View source for Dryopithecus

{{good article}}
{{Use British English|date=November 2019}}
{{short description|Extinct great ape from Europe}}
{{Automatic taxobox	
| fossil_range = [[Serravallian]]–[[Tortonian]] <br />{{Fossil range|12.5|11.1}}
| image = Dryopithecus fontani mio med francia.JPG
| image_caption = Mandible fragment of ''D. fontani'' from Saint-Gaudens, France (Middle Miocene, 11.5 mya); cast from [[National Museum of Natural History (France)|Museum national d'histoire naturelle]], Paris
| taxon = Dryopithecus
| authority = [[Édouard Lartet|Lartet]], 1856<ref name=Lartet1856>{{cite journal|first=É.|last=Lartet|authorlink=Édouard Lartet|year=1856|title=Note sur un grand Singe fossile qui se rattache au groupe des Singes Supérieurs|language=French|trans-title=Note on the large ape fossils related to the great apes|journal=Comptes Rendus de l'Académie des Sciences de Paris|volume=43|pages=219–223}}</ref>
| subdivision_ranks = Species	
| subdivision = 
*†''D. fontani''<ref name=Lartet1856/> <br /><small>[[Édouard Lartet|Lartet]], 1856</small>	
*?†''D. carinthiacus''<ref name=Mottl1957/> <br /><small>[[Von Maria Mottl|Mottl]], 1957</small>	
| subdivision_ref = <ref name=Begun2018/>	
}}

'''''Dryopithecus''''' is a [[genus]] of [[extinct]] [[great ape]]s from the [[middle Miocene|middle]]–[[late Miocene]] boundary of Europe 12.5 to 11.1 [[million years ago]] (mya). Since its discovery in 1856, the genus has been subject to taxonomic turmoil, with numerous new species being described from single remains based on minute differences amongst each other, and the fragmentary nature of the [[holotype specimen]] makes differentiating remains difficult. There is currently only one uncontested species, the [[type species]] '''''D. fontani''''', though there may be more. The genus is placed into the [[tribe (biology)|tribe]] [[Dryopithecini]], which is either an offshoot of [[orangutan]]s, [[Homininae|African apes]], or is its own separate branch.

A male specimen was estimated to have weighed {{cvt|44|kg|lbs}} in life. ''Dryopithecus'' likely predominantly ate ripe fruit from trees, suggesting a degree of [[suspensory behaviour]] to reach them, though the anatomy of a [[humerus]] and [[femur]] suggest a greater reliance on walking on all fours ([[quadrupedalism]]). The face was similar to [[gorilla]]s, and males had longer canines than females, which is typically correlated with high levels of [[agonistic behaviour|aggression]]. They lived in a seasonal, paratropical climate, and may have built up fat reserves for winter. European great apes likely went extinct during a drying and cooling trend in the Late Miocene which caused the retreat of warm-climate forests.

==Etymology==
The [[genus name]] ''Dryopithecus'' comes from [[Ancient Greek]] ''drus'' 'oak tree' and ''pithekos'' '[[ape]]' because the [[author citation (zoology)|authority]] believed it inhabited an [[oak]] or [[pine]] forest in an environment similar to modern day Europe.<ref>{{cite book|first=D. W.|last=Csmeron|year=2004|title=Hominid Adaptations and Extinctions|publisher=UNSW Press|pages=138–139|isbn=978-0-86840-716-6}}</ref> The species ''D. fontani'' was named in honour of its discoverer, local collector Monsieur Alfred Fontan.<ref name=Lartet1856/>

