Anadoluvius Sevim-Erol & Begun & Sözer & Mayda & van den Hoek Ostende & Martin & Alçiçek 2023, gen. nov.

<i>Anadoluvius</i> gen. nov. <i>Synonomy</i>. <i>Ouranopithecus</i> Bonis and Melentis: Güleç et al. 28 <i>Type species</i>. <i>Anadoluvius turkae</i> comb. nov. Sevim Erol et al. 2023. <i>Etymology</i>. Anadolu is the modern Turkish word for Anatolia and Anatolian. <i>Holotype</i>. CO-205, a fragmented but largely complete male palate with LI1-M3 and RC-M2 (Supplementary Figs. 1, 2). <i>Paratypes</i>. CO-300 (RM 2); CO-305 (male mandibular fragment with RC-M 1); CO-710 (female mandibular fragment with RP 3 - M 2); CO-2100 (RI 1); CO-2800 (female partial cranium with RC- M 2, portions of the right maxilla, maxillary frontal processes, frontal maxillary processes and most of the frontal bone) (Fig. 1; Supplementary Figs. 3–5) Detailed specimen descriptions and a revised diagnosis for this new taxon appear in the Supplementary Notes 2, 3. The hypodigm is curated in the Department of Anthropology, Ankara University. All samples used in this analysis are listed in Supplementary Tables 1, 2. Measurements are provided in Supplementary Tables 3, 4. Supplementary Note 4 presents the results of a comprehensive quantitative analysis of the <i>Anadoluvius</i> hypodigm. Supplementary Note 5 provides details of the phylogenetic analysis. Supplementary Note 6 provides historical, geological and biochronologic background. <i>Cranium</i>. The CO-2100/2800 partial cranium (Fig. 1, Supplementary Figs. 2, 3) was recovered with some crushing and displacement of several broken pieces (see SM for restoration details and detailed description.) The frontal bone is nearly intact, missing only portions within the temporal fossa and the squama approaching bregma. This distinguishes it from the most complete facial specimen of <i>Ouranopithecus</i> (XIR-1), which is broken just beyond the superior orbital margins and preserves almost nothing of the frontal squama (Supplementary Fig. 2). Though damaged, the position and orientation of the premaxilla is better preserved in CO-2100/2800 than in CO-205 and <i>Ouranopithecus</i> (XIR-1 and RPl 128), confirming previous interpretations of a stepped and overlapping morphology in these specimens 1 – 3, 11, 13 – 15 (Fig. 2). The premaxilla of <i>Anadoluvius</i> is short and vertical compared with <i>Pan</i>, <i>Pongo</i>, and australopithecines, and is most like <i>Gorilla</i> and dryopithecins, being relatively short in the alveolar portion but expanded nasally to overlap with the palatine process of the maxilla (Fig. 2) The incisor alveoli are positioned along the mesial transverse plane of the canine crowns (Figs. 1, 2; Supplementary Fig. 5) In two specimens of <i>Ouranopithecus</i> (RPl 128 and XIR 1) the upper incisors are well anterior to the canines (Supplementary Fig. 5) In NKT 89 the premaxilla is severely damaged, but the posterior edge of the lateral incisor appears to be aligned with the anterior transverse plane of the canines, a position most like <i>Anadoluvius</i>. The frontal bone of <i>Anadoluvius</i> differs strongly from that of <i>Ouranopithecus</i> in the smooth biconvex squama of the former, contrasting with a broad concavity above glabella in the latter. The superior orbital margins of <i>Ouranopithecus</i> are broad, rounded and slightly projecting while they are sharp and flat in <i>Anadoluvius</i>. <i>Mandible</i>. A principal components analysis based on mandibular measurements available for <i>Graecopithecus, Ouranopithecus</i> and <i>Anadoluvius</i> is presented in Supplementary Fig. 6 and Supplementary Data 1). The Çorakyerler, Nikiti 1 and <i>Graecopithecus</i> mandibles are separated from each other, especially along PC 2, and from <i>Ouranopithecus</i>, illustrating the diversity present in these samples. Supplementary Data 1 includes the data matrix, summary statistics, scores, and loadings. <i>Anadoluvius</i>, like <i>Graecopithecus</i>, and NKT 21, has a relatively narrow mandible compared with the combined sex sample of <i>Ouranopithecus</i> (Supplementary Fig. 7a). Supplementary Fig. 7b compares relative mandibular corpus breadth at each tooth position (P 3 -M 2) in <i>Anadoluvius, Graecopithecus</i>, and <i>Ouranopithecus</i>. <i>Anadoluvius</i> is similar in mandibular robusticity at the premolar level but at the level of the molars it matches or strongly exceeds the maximum value in the other taxa (Supplementary Fig. 7b). There is diversity in mandibular robusticity (breadth relative to height) and in dental size ratios as well among the samples of eastern Mediterranean apes (Supplementary Fig. 8a–e). <i>Anadoluvius</i> is distinct from <i>Graecopithecus</i> in all mandibular and dental ratios. <i>Anadoluvius</i> falls beyond the range of variation of <i>Ouranopithecus</i> in relative corpus breadth at M 1 -M 2 (Supplementary Fig. 8a) and M 2 size (Supplementary Fig. 8d). Interestingly, NKT 21 falls outside the <i>Ouranopithecus</i> range in relative P 4 length and M 2 size (Supplementary Fig. 8d–f). It has a relatively short