXI Akisim Africa Uganda 20,5 20,5 Early Miocene Pickford et al. (2020) Napak XXXII Akisim Africa Uganda 20,1 20,1 Early Miocene Van Couvering and Delson (2020) Neuhausen Swabian Alb Europe Germany 11,0 9,0 Late Miocene Pickford (2012) Nikiti 1 Chalkidiki Peninsula Europe Greece 9,7 8,8 Late Miocene Casanovas-Vilar et al. (2011) North Napudet Napudet Africa Kenya 13,3 13,3 Middle Miocene Nengo et al. (2017) Nyakach Nyakach Africa Kenya 15,0 15,0 Middle Miocene Van Couvering and Delson (2020) Ombo Ombo Africa Kenya 16,0 16,0 Early Miocene Van Couvering and Delson (2020) Paşalar Paşalar Asia Turkey 14,5 14,0 Middle Miocene Casanovas-Vilar et al. (2011) Pikermi Fm. (Red Conglomerate Member) Pyrgos Europe Greece 7,2 7,2 Late Miocene Böhme et al. (2017) Polinyà 2 Polinyà Europe Spain 11,6 9,7 Late Miocene Casanovas-Vilar et al. (2011) Potwar Plateau [Chinji] Potwar Plateau Asia Pakistan 12,7 11,2 Middle-Late Miocene Barry et al. (2002); Kelley (2005b) [used for specimens collected in early 20th century] Potwar Plateau [Dhok Pathan] Potwar Plateau Asia Pakistan 9,8 8,5 Late Miocene Barry et al. (2002); Kelley (2005b) [used for specimens collected in early 20th century] Ramnagar Ramnagar Asia India 13,8 12,5 Middle Miocene Pillans et al. (2005); Gilbert et al. (2020) Rashole 3 Ramnagar Asia India 13,8 12,5 Middle Miocene Gilbert et al. (2020) Ravin de la Pluie Axios Valley Europe Greece 9,4 9,3 Late Miocene Casanovas-Vilar et al. (2011) Redbed Member Unit 5 Koru-Muhoroni area (Songhor) Africa Kenya 20,0 18,5 Early Miocene Bishop et al. (1969) Redbed Member Unit 6 Koru-Muhoroni area (Songhor) Africa Kenya 20,0 18,5 Early Miocene Bishop et al. (1969) Rudabánya 2 Pannonian Basin Europe Hungary 10,0 9,7 Late Miocene Casanovas-Vilar et al. (2011) Ryskop Ryskop Africa South Africa 16,0 16,0 Early Miocene Van Couvering and Delson (2020) Salmendingen Swabian Alb Europe Germany 11,6 7,5 Late Miocene Casanovas-Vilar et al. (2011) Shihuiba 75033 D2-6 Lufeng Asia China 6,9 6,2 Late Miocene Yue and Zhang (2006) Shuitangba Shuitangba Asia China 6,2 6,2 Late Miocene Jablonski et al. (2014) Sinap Locality 12 Sinap Formation Asia Turkey 9,8 9,8 Late Miocene Kappelman et al. (2003) Siwaliks B0009 Hasnot (Nagri Fm. - Siwalik Group) Asia Pakistan 11,2 10,8 Late Miocene Brown et al. (1924); Barry et al. (2002); Barry et al. (pers. comm.) Siwaliks B0145 ? (Chinji Fm. - Siwalik Group) Asia Pakistan ? ? - Barry et al. (pers. comm.) Siwaliks KL0001 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry et al. (pers. comm.) Siwaliks L0021 Hutch Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 11,4 11,4 Late Miocene Barry (1986); Barry et al. (pers. comm.) Siwaliks L0024 Hutch Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 12,4 12,2 Middle Miocene Barry et al. (pers. comm.) Siwaliks L0035 Haritalyangar Asia India ? ? - Barry (1986); Barry et al. (pers. comm.) Siwaliks L0039 Haritalyangar Asia India ? ? - Barry et al. (pers. comm.) Siwaliks L0040 Haritalyangar Asia India ? ? - Barry et al. (pers. comm.) Siwaliks L0081 Andar Kas (Chinji Fm. - Siwalik Group) Asia Pakistan 11,2 10,8 Late Miocene Barry (1986); Barry et al. (pers. comm.) Siwaliks L0094 Dhala Nala (Lower) (Nagri Fm. - Siwalik Group) Asia Pakistan 10,3 10,1 Late Miocene Barry et al. (pers. comm.) Siwaliks S0008 ? (Manchar Fm. - Siwalik Group) Asia Pakistan ? ? - Khan et al. (1984); Raza et al. (1984); Barry et al. (pers. comm.) Siwaliks