Anadoluvius, Sevim-Erol & Begun & Sözer & Mayda & van den Hoek Ostende & Martin & Alçiçek, 2023
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publication ID |
https://doi.org/ 10.1038/s42003-023-05210-5 |
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DOI |
https://doi.org/10.5281/zenodo.8279632 |
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persistent identifier |
https://treatment.plazi.org/id/03D487E7-FF9B-0F5A-FCC0-FEB98E7CFE84 |
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treatment provided by |
Tatiana |
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scientific name |
Anadoluvius |
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gen. nov. |
Anadoluvius gen. nov.
Synonomy. Ouranopithecus Bonis and Melentis : Güleç et al. 28
Type species. Anadoluvius turkae comb. nov. Sevim Erol et al. 2023.
Etymology. Anadolu is the modern Turkish word for Anatolia and Anatolian.
Holotype. CO-205, a fragmented but largely complete male palate with LI1-M3 and RC-M2 GoogleMaps (Supplementary Figs. 1 View Fig , 2 View Fig ).
Paratypes. CO-300 (RM 2); GoogleMaps CO-305 (male mandibular fragment with RC-M 1); GoogleMaps CO-710 (female mandibular fragment with RP 3 - M 2); CO-2100 (RI 1); GoogleMaps 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 View Fig ; Supplementary Figs. 3–5 View Fig View Fig View Fig ) GoogleMaps
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 View Table 1 , 2 View Table 2 . Measurements are provided in Supplementary Tables 3, 4. Supplementary Note 4 presents the results of a comprehensive quantitative analysis of the Anadoluvius hypodigm. Supplementary Note 5 provides details of the phylogenetic analysis. Supplementary Note 6 provides historical, geological and biochronologic background.
Cranium. The CO-2100/2800 partial cranium ( Fig. 1 View Fig , Supplementary Figs. 2 View Fig , 3 View Fig ) 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 Ouranopithecus (XIR-1), which is broken just beyond the superior orbital margins and preserves almost nothing of the frontal squama (Supplementary Fig. 2 View Fig ). Though damaged, the position and orientation of the premaxilla is better preserved in CO-2100/2800 than in CO-205 and Ouranopithecus (XIR-1 and RPl 128), confirming previous interpretations of a stepped and overlapping morphology in these specimens 1 – 3, 11, 13 – 15 ( Fig. 2 View Fig ).
The premaxilla of Anadoluvius is short and vertical compared with Pan , Pongo , and australopithecines, and is most like Gorilla and dryopithecins, being relatively short in the alveolar portion but expanded nasally to overlap with the palatine process of the maxilla ( Fig. 2 View Fig ) The incisor alveoli are positioned along the mesial transverse plane of the canine crowns ( Figs. 1 View Fig , 2 View Fig ; Supplementary Fig. 5 View Fig ) In two specimens of Ouranopithecus (RPl 128 and XIR 1) the upper incisors are well anterior to the canines (Supplementary Fig. 5 View Fig ) 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 Anadoluvius . The frontal bone of Anadoluvius differs strongly from that of Ouranopithecus in the smooth biconvex squama of the former, contrasting with a broad concavity above glabella in the latter. The superior orbital margins of Ouranopithecus are broad, rounded and slightly projecting while they are sharp and flat in Anadoluvius .
Mandible. A principal components analysis based on mandibular measurements available for Graecopithecus , Ouranopithecus and Anadoluvius is presented in Supplementary Fig. 6 and Supplementary Data 1). The Çorakyerler, Nikiti 1 and Graecopithecus mandibles are separated from each other, especially along PC 2, and from Ouranopithecus , illustrating the diversity present in these samples. Supplementary Data 1 includes the data matrix, summary statistics, scores, and loadings.
