Darwin may have been wrong when he opined that the most likely place of origin of the African apes and humans was Africa, an opinion still favored by many researchers. This new research proposes that hominines not only evolved in western and central Europe but spent over five million years evolving there and spreading to the eastern Mediterranean before eventually dispersing into Africa, probably because of changing environments and diminishing forests.

 Figure 1: CO-2100/2800, a new face and partial brain case of Anadoluvius turkae, a fossil hominine (African apes and humans) from Çorakyerler (Central Anatolia, Türkiye).

            In their study, the authors analyzed the new face (figure 1) and had a second look at the other ape fossils previously described from Çorakyerler. Their reanalysis uncovered some errors in the first analysis and led them to recognize the new genus. Their new phylogenetic analysis provides the strongest evidence to date that hominines originate in Europe, expand into the Balkans and Anatolia, and eventually disperse into Africa.

 Figure 2. A consensus cladogram resulting from the analysis of 112 characters of the cranium and dentition. Fossil apes from Europe and Anatolia (from top to bottom, Anoiapithecus to Ouranopithecus) are identified as hominines along with African apes and Ardipithecus. Nakalipithecus, which is sometimes identified as hominine, is identified here as a stem hominid.

             Anadoluvius turkae is part of the evolutionary radiation that gave rise to chimpanzees, bonobos, gorillas, and humans (figure 2). Although African apes today are only known from Africa, as are the earliest known humans, the ancestors of both came from Europe and the eastern Mediterranean. The study’s detailed analysis reveals that the Balkan and Anatolian apes evolved from ancestors in western and central Europe. This contrasts with the long-held view that African apes and humans evolved exclusively in Africa. While the remains of early hominines are abundant in Europe and Anatolia (figure 3), they are completely absent from Africa until the first hominin appears there about 7 million years ago.

Figure 3. A continent full of apes in motion. Site distributions and dispersals of middle and late Miocene apes from Europe and southwest Asia. The large red arrow represents the dispersal of thickly enameled hominids into Eurasia from Africa after about 17 Ma. Afropithecus from Kenya or Helipithecus from Saudi Arabia may be broadly ancestral to the earliest hominids in Eurasia. The oldest hominids are the thickly enameled apes from Türkiye, Slovakia, and Germany (red circles). Thinner red arrows depict the dispersal of early hominids into central and western Europe later in the middle Miocene. Toward the end of the middle Miocene, the earliest hominines, which are thinly enameled (violet circles), are found in central and western Europe. In the late Miocene, more advanced hominines of differing enamel thickness evolve in situ in central and western Europe (yellow). Slightly later in time thickly enameled hominines with noticeably more robust jaws appear in the Balkans and Anatolia, probably derived from a western or central European hominine (blue arrows and circles). Notably, among the central European hominines Danuvius shares thick enamel with ouranopithecins. The grey circles and double arrow represent localities and the dispersal of the pongines between Southwest and South Asia. The green arrows represent the dispersal of hominines into Asia and Africa after 8-9 Ma. The thinner green arrow to the north represents the dispersal of a central European hominine to the Caucasus (Georgia). It also represents the possible dispersal of hominines from Europe to south China (Lufengpithecus from Yunnan province). The thicker green arrow represents the dispersal of hominines from the eastern Mediterranean to Africa. It is unclear if crown hominines subsequently evolve in situ in Africa or if individual crown hominine clades disperse separately into Africa. Taxa represented on the map are listed in table 1.

 Table 1. Taxa represented in figure 3. Colors correspond to those in figure 3. Note 1: The oldest specimen in this group is a fragmentary molar from Engelswies, Germany, tentatively attributed to Griphopithecus. Affinities with Afropithecus have also been suggested. Note 2: Austriacopithecus is known from a partial ulna and humerus and may be synonymous with Griphopithecus (which is not known from any long bones). Note 3: The sample from Nikiti 1 in Macedonia (Greece), usually attributed to Ouranopithecus, may be attributable to Anadoluvius. Note 4: Oreopithecus is of disputed affinities. We think it is unlikely to be a hominid, so it is listed as a hominoid of uncertain status.

             Anadoluvius from Türkiye and other fossil apes from nearby Greece and Bulgaria (Ouranopithecus and Graecopithecus) form a group that comes closest in details of anatomy and ecology to the earliest known hominins, or humans. The new fossils from Türkiye are the best-preserved specimens of this group of early hominines (figures 1,4).

