Anadoluvius Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek 2023, gen. nov.
Creators: name: Sevim-Erol, Ayla
name: Begun, D. R.
name: Sözer, Ç. Sönmez
name: Mayda, S.
name: van den Hoek Ostende, L. W.
name: Martin, R. M. G.
name: Alçiçek, M. Cihat
Publisher: name: Zenodo
Year: 2023
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Primates
Hominidae
Anadoluvius

description: <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 including all European and Anatolian taxa, which contrasts with all previous cladistic results from analyses with large data sets and numerical cladistic methods (e.g. TNT, PAUP, etc). The 20 OTU analysis includes several other unconventional results such as an <i>Orrorin - Anadoluvius</i> clade that, along with <i>Ouranopithecus</i> as the sister to pongines, and a pongine clade that includes all European and Anatolian taxa. The 23 OTU analysis remains highly unresolved. Given the diversity of results of these all ordered analyses and the broader consistency of the analyses in which the data matrices were either unordered or partly ordered, we consider the latter to be more reliable (see Methods for a discussion of our rationale on ordering character states in this analysis). The consensus cladogram with the three taxa represented by &lt;20% of the character matrix (23 OTUs) is uninformative, given the low level of resolution. The cladograms resulting from the unordered or partly ordered analyses, which exclude these taxa (<i>Chororapithecus</i>, <i>Samburupithecus</i> and <i>Graecopithecus</i>) all recover a clade that includes <i>Anadoluvius</i> and <i>Ouranopithecus</i> as sister taxa, which in turn is either the sister taxon to the dryopithecins or in an unresolved polychotomy with the dryopithecins and the crown hominines. <i>Nakalipithecus</i>, which has been interpreted as a potential ancestor of <i>Ouranopithecus</i>, is outside the crown hominids in this analysis, as suggested elsewhere 1, 31 <i>Ankarapithecus</i> is identified as a pongine and the widely accepted <i>Pan</i> -hominin clade is supported as well. <i>Ouranopithecus, Graecopithecus</i> and <i>Anadoluvius</i> share a suite of derived characters of the jaws and dentition that support their status as a distinct clade. Although <i>Graecopithecus</i> could not be included in the analyses that yielded well resolved phylogenies, due to its many missing data (90%), all previous analyses of <i>Graecopithecus</i> associate it phylogenetically with <i>Ouranopithecus</i> 1 – 3, 5, 11, 13, 15, 32. The core attributes of the Balkan/ Anatolian late Miocene apes are large, thickly enameled molars, transversely robust mandibles, small canines, and large size. However, among these taxa there is diversity. <i>Graecopithecus</i> is distinguished from <i>Ouranopithecus</i> (<i>contra</i> 35) in its relatively large M 2 compared with both the M 1 and corpus breadth, its more vertical mandibular symphysis and in details of root morphology 1, 11, 15 <i>Anadoluvius</i> has the same lower dental root formula (P 3 to M 1) as <i>Graecopithecus</i> and both differ from <i>Ouranopithecus</i>. The frontal bone between the superior orbital margin and the anterior temporal line is preserved in XIR-1 (<i>Ouranopithecus</i>), which is sufficient to show that it was more vertically oriented than in CO-2100/2800. <i>Anadoluvius</i> is further distinguished from <i>Ouranopithecus</i> and other non-hominin hominines in having mesiodistally shorter canines (as was probably the case in <i>Graecopithecus</i> based on canine root size and shape) that lack mesial grooves and lingual cingula (unknown for <i>Graecopithecus</i>). In quantitative attributes, the PCA (Supplementary Fig. 6) illustrates the overall distinctiveness of <i>Anadoluvius</i> compared with Balkan apes. The male <i>Anadoluvius</i> mandible CO-300/305 is distinguished from male <i>Ouranopithecus</i> in many metric comparisons (canine-M 2 ratio, canine/geometric mean, relative mandibular breadth, symphyseal-molar distance, relative P 4 length and size, and relative P 4, M 1 and M 2 size (Supplementary Fig. 8a–f). <i>Anadoluvius</i> is distinguished from male <i>Ouranopithecus</i> in canine size relative to postcanine tooth size at every tooth position for both upper and lower tooth rows (Supplementary Fig. 11). <i>Anadoluvius</i> is distinguished from <i>Graecopithecus</i> in relative mandibular breadth, symphyseal-molar distance, P 4 length, M 2 relative size and M 2 size relative to corpus breadth (Supplementary Fig. 8). In addition, the mandibular arch is wider and relative mandibular breadth larger at every dental position in <i>Anadoluvius</i> (Supplementary Fig. 7). Unfortunately, <i>Nakalipithecus</i> is insufficiently preserved to be included in most of these quantitative analyses, which are scaled using the geometric mean. <i>Nakalipithecus</i> lacks one or more of the quantitative attributes needed to generate the geometric mean used in this analysis. We ran an analysis of M 2 size relative to mandibular corpus breadth at the level of mid M 1, which are data that have been published for <i>Nakalipithecus</i>. In this ratio the <i>Nakalipithecus</i> specimen, which is probably male, falls within the