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''Graecopithecus'' is an extinct species of
hominid The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the eas ...
that lived in southeast Europe during the late
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
around 7.2 million years ago. Originally identified by a single lower jaw bone bearing a
molar tooth The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
found in Pyrgos Vasilissis, Athens,
Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders with ...
, in 1944, other tooth specimens were discovered from Azmaka quarry in Bulgaria in 2012. With only little and badly preserved materials to reveal its nature, it is considered as "the most poorly known European Miocene
hominoids Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister ...
." The creature was popularly nicknamed 'El Graeco' (word play on the Spanish painter
El Greco Domḗnikos Theotokópoulos ( el, Δομήνικος Θεοτοκόπουλος ; 1 October 1541 7 April 1614), most widely known as El Greco ("The Greek"), was a Greek painter, sculptor and architect of the Spanish Renaissance. "El ...
) by scientists. In 2017, an international team of palaeontologists led by Madelaine Böhme of the Eberhard-Karls-University Tübingen, Germany, published a detailed analysis of the teeth and age of the specimens, and came to the conclusion that it could be the oldest hominin, meaning that it could be the oldest direct ancestors of humans after splitting from that of the chimpanzees. Their simultaneous study also claimed that contrary to the generally accepted evidences of the African origin of the hominin lineage, the ancestors of humans originated from the main ape ancestry in the
Mediterranean region In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and wa ...
(before migrating into Africa where they evolved into the ancestors of ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (modern humans), plus several extinct species classified as either ancestral to or closely related ...
'' species). They named the origin of human theory as the "North Side Story." These claims have been disputed by other scientists. Rick Potts and Bernard Wood argued that the evidence is too flimsy to even say it is a hominin. Tim D. White commented that the claim was only to support a biased argument that Africa is not the birthplace of humans; while Sergio Almécija stated that single characters such as teeth cannot tell the claimed evolutionary details. Systematic re-analysis by palaeontologists from the
University of the Witwatersrand The University of the Witwatersrand, Johannesburg (), is a multi-campus South African public research university situated in the northern areas of central Johannesburg. It is more commonly known as Wits University or Wits ( or ). The university ...
in 2017 did not find enough evidence to support the species as hominin or as the oldest ancestor of human lineage separating from the apes.


Discovery

The original ''Graecopithecus'' specimen was a single mandible found in southern Greece in 1944, "reportedly unearthed as the occupying German forces were building a wartime bunker". The original finder, German paleontologist Bruno von Freyberg initially believed that it belonged to an extinct
Old World monkey Old World monkey is the common English name for a family of primates known taxonomically as the Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboo ...
''
Mesopithecus ''Mesopithecus'' ("middle monkey" for being between ''Hylobates'' and ''Semnopithecus'' in build) is an extinct genus of Old World monkey that lived in Europe and Asia 7 to 5 million years ago. ''Mesopithecus'' resembled a modern macaque, with a ...
,'' as he reported in 1951. However,
Gustav Heinrich Ralph von Koenigswald Gustav Heinrich Ralph (often cited as G. H. R.) von Koenigswald (13 November 1902 – 10 July 1982) was a German-Dutch paleontologist and geologist who conducted research on hominins, including ''Homo erectus''. His discoverie ...
realised that it was the tooth of an ape family and erected the scientific name ''Graecopithecus freybergi'' in 1972, after the discoverer. Another tooth remain was discovered from Azmaka quarry in Bulgaria in 2012.


Description

The
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
(lower jaw) of ''Graecopithecus'' with a third molar that is very worn, the root of a second molar, and a fragment of a
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mou ...
is from a site called Pyrgos Vassilissis, northwest of Athens, and is dated from the late
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
around 7.2 million years old. Excavation of the site is not possible (as of 1986) due to the owner having built a swimming pool on the location. The thick enamel and large molars are the features that convinced von Koenigswald that the specimen belonged to a hominid species.
X-ray microtomography X-ray microtomography, like tomography and X-ray computed tomography, uses X-rays to create cross-sections of a physical object that can be used to recreate a virtual model (3D model) without destroying the original object. The prefix '' micro-' ...
and 3-dimensional reconstruction in 2017 revealed that it belonged to an adult individual and possibly a male. The partial fusion of the fourth premolar (P4) roots is an additional evidence that it is of a hominid, and the thick enamel resembles those of the human lineage (hominins).