Figure 4. Top-Frontal and lateral view of a male Anadoluvius (top) and a male Ouranopithecus (bottom). Bottom-CO 2100 in situ.

             The face and skull of Anadoluvius share attributes with other hominines, including European fossil apes, that are not found in other apes. Although smaller, they are generally most like gorillas in such details as the position of the front teeth relative to the palate, the shape of the nasal and orbital apertures, the region around the eyes and cheeks and the position of the forehead. Unlike living hominines except humans and our ancestors, Anadoluvius, along with Ouranopithecus and Graecopithecus, have powerfully built jaws with thickly enameled teeth, more like australopithecines than any other hominine. In fact, the radiation of hominines in Europe and Anatolia mirrors the subsequent radiation of hominines millions of years later in Africa. Some taxa developed thinly enameled teeth and smaller jaws, adapted to eating soft fruit or fibrous vegetation (most dryopithecins in Europe, Ardipithecus and African apes in Africa). Others evolved robust jaws, thickly enameled teeth, and powerful chewing muscles, adapted to processing tough and hard foods, many from terrestrial resources (Anadoluvius, Ouranopithecus and Graecopithecus in the eastern Mediterranean and australopithecines and early Homo in Africa).

Figure 5. One phylogeny consistent with the cladogram in figure 2. The “puddles” represent potential sources for the origin of subsequent taxa. The blue puddle contains taxa broadly descendant from the early Miocene Kenyan stem hominoid Ekembo. One of these taxa is likely to be closely related to the common ancestor of the orange (pongines) and green (hominines) puddles. The older among them (Kenyapithecus, Equatorius, and Nacholapithecus) are more likely candidates, given their age and more generalized morphology. The larger, more specialized Chororapithecus, Nakalipithecus, and Samburupithecus are most likely end-points of early or middle Miocene ape lineages. The early pongines Sivapithecus, Ankarapithecus and Khoratpithecus are broadly ancestral to the only living pongine, Pongo. Other members of this group include the largest known Miocene ape, Indopithecus and its probable descendant, the largest ape ever, Gigantopithecus. The green puddles represent the various groups of hominines. The lightest green puddle includes the thickly enameled eastern Mediterranean hominines and the brighter green puddle the more thinly enameled western and central European hominines (Danuvius is an exception, having relatively thick enamel). The line between the two represents the probable ancestral-descendant relationship between these groups. The remaining green puddles represent crown hominines. It is unclear from which group of early hominines each crown hominine taxon evolved, though we know that the gorilla clade diverged first, followed by the divergence of the chimpanzee and human clades. The branching pattern within the Homo puddle represents the myriad of fossil human lineages.

             The oldest samples attributed by most researchers to the Hominini (humans and their fossil relatives since the divergence with chimpanzees) are Sahelanthropus, Orrorin, and Ardipithecus kadabba[1], ranging in age from about 7 to 5.2 Ma. Sahelanthropus and Orrorin more closely resemble the ouranopiths in dental morphology, while Ardipithecus kadabba is more thinly enameled with less robust jaws, more like the dryopiths. So, it is unclear if the more likely ancestor of the earliest hominins is something akin to the large jawed Anadoluvius or the more gracile Rudapithecus. Only the recovery of more fossils will help to address this question.

            Anadoluvius lived in a dry forest setting and probably spent much time on the ground. The animals that lived with Anadolvius are those commonly associated with African grasslands and dry forests today (giraffes, wart hogs, rhinos, diverse antelopes, zebras, elephants, porcupines, hyaenas, lion-like carnivores). This ecological community appears to have dispersed into Africa from the eastern Mediterranean sometime after about 8 million years ago. The founding of the modern African dry forest/open country fauna from the eastern Mediterranean has long been known, and now we can add to the list of entrants the ancestors of the African apes and humans.

David R. Begun

Department of Anthropology

University of Toronto

Ayla Sevim-Erol

Department of Anthropology

Ankara University

[1] Ardipithecus kadabba is based on samples from several localities in Ethiopia dated between 5.8 and 5.2 Ma. The type species of the genus, Ardipithecus ramidus, is from another Ethiopian locality dated to 4.4 Ma. Ar. ramidus shares attributes of the canine-premolar complex with Australopithecus and Homo that are missing in Ar. kadabba, suggesting that the latter is not properly attributable to Ardipithecus. Whatever the “kadabba” hypodigm is, it is diagnostically hominin in canine and pedal phalangeal morphology.