Anadoluvius , like Graecopithecus , and NKT 21, has a relatively narrow mandible compared with the combined sex sample of Ouranopithecus (Supplementary Fig. 7a). Supplementary Fig. 7b compares relative mandibular corpus breadth at each tooth position (P 3 -M 2) in Anadoluvius , Graecopithecus , and Ouranopithecus . Anadoluvius 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). Anadoluvius is distinct from Graecopithecus in all mandibular and dental ratios. Anadoluvius falls beyond the range of variation of Ouranopithecus 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 Ouranopithecus 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 Ouranopithecus (Supplementary Fig. 8a, c). In Graecopithecus the symphysis is positioned closest to the molars, just barely in the range of the Ravin sample. In summary, in most quantitative comparisons Anadoluvius is distinguished from Ouranopithecus and Graecopithecus .
Tooth roots and enamel thickness. Figure 3 View Fig shows the root and root canal morphology of CO-300, the male mandible of Anadoluvius (see Methods for segmentation details). Unlike Ouranopithecus , the distal roots of P 3 to M 1 in Anadoluvius and Graecopithecus 11 are single fused roots with two root canals ( Fig. 3 View Fig and Supplementary Table 5).
Like Ouranopithecus , Anadoluvius 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. Anadoluvius has thicker enamel than most Miocene apes, falling at the upper end of the range in Afropithecus 30 Its RET is greater than RPl 641, an M 3 of Ouranopithecus . The relationship between M 2 and M 3 RET is variable in hominoids 30, but their ranges of variation always overlap. Anadoluvius falls well above the ranges in extant hominids and within the A. afarensis and A. africanus 75% quartiles.
Canine size. The results of an ANOVA examining lower canine relative size is presented in Supplementary Table 7. In canine size relative to the geometric mean Ouranopithecus 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 Ouranopithecus males and at the low end of the range of variation in Pan males and females.
Canines are small in the Balkan/Anatolian sample compared with other fossil and extant apes including Ardipithecus , being more consistent with the ranges in Australopithecus . The ratio of lower canine to M 2 size (canine maximum ln x bd/M 2 maximum ln x bd) in Anadoluvius is lower than in any male and most females except Australopithecus (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 Graecopithecus , Anadoluvius , and Nikiti 1while Ravin de la Pluie Ouranopithecus is intermediate between the other fossil apes and Gorilla (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 View Fig , Tables 1 View Table 1 , 2 View Table 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 Sahelanthropus . 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. Sahelanthropus 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 Orrorin , 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 Sahelanthropus ) 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 Orrorin - Anadoluvius clade that, along with Ouranopithecus 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 <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 ( Chororapithecus , Samburupithecus and Graecopithecus ) all recover a clade that includes Anadoluvius and Ouranopithecus 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. Nakalipithecus , which has been interpreted as a potential ancestor of Ouranopithecus , is outside the crown hominids in this analysis, as suggested elsewhere 1, 31 Ankarapithecus is identified as a pongine and the widely accepted Pan -hominin clade is supported as well.
Ouranopithecus , Graecopithecus and Anadoluvius share a suite of derived characters of the jaws and dentition that support their status as a distinct clade. Although Graecopithecus could not be included in the analyses that yielded well resolved phylogenies, due to its many missing data (90%), all previous analyses of Graecopithecus associate it phylogenetically with Ouranopithecus 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. Graecopithecus is distinguished from Ouranopithecus (contra 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 Anadoluvius has the same lower dental root formula (P 3 to M 1) as Graecopithecus and both differ from Ouranopithecus . The frontal bone between the superior orbital margin and the anterior temporal line is preserved in XIR-1 ( Ouranopithecus ), which is sufficient to show that it was more vertically oriented than in CO-2100/2800. Anadoluvius is further distinguished from Ouranopithecus and other non-hominin hominines in having mesiodistally shorter canines (as was probably the case in Graecopithecus based on canine root size and shape) that lack mesial grooves and lingual cingula (unknown for Graecopithecus ).