==Taxonomy==
[[File:DryopithecusDB15.jpg|thumb|left|upright|Restoration of ''D. fontani'' with an [[orangutan]]-like build]]
The first ''Dryopithecus'' fossils were [[species description|described]] from the French [[Pyrenees]] by French paleontologist [[Édouard Lartet]] in 1856,<ref name=Lartet1856/> three years before [[Charles Darwin]] published his ''[[On the Origin of Species]]''. Subsequent authors noted similarities to modern African [[great ape]]s. In his ''[[The Descent of Man]]'', Darwin briefly noted that ''Dryopithecus'' casts doubt on the African origin of apes:
{{blockquote|
...it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject; for two or three anthropomorphous apes, one the Dryopithecus of Lartet, nearly as large as a man, and closely allied to [[gibbon|Hylobates]], existed in Europe during the Miocene age; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.<ref>{{cite book|first=C.|last=Darwin|authorlink=Charles Darwin|year=1875|title=The Descent of Man|publisher=D. Appleton and Company|page=199|title-link=wikisource:The Descent of Man (Darwin)/Chapter VI}}</ref>
}}

''Dryopithecus'' taxonomy has been the subject of much turmoil, with new specimens being the basis of a new species or genus based on minute differences, resulting in several now-defunct species.<ref name=Begun2018/> By the 1960s, all non-[[human]] apes were classified into the now-[[paraphyly|obsolete]] [[family (biology)|family]] [[Pongidae]], and extinct apes into [[Dryopithecidae]].<ref name=Begun2018>{{cite book|first=D. R.|last=Begun|year=2018|title=''Dryopithecus''|pages=1–4|publisher=Wiley Online Library|doi=10.1002/9781118584538.ieba0143|chapter=Dryopithecus|isbn=9781118584422|s2cid=240097940}}</ref> In 1965, English palaeoanthropologist [[David Pilbeam]] and American palaeontologist [[Elwyn L. Simons]] separated the genus–which included specimens from across the [[Old World]] at the time–into three [[subgenera]]: ''Dryopithecus'' in Europe, ''Sivapithecus'' in Asia, and ''Proconsul'' in Africa. Afterwards, there was discussion over whether each of these subgenera should be elevated to genus. In 1979, ''[[Sivapithecus]]'' was elevated to genus, and ''Dryopithecus'' was subdivided again into the subgenera ''Dryopithecus'' in Europe, and ''Proconsul'', ''Limnopithecus'', and ''Rangwapithecus'' in Africa.<ref>{{cite book|first1=F. S.|last1=Szalay|first2=E.|last2=Delson|year=1979|title=Evolutionary History of the Primates|url={{google books|plainurl=yes|id=jE7gBAAAQBAJ|page=470}}|publisher=Academic Press|pages=470–490|isbn=978-1-4832-8925-0}}</ref> Since that time, several more species were assigned and moved, and by the 21st century, the genus included ''D. fontani'', ''[[Rudapithecus|D. brancoi]]'',<ref>{{cite journal|first1=G.|last1=Merceron|first2=E.|last2=Schulz|first3=L.|last3=Kordos|first4=T. M.|last4=Kaiser|year=2007|title=Paleoenvironment of ''Dryopithecus brancoi'' at Rudabánya, Hungary: evidence from dental meso- and micro-wear analyses of large vegetarian mammals|journal=Journal of Human Evolution|volume=53|issue=4|pages=331–349|doi=10.1016/j.jhevol.2007.04.008|pmid=17719619|bibcode=2007JHumE..53..331M }}</ref><ref>{{cite journal|first1=L.|last1=Kordos|first2=D. R.|last2=Begun|s2cid=14955805|year=2001|title=A new cranium of Dryopithecus from Rudabánya, Hungary|journal=Journal of Human Evolution|volume=41|issue=6|pages=689–700|doi=10.1006/jhev.2001.0523|pmid=11782114|bibcode=2001JHumE..41..689K }}</ref> ''[[Hispanopithecus|D. laietanus]]'',<ref>{{cite journal|first1=S.|last1=Moyà-Solà|authorlink=Salvador Moyà-Solà|first2=M.|last2=Köhler|year=1996|title=A ''Dryopithecus'' skeleton and the origins of great-ape locomotion|journal=Nature|volume=379|issue=6,561|pages=156–159|doi=10.1038/379156a0|pmid=8538764|bibcode=1996Natur.379..156M|s2cid=4344346}}</ref> and ''[[Hispanopithecus|D. crusafonti]]''.<ref>{{cite journal|first=D. R.|last=Begun|year=1992|title=''Dryopithecus crusafonti'' sp. nov., a new Miocene Hominoid species from Can Ponsic (northeastern Spain)|journal=American Journal of Physical Anthropology|volume=87|issue=3|pages=291–309|doi=10.1002/ajpa.1330870306|s2cid=41299778 }}</ref> However, the 2009 discovery of a partial skull of ''D. fontani'' caused many of them to be split off into different genera, such as the newly erected ''[[Hispanopithecus]]'', because part of the confusion was caused by the fragmentary nature of the ''Dryopithecus'' [[holotype]] with vague and incomplete diagnostic characteristics.<ref name=MoyaSola2009>{{cite journal|first1=S.|last1=Moyà-Solà|authorlink=Salvador Moyà-Solà|first2=M.|last2=Köhler|first3=D. M.|last3=Alba|year=2009|title=First partial face and upper dentition of the Middle Miocene hominoid ''Dryopithecus fontani'' from Abocador de Can Mata (Vallès-Penedès Basin, Catalonia, NE Spain): taxonomic and phylogenetic implications|journal=American Journal of Physical Anthropology|volume=139|issue=2|pages=126–145|doi=10.1002/ajpa.20891|pmid=19278017|doi-access=free}}</ref><ref name=Begun2018/>