range of gorilla females, <i>Pan</i> of both sexes, and female <i>Ouranopithecus</i>, outside the range of gorilla males, <i>Ouranopithecus</i> males, Nikiti, <i>Graecopithecus</i>, and <i>Anadoluvius</i> (Supplementary Fig. 8f). <i>Ouranopithecus</i> and <i>Anadoluvius</i> lack shared derived characters of the pongines (greatly elongated premaxilla substantially or completely overlapping the maxillary palatine process, expanded zygoma, tall orbits, narrow interorbital space, reduced or absent ethmoidal frontal sinus, circumorbital costae.) There is no evidence for their inclusion in Ponginae. <i>Ouranopithecus</i> and <i>Anadoluvius</i> share with dryopithecins (European middle and late Miocene apes with affinities to <i>Dryopithecus</i>) a series of characters found among hominines 1 – 3 These include a ventrally rotated palate, a stepped subnasal fossa, broad, flat nasal aperture base, short nasal bones, nasal aperture apex superior to the infraorbital margins, robust lateral orbital pillars, frontal sinus expanded below nasion, incipient supraorbital torus, more horizontal frontal squama 1 – 3 The phylogenetic significance of some of these shared attributes is disputed, particularly concerning the dryopithecins 9, 24. However, there is broad agreement that <i>Ouranopithecus</i> shares enough derived characters with hominines to warrant inclusion in that taxon 1 – 3, 8 – 20 (Table 1). The phylogenetic results presented here regarding <i>Ouranopithecus</i> and its sister, <i>Anadoluvius</i>, are consistent with many previous analyses 1 – 3, 8 – 20 They are robust in terms of the number of synapomorphies and Bremer support values for many clades (Supplementary Figs. 12, 13). The results also recover the widely accepted relations among crown hominids and hominines and relations within all fossil clades. The Bremer support values for the hominid clade as defined here (15-16) are extremely strong. The hominine clade as defined here is moderate to strong (2-3), and very strong (4 to 8) for the pongine clade (Supplementary Figs. 12, 13). European and eastern Mediterranean apes are classified as hominine in three of four consensus cladograms. It fails to be resolved only in the analysis of all 23 taxa, including <i>Samburupithecus, Chororapithecus</i> and <i>Graecopithecus</i>, missing 82%, 87% and 90% of the data respectively. The latter cladogram also fails to distinguish between pongines and hominines. The relations among the dryopithecins are consistent with most detailed analyses focused on this group, as are the relations within the Asian clade (<i>Ankarapithecus, Sivapithecus</i> and <i>Pongo</i>) and the crown hominines 1 – 3, 8 – 20 The consistencies of these cladograms in many details with previous research lend credibility to these results. Recent analyses by (9 and 10) are broadly similar in their results. Among the most parsimonious cladograms reported in ref. 10 at least one also recovers a hominine clade that includes European fossil taxa9. also recovers <i>Ouranopithecus</i> as a hominine though in that analysis the dryopithecins are recovered as stem hominids. Other taxa have been linked with <i>Ouranopithecus</i> or hominines. Three fossil apes from Africa, <i>Nakalipithecus</i>, <i>Samburupithecus</i> and <i>Chororapithecus</i>, broadly overlap in time with <i>Ouranopithecus, Graecopithecus</i> and <i>Anadoluvius</i> 21 – 23 <i>Nakalipithecus</i> has been identified as potentially ancestral to <i>Ouranopithecus</i> 22. However, <i>Nakalipithecus</i> differs in many details of dental morphology from <i>Anadoluvius</i> and <i>Ouranopithecus</i> (see Supplementary Note 3). As noted, <i>Nakalipithecus</i>, represented in the data matrix by 43% of the total number of characters, consistently falls outside the crown hominids, which fails to support the hypothesis of an ancestral-descendant relationship with <i>Ouranopitheus</i>. It has been hypothesized that both <i>Chororapithecus</i> and <i>Ouranopithecus</i> have phylogenetic affinities with gorillas 8, 22, 23, 33. However, <i>Chororapithecus</i> is readily distinguished from both <i>Anadoluvius</i> and Our <i>anopithecus</i> (see differential diagnosis), and its affinity with gorillas has been questioned 1, 34 A close phylogenetic relationship with gorillas is not supported by our results21. claim that <i>Samburupithecus</i> has phylogenetic affinities with African apes and humans, though this conclusion has also been challenged by 1, 35, who conclude that <i>Samburupithecus</i> is not a hominid but instead a vestige of the early Miocene proconsuloid radiation. As with <i>Chororapithecus</i> and <i>Nakalipithecus, Samburupithecus</i> falls outside the crown hominid clade. <i>Diversity and paleobiogeography</i>. A comprehensive review of the taxonomy and phylogeny of lat
description: Published as part of <i>Sevim-Erol, Ayla, Begun, D. R., Sözer, Ç. Sönmez, Mayda, S., van den Hoek Ostende, L. W., Martin, R. M. G. &amp; Alçiçek, M. Cihat, 2023, A new ape from Türkiye and the radiation of late Miocene hominines, pp. 1-10 in Communications Biology 6 (1)</i> on pages 2-7, DOI: 10.1038/s42003-023-05210-5, http://zenodo.org/record/8279657
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