In quantitative attributes, the PCA (Supplementary Fig. 6) illustrates the overall distinctiveness of Anadoluvius compared with Balkan apes. The male Anadoluvius mandible CO-300/305 is distinguished from male Ouranopithecus 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). Anadoluvius is distinguished from male Ouranopithecus in canine size relative to postcanine tooth size at every tooth position for both upper and lower tooth rows (Supplementary Fig. 11). Anadoluvius is distinguished from Graecopithecus 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 Anadoluvius (Supplementary Fig. 7). Unfortunately, Nakalipithecus is insufficiently preserved to be included in most of these quantitative analyses, which are scaled using the geometric mean. Nakalipithecus 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 Nakalipithecus . In this ratio the Nakalipithecus specimen, which is probably male, falls within the range of gorilla females, Pan of both sexes, and female Ouranopithecus , outside the range of gorilla males, Ouranopithecus males, Nikiti, Graecopithecus , and Anadoluvius (Supplementary Fig. 8f).
Ouranopithecus and Anadoluvius 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.
Ouranopithecus and Anadoluvius share with dryopithecins (European middle and late Miocene apes with affinities to Dryopithecus ) 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 Ouranopithecus shares enough derived characters with hominines to warrant inclusion in that taxon 1 – 3, 8 – 20 ( Table 1 View Table 1 ).
The phylogenetic results presented here regarding Ouranopithecus and its sister, Anadoluvius , 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 Samburupithecus , Chororapithecus and Graecopithecus , 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 ( Ankarapithecus , Sivapithecus and Pongo ) 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 Ouranopithecus as a hominine though in that analysis the dryopithecins are recovered as stem hominids.
Other taxa have been linked with Ouranopithecus or hominines. Three fossil apes from Africa, Nakalipithecus , Samburupithecus and Chororapithecus , broadly overlap in time with Ouranopithecus , Graecopithecus and Anadoluvius 21 – 23 Nakalipithecus has been identified as potentially ancestral to Ouranopithecus 22. However, Nakalipithecus differs in many details of dental morphology from Anadoluvius and Ouranopithecus (see Supplementary Note 3). As noted, Nakalipithecus , 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 Ouranopitheus. It has been hypothesized that both Chororapithecus and Ouranopithecus have phylogenetic affinities with gorillas 8, 22, 23, 33. However, Chororapithecus is readily distinguished from both Anadoluvius and Our anopithecus (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 Samburupithecus has phylogenetic affinities with African apes and humans, though this conclusion has also been challenged by 1, 35, who conclude that Samburupithecus is not a hominid but instead a vestige of the early Miocene proconsuloid radiation. As with Chororapithecus and Nakalipithecus , Samburupithecus falls outside the crown hominid clade.
Diversity and paleobiogeography. A comprehensive review of the taxonomy and phylogeny of late Miocene apes is needed, given recent discoveries and reinterpretations. Here we focus on the diversity and paleobiogeographic implications of Anadoluvius (Supplementary Fig. 15; Supplementary Table 8). Anadoluvius and Ouranopithecus share attributes with other European middle and late Miocene hominids that distinguish them from late Miocene ape fossils from Africa and the broadly contemporaneous pongines Sivapithecus , Ankarapithecus , and Khoratpithecus (Supplementary Note 3). Ouranopithecus from Ravin de la Pluie and Xirochori are dated to 9.6 and 9.3 Ma respectively 6, 7. Ouranopithecus from Nikiti 1 is dated to 8.9 Ma 6, 7. Çorakyerler is dated to 8.7 Ma (Supplementary Figs. 16, 17; Supplementary Table 9 and Supplementary Note 6). The Nikiti mandible and maxilla are distinct from Ouranopithecus from the more northern Macedonian sites and may represent a different taxon 13. This possibility, which needs further study, is interesting in terms of regional evolution as Nikiti 1 is likely to be slightly older than Çorakyerler 6, 7, 29. Nikiti, Anadoluvius and Graecopithecus are distinguished from Ouranopithecus in our PCA and in having a greater degree of canine reduction, elongated P 4 and large molars relative to mandibular corpus size. Anadoluvius and Graecopithecus are distinguished from Ouranopithecus in root morphology (unknown in NKT 21.) NKT 21 and Graecopithecus have inferior transverse tori positioned posterior to the mesial edge of the M 1 (unknown in Anadoluvius ), while the torus is anterior to the M 1 in Ouranopithecus . Graecopithecus , which is considerably younger (~7.2 Ma) 4, has been shown to differ from Ouranopithecus in morphology that replicates differences between late Miocene apes and early hominins, such as reduced relative canine size and premolar root morphology 11 However, the Nikiti specimens have not previously been included in these comparisons.