[[File:Dryopithecus carinthiacus.png|thumb|The lower [[molar tooth|molars]] of "''D. carinthiacus''"]]
Currently, there is only one uncontested species, ''D. fontani''. Specimens are:
* Holotype MNHNP AC 36, three pieces of a male mandible with teeth from [[Saint-Gaudens, Haute-Garonne|Saint-Gaudens]] in the French Pyrenees.<ref name=Begun2018/><ref>{{cite journal|first1=C.|last1=Casanovas-Vilar|first2=D. M.|last2=Alba|first3=M.|last3=Garcés|first4=J. M.|last4=Robles|first5=S.|last5=Moyà-Solà|authorlink5=Salvador Moyà-Solà|year=2011|title=Updated chronology for the Miocene hominoid radiation in Western Eurasia|journal=Proceedings of the National Academy of Sciences|volume=108|issue=14|pages=5554–5559|doi=10.1073/pnas.1018562108|pmid=21436034|bibcode=2011PNAS..108.5554C|doi-access=free|pmc=3078397}}</ref> Based on dental development in [[chimpanzee]]s, it was 6 to 8 years old, and several diagnostic characteristics made from the holotype would be lost in mature ''D. fontani'';<ref>{{Cite journal| last1 = Simons | first1 = E. L.| last2 = Meinel | first2 = W.| title = Mandibular ontogeny in the Miocene great ape ''Dryopithecus''| journal = International Journal of Primatology| volume = 4| issue = 4| pages = 331–337| year = 1983| doi = 10.1007/BF02735598| s2cid = 26442894}}</ref> A partial left [[humerus]] arm bone, an additional mandible (MNHNP 1872-2), a left lower jaw and five isolated teeth are also known from the site.<ref name=Pilbeam1971/> 
*An upper [[incisor]], NMB G.a.9., and female upper [[molar tooth|molar]], FSL 213981, come from [[Saint-Alban-de-Roche]], France.<ref>{{cite journal|first1=M. P.|last1=de los Ríos|first2=D. M.|last2=Alba|first3=S.|last3=Moyà-Solà|authorlink3=Salvador Moyà-Solà|year=2013|title=Taxonomic attribution of the La Grive hominoid teeth|journal=American Journal of Physical Anthropology|volume=151|issue=4|pages=558–565|doi=10.1002/ajpa.22297|pmid=23754569}}</ref> 
*A male partial face, IPS35026, and femur, IPS41724, from Vallès Penedès in [[Catalonia]], Spain.<ref name=MoyaSola2009/><ref name=Begun2018/> 
*A female mandible with teeth, LMK-Pal 5508, from St. Stefan, [[Carinthia]], Austria 12.5 mya, which could possibly be considered a separate species, "''D. carinthiacus''".<ref name=Begun2018/><ref name=Fuss2018/><ref name=Mottl1957>{{cite journal|url=https://www.zobodat.at/pdf/CAR_147_67_0039-0084.pdf|first=V. M.|last=Mottl|year=1957|title=Bericht über die neuen Menschenaffenfunde aus Österreich, von St. Stefan im Lavanttal, Kärnten|language=German|trans-title=Report on new apes from Austria, from St. Stefan im Lavanttal, Carinthia|journal=Carinthia II|volume=67|pages=39–84}}</ref>

