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Supporting Information Moya` -Sola` et al. 10.1073/pnas.0811730106 SI Text netic relationships between all these taxa by classifying them all Systematic Framework. In Table S1 we provide a systematic into a single family Afropithecidae with 2 subfamilies (Keny- classification of living and fossil Hominoidea to the tribe level, apithecinae and Afropithecinae). by further including extant taxa and extinct genera discussed in The systematic scheme used here requires several nomencla- this paper. Hominoidea are defined as the group constituted by tural decisions, which deserve further explanation. The nomina Hylobatidae and Hominidae, plus all extinct taxa more closely Kenyapithecini and Kenyapithecinae are adopted instead of related to them than to Cercopithecoidea. Hominidae, in turn, Griphopithecinae and Griphopithecini (see also ref. 25) merely are defined as the group containing Ponginae and Homininae, because the former have priority. It is unclear why neither Begun plus all extinct forms more closely related to them than to (1) nor Kelley (2) specify the authorship of Griphopithecinae (or Hylobatidae. While this broad concept of Hominidae is currently Griphopithecidae), but, to our knowledge, the authorship of the used by many paleoprimatologists (e.g., refs. 1–2), the systematic latter nomina must be attributed to Begun (ref. 4, p. 232: Table position of primitive (or archaic) putative hominoids is far from 10.1), which therefore do not have priority over Kenyapithecinae clear (see below). Begun (3–5) employs the terms ‘‘Eohomi- Andrews, 1992. Griphopithecinae thus remains potentially valid noidea’’ and ‘‘Euhominoidea’’ to informally refer to hominoids only if Kenyapithecus (and Afropithecus, see below) are excluded of primitive and modern aspect, respectively. These terms, from it. This notwithstanding, there has been some confusion however, are roughly equivalent to stem-lineage and crown- regarding the authorship of the nomen Kenyapithecinae. Both group hominoids, respectively, and are not further used here. Begun (1) and Ward and Duren (20) attribute its authorship to The systematic status of many Late Oligocene and Early- Leakey (26). However, the earliest usage of a suprageneric Middle Miocene fossil catarrhines, lacking both cercopithecoid nomen with Kenyapithecus as the type genus is attributable to synapomorphies and crown hominoid synapomorphies, has been Andrews (6), who erected it as a previously undescribed tribe, subject to different interpretations (6–8). Harrison (8) argues Kenyapithecini, in the same paper that he erected Afropithecini that most fossil ‘‘apes’’ from the Late Oligocene and Early (27). To our knowledge, Kenyapithecus and Afropithecus have not Miocene of Africa have not crossed the hominoid ‘‘cladistic been previously included into a single family with the exclusion threshold’’ and classifies them into 2 distinct superfamilies of Hominidae (as in ref. 20), but we have chosen the nomen (Dendropithecoidea and Proconsuloidea), which he considers to Afropithecidae (instead of Kenyapithecidae) because the former be successive sister taxa of stem catarrhines, more derived than has been already used at the family level—albeit with a different propliopithecoids and pliopithecoids, but presumably preceding meaning—by some previous workers (2, 5). If, as suggested in the cercopithecoid–hominoid split (8–12). On the other ex- this paper, there is a close phylogenetic relationship between treme, a few authors have considered that proconsuloids might Kenyapithecinae (in particular, the Kenyapithecini) and Hom- be stem hominids (13–15). Although the latter view is apparently inidae, Afropithecidae as conceived here would be paraphyletic. abandoned (16), most authors consider that Early Miocene Transferring the Kenyapithecinae into the Hominidae (25), forms, especially Proconsul, are more closely related to extant however, would not solve this problem, since the remaining hominoids than to cercopithecoids (3, 6–7, 16–20). The system- Afropithecidae (including only the Afropithecinae) would re- atic scheme used here (Table S1) follows Harrison (8) in main paraphyletic in excluding the Kenyapithecinae. Paraphyly recognizing that proconsulids and dendropithecids are distinct can be transferred from one group or rank to another, but cannot clades, but considers that at least the former are stem hominoids. be completely eliminated unless Linnean ranks are aban- These putative stem hominoids lack crown hominoid synapo- doned—a view that is not advocated here. morphies, in particular, features functionally related to ortho- The Late Miocene genera Hispanopithecus and Ouranopithe- grady that are presumably homologous between hylobatids and cus are not classified here at the tribe level. Both genera