Classification

''G. freybergi'' is considered to be possibly the same taxon as ''
Ouranopithecus macedoniensis ''Ouranopithecus macedoniensis'' is a prehistoric species of ''Ouranopithecus'' from the Late Miocene of Greece. See more detail at ''Ouranopithecus''. This species is known from three localities in Northern Greece. The type location is Ravin d ...
'', another extinct hominid described in 1977 from northern Greece. Due to paucity of specimens and poor quality of the fossils, it remains the least well known extinct hominid found within Europe. In 1984, British palaeontologists Peter Andrews and Lawrence B. Martin classified ''Graecopithecus'' and ''Ouranopithecus'' as synonyms (same taxon) and treated them as members of the genus ''
Sivapithecus ''Sivapithecus'' () (syn: ''Ramapithecus)'' is a genus of extinct apes. Fossil remains of animals now assigned to this genus, dated from 12.2 million years old in the Miocene, have been found since the 19th century in the Siwalik Hills of the In ...
.'' This classification persisted for several years until additional ''Ouranopithecus'' fossils were discovered including part of the skull in the 1990s that indicated better distinction as different hominids. Based on new evidences, in 1997, Australian palaeontologist David W. Cameron proposed renaming and inclusion of ''Ouranopithecus'' into ''Graecopithecus'' based on taxonomic priority with ''Graecopithecus macedoniensis'' as a new name for ''O. macedoniensis''. However, better ''O. macedoniensis'' specimens were found including a new species ''
Ouranopithecus turkae ''Ouranopithecus turkae'' is a prehistoric species of ''Ouranopithecus'' from the Late Miocene of Turkey. This is known from the Corakyerler locality, central Anatolia. It is known only from three cranial fossils. Dated faunal remains associate ...
'' from Turkey that supported separation of the genus, and such is continue to be generally adopted.


Re-examination and reinterpretation

In 2017, an international team of palaeontologists led by Madelaine Böhme ( Eberhard-Karls-University Tübingen, Germany) published detailed reanalysis and new interpretation in the journal '' PLOS One.'' One paper deals with an examination of the detailed morphology of molar teeth of ''G. freybergi'' from Greece and Bulgaria, and compared it with that of ''Ouranopithecus.'' The study concluded that ''Graecopithecus'' was a hominin, sharing ancestry with ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (modern humans), plus several extinct species classified as either ancestral to or closely related ...
'' but not with the chimpanzees ('' Pan''), and distinct from ''Ouranopithecus'', which has more ape-like traits. If this classification is correct, ''Graecopithecus'' would be the oldest known representative of the human lineage after the human-chimpanzee split, in 19th-century terminology, the " missing link" between human and non-human primates. The species was found to be some two hundred thousand years older than the oldest known hominid found in Africa (not necessarily ancestral to the human lineage), '' Sahelanthropus tchadensis''. The study concludes:
hedental root attributes of ''Graecopithecus'' suggest hominin affinities, such that its hominin status cannot be excluded. If this status is confirmed by additional fossil evidence, ''Graecopithecus'' would be the oldest known hominin and the oldest known crown hominine, as the evidence for the gorillin status of '' Chororapithecus'' is much weaker than the hominin status of ''Graecopithecus''. More fossils are needed but at this point it seems likely that the Eastern Mediterranean needs to be considered as just as likely a place of hominine diversification and hominin origins as tropical Africa.''''
An accompanying paper presents the study of the geological environments of the areas where the fossils were discovered. Until then, the precise date of ''Graecopithecus'' has not been resolved and usually inferred from geological data of materials related the fossils and surrounding areas that add to uncertainty in its evolutionary importance and relationship with other hominids. It is often broadly described as 6.6 to 8 million years old. The ''PLOS One'' paper resolved that the hominid lived 7.37 to 7.11 million years ago, with the specime from Greece dated to 7.18 Ma and that from Bulgaria to 7.24 Ma. It also indicates that as the species lived in Europe, it suggest "that major splits in the hominid family occurred outside Africa." It has also been proposed the ''Graecopithecus'' may not be a direct ancestor of the human lineage, but instead may have evolved its hominin-like traits independently. The emergence of ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (modern humans), plus several extinct species classified as either ancestral to or closely related ...
'' itself is dated to close to 4 million years later than ''Graecopithecus'', so that the appearance of ''Graecopithecus'' in Europe does not preclude the development of ''Homo'' proper in East Africa (as suggested by ''
Homo habilis ''Homo habilis'' ("handy man") is an extinct species of archaic human from the Early Pleistocene of East and South Africa about 2.31 million years ago to 1.65 million years ago (mya). Upon species description in 1964, ''H. habilis'' was highly c ...
'' being found in Tanzania); however, the popular press reporting on the 2017 study did cast its result in terms of determining the "birthplace of mankind". ''Graecopithecus'' lived in southeast Europe 7.2 million years ago, and if the premise of the study is correct, ''Graecopithecus'', after evolving in Europe, would have migrated to Africa about 7 million years ago where its descendants would eventually evolve into the genus ''Homo''.