Y0076 Gambhir Section (Chinji Fm. - Siwalik Group) Asia Pakistan 11,5 11,4 Late Miocene Kappelman et al. (1991); Barry et al. (2002, pers. comm.); Flynn et al. (2020) Siwaliks Y0083 Hutch Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 11,6 11,6 Late Miocene Flynn et al. (2020); Barry et al. (pers. comm.) Siwaliks Y0137 Gandakas Road (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0158 (=Y0388) Malhuwala Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 8,8 8,7 Late Miocene Barry et al. (pers. comm.) Siwaliks Y0182 Gandakas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0191 Gandakas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0207 Malhuwala Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0211 Dinga Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0221 Dinga Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0224 Dinga Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0226 Dinga Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0227 227 Section Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0230 227 Section Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0251 Ratha Kas (Nagri Fm. - Siwalik Group) Asia Pakistan 10,0 10,0 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0259 Kaulial Kas (Nagri Fm. - Siwalik Group) Asia Pakistan 10,5 10,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0260 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0261 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,7 9,7 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0309 Dhok Mila, Khaur Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0310 Dhok Mila, Khaur Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0311 Gambhir Section (Nagri Fm. - Siwalik Group) Asia Pakistan 10,1 10,0 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0314 Dhok Mila, Khaur Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0317 Choutriwali Kas (Nagri Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0327 Kot Maliaran (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0328 Hasal Kas, Bora Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0350 269 Section (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0409 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0410 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0414 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0416 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,3 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0442 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 8,6 8,5 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0463 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0494 Gambhir Section (Chinji Fm. - Siwalik Group) Asia Pakistan 12,4 12,4 Middle Miocene Barry (1986); Barry et al. (2002, pers. comm.) Siwaliks Y0495 Gambhir Section (Chinji Fm. - Siwalik Group) Asia Pakistan 11,6 11,6 Late Miocene Barry (1986); Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0496 Gambhir Section (Chinji Fm. - Siwalik Group) Asia Pakistan 12,4 12,4 Middle Miocene Barry (1986); Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0498 Hutch Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 11,7 11,7 Middle Miocene Barry (1986); Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0499 Kanatti Upper (Chinji Fm. - Siwalik Group) Asia Pakistan 12,1 12,1 Middle Miocene Barry (1986); Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0500 