have hominids. Yet stem hominoids already share with both hylo- been previously suggested to be either pongines (28) or homi- batids and hominids some facial (16) and several postcranial nines (4), but this issue is not definitively resolved and lies (17–18, 21) synapomorphies. Prominently, the lack of external outside the scope of this paper. Nevertheless, it is worth men- tail in Proconsul (17, 19, 21), albeit disputed by some authors tioning that Hispanopithecus was recently resurrected (29) for (22), is now firmly established (23) and has been further ascer- Late Miocene species previously included in Dryopithecus,so tained in the putative stem hominoid Nacholapithecus (24). that the latter genus is restricted to its type species, Dryopithecus Among putative stem hominoids, the position of Afropithecus fontani. As a result, the tribe Dryopithecini is here used with a and other related taxa is the most problematic. Harrison (8) different meaning from previous usages, to refer to Middle distinguishes a single family Proconsulidae with 3 distinct sub- Miocene stem hominids that apparently do not belong to any of families (Proconsulinae, Nyanzapithecinae, and Afropitheci- the 2 crown-hominid subfamilies; this tribe includes Pierolapithe- nae), while other authors (20) take an opposite approach by cus, Dryopithecus s.s., and Anoiapithecus gen. nov., but most classifying their Afropithecinae (including Nacholapithecus and likely excludes Hispanopithecus. Given the uncertain phyloge- Equatorius) within the Hominidae, and others (2) consider the netic relationships between the several dryopithecin genera, it is former to be a distinct family. Especially problematic is the currently unclear whether this tribe is paraphyletic or represents placement of Kenyapithecus and Griphopithecus: in some sys- a clade of stem European hominids. tematic schemes (20, 25), the latter taxa are classified into the subfamily Kenyapithecinae within the Hominidae; others (2) Results distinguish a subfamily Griphopithecinae (for Griphopithecus) Morphometric Analyses. ANOVA comparisons show that there within the Afropithecidae; and Begun reunites both Keny- are significant differences (P Ͻ 0.001; F ϭ 178.6) among several apithecinae and Griphopithecinae into a distinct family Gripho- extant catarrhines regarding the CFA (Table S2 and S3), with pithecidae (2) or combines these taxa into an informal grouping gorillas, colobines, and hylobatids displaying the highest values, (‘‘griphopiths’’) of stem hominoids (5). The systematic scheme which nevertheless generally do not surpass 60°. Living humans, followed in this paper (Table S1) recognizes the close phyloge- on the contrary, differ from all of the remaining taxa (P Ͻ 0.001) Moya` -Sola` et al. www.pnas.org/cgi/content/short/0811730106 1of10 by the much higher CFA of the former, while extinct hominins ape to have ever displayed a colobine-like facial profile, to which display intermediate values. Most extinct catarrhines clearly fall it may have autapomorphically converged from an ancestral within the ranges of extant nonhuman cercopithecoids and condition more similar to that displayed by stem hominoids and hominoids, with the exception of Anoiapithecus. The latter taxa living cercopithecines. display values of CFA beyond the maximum value recorded in Squared distances based on the CVA discriminant scores, and extant nonhuman catarrhines, and Ϸ20° higher than other fossil computed for pairs of fossil individuals included (Table S5), apes, most closely resembling the values displayed by extinct indicate that Anoiapithecus is particularly far from the 2 other members of the genus Homo. Miocene hominoids from Spain included in the analysis (Piero- The canonical variate analysis (CVA) indicates that Anoia- lapithecus and Hispanopithecus); D. fontani could not be included pithecus displays a unique morphology, previously unknown in the analysis because of incomplete preservation, although a among living and fossil hominoids, which confirms the need to previous analysis of facial morphology based on new remains erect a previously undescribed genus. This analysis (Table S4 and from Abocador de Can Mata indicates a gorilla-like morphology Table S5) correctly classifies 93% of the original cases, only with (29), which is thus quite different from the Anoiapithecus minor confusion between some chimpanzees and gorillas and condition. The results of the randomization approach further between very few colobines, cercopithecines, and hylobatids. confirm that differences between Anoiapithecus and other fossil Stem catarrhines, stem hominoids, and even the stem great ape individuals cannot be accommodated within the range of vari- Pierolapithecus most closely resemble cercopithecines, whereas ation of a
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