Quantitative and qualitative comparisons reveal many differences among Balkan/Anatolian taxa, indicative of a greater diversity of late Miocene eastern Mediterranean hominids than previously recognized. Hominines appear to have been present and diverse for millions of years in the late Miocene of Europe. Based on the large number of qualitative and quantitative differences among the samples from Macedonia, Attica and Anatolia, we conclude that they represent at least three hominine genera, Ouranopithecus , Graecopithecus , and Anadoluvius . The diversity of hominines in the eastern Mediterranean mirrors that among australopithecines in the Plio-Pleistocene hominin record in Africa. In the phylogenetic analysis presented here the Balkan/ Anatolian taxa are in the sister clade of crown hominines. The fact that dryopithecins are also classified as hominines in this analysis suggests that there was an in situ evolution of thickly enameled late Miocene eastern Mediterranean hominines from more thinly enameled precursors in central and western Europe, though this conclusion has been challenged 9, 24. Pierolapithecus, Anoiapithecus , Dryopithecus , Hispanopithecus and Rudapithecus all share attributes with extant hominines and are distinguished from pongines such as Sivapithecus and Ankarapithecus ( Tables 1 View Table 1 , 2 View Table 2 ) 1 – 3, 36, 37, (but see refs. 9, 24 for alternative views). A clade that includes both thinly and thickly enameled taxa, in this case the dryopithecins and the Balkan/Anatolian apes, has a parallel in Africa with Ardipithecus and australopithecines and with Pan and hominins.
The parallel evolution of thickly and thinly enameled members of a clade in Africa and Europe is not proof that the late Miocene European apes are all hominines, but it does make this hypothesis, supported by the results of our cladistic analysis, plausible. It is possible that the generally more thinly enameled dryopithecins and the later occurring thickly enameled Balkan/ Anatolian hominines do not share an ancestor-descendant relationship and represent separate dispersal events into Europe from Africa (e.g. ref. 22), though this is less parsimonious biogeographically and contrasts with the results of the phylogenetic analysis presented here. While independent dispersal events are possible, we regard the in situ European hypothesis as more likely and more parsimonious given the current evidence. Other independent lines of evidence are also consistent with the widespread presence of hominines in Europe 38.
Table 1 Hominine and pongine synapomorphies (18 taxa, unordered).
| Hominine | Pongine |
|---|---|
| Supraorbital torus | Supraorbital margin |
| Frontal sinus/glabella | Lateral orbital pillar surface |
| Ethmoidal sinus | Nasal bone length |
| I 1 marginal ridge shape | Nasal bone breath |
| C implantation | Clivus orientation |
| Upper premolar length | C inclination |
| Lower molar buccal cristids | Upper premolar crests |
| P 3 hypoproto-postprotocristid | |
Table 2 Hominine and pongine synapomorphies (18 taxa, ordered).
| Hominine | Pongine |
|---|---|
| Supraorbital torus | Supraorbital margin |
| Frontal sinus/glabella | Interorbital space |
| Ethmoidal sinus | Lateral orbital pillar surface |
| I 1 marginal ridge shape | Nasal bone length |
| C implantation | Nasal bone breath |
| Upper premolar length | Clivus length |
| Lower molar buccal cristids | Clivus orientation |
| Supraorbital torus | C inclination |
| Incisive canal | |
| Incisive foramen size | |
| Upper premolar crests | |
| P 3 hypoproto-postprotocristid | |
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