Kanatti Upper (Chinji Fm. - Siwalik Group) Asia Pakistan 12,1 12,1 Middle Miocene Barry (1986); Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0502 Hutch Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 12,0 12,0 Middle Miocene Barry et al. (pers. comm.) Siwaliks Y0604 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 8,6 8,5 Late Miocene Barry (1986); Barry et al. (pers. comm.) Siwaliks Y0647 Gambhir Section (Chinji Fm. - Siwalik Group) Asia Pakistan 12,4 12,4 Middle Miocene Morgan et al (2015); Barry et al. (pers. comm.) Siwaliks Y0663 Parrawali (Chinji Fm. - Siwalik Group) Asia Pakistan 11,6 11,6 Late Miocene Barry (1986); Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0750 Samriala Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 12,8 12,7 Middle Miocene Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0767 Gambhir Section (Chinji Fm. - Siwalik Group) Asia Pakistan 12,4 12,3 Middle Miocene Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0775 Rata Dala Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 12,4 12,2 Middle Miocene Kappelman et al. (1991); Barry et al. (pers. comm.) Siwaliks Y0829 Samriala Nala (Chinji Fm. - Siwalik Group) Asia Pakistan 11,9 11,1 Middle-Late Miocene Barry et al. (pers. comm.) Siwaliks Y0885 ? (Siwalik Group) Asia Pakistan ? ? - Barry et al. (pers. comm.) Siwaliks Y0891 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 8,8 8,7 Late Miocene Barry et al. (pers. comm.) Siwaliks Y0990 ? (Chinji Fm. - Siwalik Group) Asia Pakistan ? ? - Barry et al. (pers. comm.) Siwaliks Y0997 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,3 9,2 Late Miocene Barry et al. (pers. comm.) Siwaliks Y1002 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,4 Late Miocene Barry et al. (pers. comm.) Siwaliks Y1004 Kaulial Kas (Dhok Pathan Fm. - Siwalik Group) Asia Pakistan 9,4 9,3 Late Miocene Barry et al. (pers. comm.) SMG1 Irrawaddy Fm. Asia Myanmar 10,0 8,0 Late Miocene Takai et al. (2021) Somsak Nakhon Ratchasima Province (Khorat) Asia Thailand 9,0 6,0 Late Miocene Chaimanee et al. (2006) Songhor Koru-Muhoroni area Africa Kenya 20,3 20,3 Early Miocene Van Couvering and Delson (2020) St. Gaudens Haute-Garonne Europe France 12,5 11,2 Middle-Late Miocene Casanovas-Vilar et al. (2011) St. Stefan im Lavanttal Gratkonr Basin Europe Austria 12,2 12,0 Middle Miocene Casanovas-Vilar et al. (2011) Sunetar 2 Lower Siwalik Asia India 13,8 12,5 Middle Miocene Gilbert et al. (2020) Tapar Kutch Asia India 10,8 10,0 Late Miocene Flynn et al. (2013); Patnaik et al. (2022) Teuleria del Firal Teuleria del Firal Europe Spain 10,3 10,0 Late Miocene Casanovas-Vilar et al. (2011) Thannhausen Thannhausen Europe Germany 14,0 14,0 Middle Miocene Pickford (2013) Tinau Khola Dang Valley Asia Nepal 9,5 9,0 Late Miocene Munthe et al. (1983); West et al. (1991) Tinau Khola Dang Valley Asia Nepal 9,5 9,0 Late Miocene Munthe et al. (1983) Trochtelfingen Swabian Alb Europe Germany 11,0 9,0 Late Miocene Pickford (2012) Udabno 1 Udabno Asia Georgia 8,1 7,7 Late Miocene Agustí et al. (2020) V1 Baccinello Europe Italy 8,3 8,1 Late Miocene Rook et al. (2011) V2 Baccinello Europe Italy 8,1 6,7 Late Miocene Rook et al. (2011) VPL-1 Hari Mandar Dhar Asia India 9,0 9,0 Late Miocene Pillans et al. (2005) Wissberg Gau-Weinheim Europe Germany 13,7 7,5 Middle-Late Miocene Casanovas-Vilar et al. (2011) Wudu Gansu Asia China 8,3 8,3 Late Miocene Woodburne et