Criticism

The 2017 ''PLOS One'' papers made two critical conclusions: that ''Graecopithecus'' is a hominin suggesting it as the oldest ancestor of humans after splitting from chimpazees, and that as ''Graecopithecus'' is a human ancestor, Europe is the birthplace of hominins. This directly challenges the prevailing knowledge that humans originated in East Africa. David R. Begun of the
University of Toronto The University of Toronto (UToronto or U of T) is a public research university in Toronto, Ontario, Canada, located on the grounds that surround Queen's Park. It was founded by royal charter in 1827 as King's College, the first institution ...
, Canada, one of the co-authors, was quoted as saying that " is dating allows us to move the human-chimpanzee split into the
Mediterranean area In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and wa ...
." This was set against a quote by an uninvolved anthropologist saying that " is possible that the human lineage originated in Europe, but very substantial fossil evidence places the origin in Africa ..I would be hesitant about using a single character from an isolated fossil to set against the evidence from Africa." Since 1994, Begun had adhered to a hypothesis that African hominids (including living apes) descended from Eurasian apes since the older ape fossils are found in Europa and Asia. This is a feasible explanation as it is possible that the African ape ancestors could move to Africa around 9 million years ago from Europe. However, claiming that ''Graecopithecus'' is an evidence of human origin in Europe is illogical since all human ancestral species known so far are strictly found in Africa; as Rick Potts, head of the Smithsonian’s Human Origins Program, remarked: "I think the principal claim of the main paper goes well beyond the evidence in hand... A hominin or even a hominine (modern African ape) ancestor located in a fairly isolated place in southern Europe doesn’t make much sense geographically as the ancestor of modern African apes, or particular the oldest ancestor of African hominins." David Alba at the Catalan Institute of Palaeontology in Barcelona was the first to point out that "It is not surprising at all that Begun is now arguing that hominins as well originated in Europe." Julien Benoit of the
University of the Witwatersrand The University of the Witwatersrand, Johannesburg (), is a multi-campus South African public research university situated in the northern areas of central Johannesburg. It is more commonly known as Wits University or Wits ( or ). The university ...
, Johannesburg, South Africa, also commented: "Any study that counters this consensus (Out of Africa theory) would have to provide very strong evidence and perfect methodology to support its claim. In my opinion, this article doesn't meet those criteria." Other scientists have also expressed skepticism of Begun's classification. Bernard Wood at
George Washington University , mottoeng = "God is Our Trust" , established = , type = Private federally chartered research university , academic_affiliations = , endowment = $2.8 billion (2022) , preside ...
described the hypothesis as "relatively weak" and Sergio Almécija, also at George Washington University, says it is important to bear in mind that primates seem particularly prone to evolving similar features independently. "Single characters are not reliable to make big evolutionary laims" Tim White at the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant univ ...
, asserted that the study was merely an attempt "to resurrect Begun’s tired argument with a long-known crappy fossil, newly scanned."


Reassessment

In late 2017, Julien Benoit and Francis J. Thackeray re-analysed the claims of the ''PLOS One'' papers and found key issues in the major conclusions: # The partial fusion of the fourth premolar (P4) roots does not define ''Graecopithecus'' as a hominin since the feature is common in hominids, even in the chimpanzees. # Thick enamel and relatively large molar teeth are not exclusive to hominins as they are also present in other Miocene apes and gorillas. # The claim that ''Graecopithecus'' is the ancestral ape of human lineage and that humans originate in Europe is not justified. Even if ''Graecopithecus'' is the basal (root ancestor) ape, all other human ancestral species starting from ''
Sahelanthropus ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene epoch. The species, and its genus ''Sahelanthropus'', was announced in 2002, based mainly on a partial cranium, nicknamed ''T ...
'' were in Africa, thus, still making Africa as the birthplace of humans. The study concludes:
erecognise a small signal for placing ''Graecopithecus'' at the root of the Hominini clade. This means that the phylogenetic relationship between ''Graecopithecus'' and Hominini is as yet not confirmed. Our analysis supports the view that ''Graecopithecus'' is potentially an important taxon for the origin of Hominini, but this is not certain and deserves further investigation and more material.


Response

In 2018, Fuss, Spassov, Böhme, and Begun published a response to Benoit and Thackeray, claiming that their original publication had been misrepresented and misconstrued. The conclusion of the 2017 paper had not been that ''Graecopithecus'' was certainly a hominin, but that its status as a hominin could not be ruled out, and that more research and evidence would be needed to make a conclusion—a conclusion that Benoit and Thackeray make in their own paper as well. Fuss et al. also point out that, contrary to what Benoit and Thackeray write, they did not judge canine root derivation of ''Graecopithecus'' and ''Salehanthropus'' against each other, stating that the differences between them were within the range of sexual variation. Additionally, when Benoit and Thackeray claim that the characteristics mentioned in the 2017 paper are not unique to Hominini, they do not mention that the 2017 paper discusses canine root size and premolar root complexity reduction, which could be indications of Hominini. Benoit and Thackeray also refer to taxonomy that combines ''Graecopithecus'' and ''Ouranopithecus'', despite the two generally being considered to be separate species, with ''Graecopithecus'' being more closely related to hominins than ''Ouranopithecus''. Fuss et al. emphasize that they do not present ''Graecopithecus'' as being a hominin without question, but that it has traits similar to hominids, and that more evidence is needed before ''Graecopithecus''s status as a hominin can be confirmed or rejected.


See also

* * * * * * *


References

{{Taxonbar, from=Q5592170, from2=Q32947788, from3=Q32947778, from4=Q14516253, from5=Q140198 Miocene primates of Europe Hominin fossils Prehistoric Greece Graecopithecus Prehistoric primate genera Neogene fossil record Fossil taxa described in 1972 Fossils of Serbia