symphyseal-molar distance, at the 25% quartile for the Ravin sample, and a relatively robust mandible at M 1 -M 2, at the 75% quartile for <i>Ouranopithecus</i> (Supplementary Fig. 8a, c). In <i>Graecopithecus</i> the symphysis is positioned closest to the molars, just barely in the range of the Ravin sample. In summary, in most quantitative comparisons <i>Anadoluvius</i> is distinguished from <i>Ouranopithecus</i> and <i>Graecopithecus</i>. <i>Tooth roots and enamel thickness</i>. Figure 3 shows the root and root canal morphology of CO-300, the male mandible of <i>Anadoluvius</i> (see Methods for segmentation details). Unlike <i>Ouranopithecus</i>, the distal roots of P 3 to M 1 in <i>Anadoluvius</i> and <i>Graecopithecus</i> 11 are single fused roots with two root canals (Fig. 3 and Supplementary Table 5). Like <i>Ouranopithecus, Anadoluvius</i> has thick enamel. Supplementary Fig. 9 illustrates ranges of variation in relative enamel thickness (RET) in the M 2 of Miocene, Plio-Pleistocene and living hominoids. <i>Anadoluvius</i> has thicker enamel than most Miocene apes, falling at the upper end of the range in <i>Afropithecus</i> 30 Its RET is greater than RPl 641, an M 3 of <i>Ouranopithecus</i>. The relationship between M 2 and M 3 RET is variable in hominoids 30, but their ranges of variation always overlap. <i>Anadoluvius</i> falls well above the ranges in extant hominids and within the <i>A. afarensis</i> and <i>A. africanus</i> 75% quartiles. <i>Canine size</i>. The results of an ANOVA examining lower canine relative size is presented in Supplementary Table 7. In canine size relative to the geometric mean <i>Ouranopithecus</i> is significantly different from extant African apes in having relatively small canines. The relative size of the CO-305 mandibular canine (0.53) is equal to the mean of <i>Ouranopithecus</i> males and at the low end of the range of variation in <i>Pan</i> males and females. Canines are small in the Balkan/Anatolian sample compared with other fossil and extant apes including <i>Ardipithecus</i>, being more consistent with the ranges in <i>Australopithecus</i>. The ratio of lower canine to M 2 size (canine maximum ln x bd/M 2 maximum ln x bd) in <i>Anadoluvius</i> is lower than in any male and most females except <i>Australopithecus</i> (Supplementary Fig. 10a). The Balkan/Anatolian specimens have relatively small canines compared with African apes when scaled to the individual geometric means (Supplementary Fig. 10b and Supplementary Tables 6, 7; geometric mean of 12 variables). Scaled M 2 size is large and beyond the range of variation of African apes in <i>Graecopithecus, Anadoluvius,</i> and Nikiti 1while Ravin de la Pluie <i>Ouranopithecus</i> is intermediate between the other fossil apes and <i>Gorilla</i> (Supplementary Fig. 10c). Canine size is compared to each tooth position in Supplementary Fig. 11a, b. The results of a cladistic analysis using a data matrix of 112 characters and between 18 and 23 taxa are presented in Fig. 4, Tables 1, 2 and in the supplementary materials (Supplementary Figs. 12–14; Supplementary Data 2, 3; Supplementary Note 5). All four cladograms are strict consensus. The analysis was run both with all characters unordered and 21 of the 112 characters ordered (see Methods and Supplementary Note 5 for character matrix assumptions). Both analyses yield similar results. The tree topologies are identical, and predictably the tree values are lower in the analyses using ordered characters. All but the cladograms that include the taxa with more than 80% missing data recover a clade that includes Eurasian apes and hominines to the exclusion of pongines and stem hominids (Fig. 4). The potential problem of using data from published sources as opposed to direct observation is illustrated in the results for <i>Sahelanthropus</i>. This taxon, universally attributed to the Homininae and most commonly to the Hominini, is never recovered in these positions in these analyses, being consistently a stem hominid. <i>Sahelanthropus</i> could be coded for 72% of the characters. The potential for coding inconsistent with the criteria used to code other taxa is therefore larger than for <i>Orrorin</i>, which is also coded from the literature, but for which only 29% of the characters could be coded. We mapped synapomorphies and Bremer support values onto two consensus cladograms (Supplementary Figs. 12, 13). Table 1 lists the hominine synapomorphies of the cladogram (ordered and unordered) with the fewest missing data (18 OTUs). A phylogeny consistent with a large majority of the cladograms presented here appears in Fig. 5. Supplementary Fig. 14 shows the results of the analyses of all four taxon sets with all character states ordered. Unlike the unordered and partly ordered analyses, the cladograms with differing OTUs vary widely. The 19 OTU (including <i>Sahelanthropus</i>) fully ordered analysis is consistent with previous ones while the 18 and 20 OTU analyses result in a pongine clade includin

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