al. (2013) Xiaolungtan Kaiyuan Asia China 12,5 11,6 Middle-Late Miocene Li et al. (2015) Xirochori 1 Axios Valley Europe Greece 9,7 9,7 Late Miocene Casanovas-Vilar et al. (2011) YS1 Irrawaddy Fm. Asia Myanmar 10,4 8,8 Late Miocene Jaeger et al. (2011) Zhupeng Xiaohe Fm. Asia China 7,4 7,2 Late Miocene Zhu et al. (2005) a When indicated as 'Barry et al. (pers. comm.)", age of Siwalik localities checked or updated based on unpublished data kindly provided by John Barry, Larry Flynn, Jay Kelley, Michèle Morgan, and David Pilbeam on September 2022. SOM Table S3 Chronostratigraphic ranges of Miocene apes divided by 1 Myr-duration bins and the corresponding diversity metrics. Miocene ape species Range / Bin 23-22 22-21 21-20 20-19 19-18 18-17 17-16 16-15 15-14 14-13 13-12 12-11 11-10 10-9 9-8 8-7 7-6 6-5 Proconsul africanus 20.3–18.5 x x x Proconsul major 20.5–19 x x Proconsul gitongai 15.8 x Proconsul meswae 22.5 x Proconsul legetetensis 20.5–19.0 x x Kalepithecus songhorensis 20.5–19.0 x x Kalepithecus kogolensis 21.0-20.0 x Ekembo heseloni 18.5–16.0 x x x Ekembo nyanzae 18.5–16.0 x x x Xenopithecus koruensis 20.0–19.0 x Mabokopithecus clarki 15.3 x Rangwapithecus gordoni 20.0–17.0 x x x Nyanzapithecus pickfordi 15.8–15.3 x Nyanzapithecus vancouveringorum 18.3 x Nyanzapithecus alesi 13.3 x Nyanzapithecus harrisoni 15.0 x Turkanapithecus kalakolensis 17.5–17.0 x Turkanapithecus rusingensis 20.5–18.3 x x x Samburupithecus kiptalami 8.5 x Afropithecus turkanensis 17.6–17.2 x Heliopithecus leakeyi 16.0 x Morotopithecus bishopi 21-20 x Equatorius africanus 16.0–15.0 x Nacholapithecus kerioi 15.0 x Otavipithecus namibiensis 12.0 x Yuanmoupithecus xiaoyuan 8.2–7.1 x x Griphopithecus suessi 13.8–12.7 x x Griphopithecus alpani 14.5–13.4 x x Kenyapithecus wickeri 13.8 x Kenyapithecus kizili 14.5–14.0 x Dryopithecus fontani 12.5–11.0 x x Pierolapithecus catalaunicus 12.0 x Anoiapithecus brevirostris 12.4–12.0 x Hispanopithecus laietanus 10.0–9.6 x Hispanopithecus crusafonti 10.3–10.0 x Rudapithecus hungaricus 10.0–9.8 x Danuvius guggenmosi 11.6 x ?Udabnopithecus garedziensis 8.1–7.7 x x Khoratpithecus chiangmuaneneis 12.4–12.2 x Khoratpithecus piriyai 9.0–6.0 x x x Khorapithecus ayeyarwadyenis 10.4–8.0 x x x Khoratpithecus magnus 9.0–6.0 x x x Ankarapithecus meteai 9.8 x Lufengpithecus lufengensis 6.9–6.2 x Sivapithecus sivalensis 10.4–7.5 x x x x Sivapithecus indicus 13.0–10.8 x x x Sivapithecus parvada 10.1–10.0 x Indopithecus giganteus 8.9–8.6 x Chororapithecus abyssinicus 8.0 x Nakalipithecus nakayamai 9.9–9.8 x Graecopithecus freybergi 7.2 x Ouranopithecus macedoniensis 9.7–8.8 x x Ouranopithecus turkae 8.1–7.6 x x Sinopithecus keiyuanensis 12.5–11.6 x x Sinopithecus hudienensis 8.2–7.1 x x Oreopithecus bambolii 8.3–6.7 x x x ‘Dryopithecus ’ wuduensis 8.3 x Variable 23-22 22-21 21-20 20-19 19-18 18-17 17-16 16-15 15-14 14-13 13-12 12-11 11-10 10-9 9-8 8-7 7-6 6-5 Nbt 0 0 0 1 1 2 0 0 0 0 0 1 0 2 0 3 0 0 NbL 0 0 0 4 2 1 2 0 0 1 1 2 1 0 2 5 3 0 NFt 0 0 5 2 2 0 0 0 1 1 4 0 2 1 7 0 0 0 NFL 1 0 2 0 1 2 0 5 3 2 1 3 1 5 4 2 1 0 Ntot = Nbt + NbL + NFt + NFL 1 0 7 7 6 5 2 5 4 4 6 6 4 8 13 10 4 0 Nst = (NbL + NFt + 2Nbt) / 2 0 0 2,5 4 3 2,5 1 0 0,5 1 2,5 2 1,5 2,5 4,5 5,5 1,5 0 Abbreviations: Nbt = range-through taxa; NbL = bottom-only boundary crossers; NFt = top-only boundary crossers; NFL = single-interval taxa; Ntot = range-through or total diversity; Nst = standing diversity.