''Dryopithecus'' is classified into the namesake great ape [[tribe (biology)|tribe]] [[Dryopithecini]], along with  ''[[Hispanopithecus]]'', ''[[Rudapithecus]]'', ''[[Ouranopithecus]]'', ''[[Anoiapithecus]]'', and ''[[Pierolapithecus]]'', though the latter two may belong to ''Dryopithecus'',<ref name=Begun2018/> the former two may be synonymous, and the former three can also be placed into their own tribes.<ref name=Rios2014>{{cite thesis|first=M. P.|last=de los Ríos|s2cid=90027032|year=2014|title=The craniodental anatomy of Miocene apes from the Vallès-Penedès Basin (Primates: Hominidae): Implications for the origin of extant great apes|publisher=Universitat Autònoma de Barcelona|type=PhD|pages=20–21}}</ref> Dryopithecini is either regarded as an offshoot of [[orangutan]]s ([[Ponginae]]),<ref name=Alba2012>{{cite journal|first=D. M.|last=Alba|year=2012|title=Fossil apes from the Vallès-Penedès basin|journal=Evolutionary Anthropology|volume=21|issue=6|pages=254–269|doi=10.1002/evan.21312|pmid=23280922|s2cid=39830696}}</ref> an ancestor to African apes and humans ([[Homininae]]),<ref name=Begun2005>{{cite journal|first=D. R.|last=Begun|year=2005|title=''Sivapithecus'' is east and ''Dryopithecus'' is west, and never the twain shall meet|journal=Anthropological Science|volume=113|issue=1|pages=53–64|doi=10.1537/ase.04S008|url=https://www.jstage.jst.go.jp/article/ase/113/1/113_1_53/_pdf/-char/en|citeseerx = 10.1.1.88.1617}}</ref><ref>{{cite journal|first=D. R.|last=Begun|year=2009|title=Dryopithecins, Darwin, de Bonis, and the European origin of the African apes and human clade|journal=Geodiversitas|volume=31|issue=4|pages=789–816|doi=10.5252/g2009n4a789|s2cid=131688092|url=http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/g2009n4a7.pdf}}</ref><ref>{{cite journal|first1=D. R.|last1=Begun|first2=M. C.|last2=Nargolwalla|first3=L.|last3=Kordos|year=2012|title=European Miocene hominids and the origin of the African ape and human clade|journal=Evolutionary Anthropology|volume=21|issue=1|pages=10–23|doi=10.1002/evan.20329|pmid=22307721|s2cid=22792031}}</ref> or its own separate branch ([[Dryopithecinae]]).<ref name=Rios2014/><ref>{{cite book|first1=M.|last1=Köhler|first2=S.|last2=Moyà-Solà|authorlink2=Salvador Moyà-Solà|first3=D. M.|last3=Alba|year=2001|chapter=Eurasian hominoid evolution in the light of recent ''Dryopithecus'' findings|title=Hominoid evolution and climatic change in Europe|volume=2|publisher=Cambridge University Press|editor1-first=L.|editor1-last=de Bonis|editor2-first=G. D.|editor2-last=Koufos|editor3-first=P.|editor3-last=Andrews|isbn=978-0-521-66075-4|chapter-url=https://books.google.com/books?id=jmpwQA5ZPLsC&pg=PA192}}</ref>

''Dryopithecus'' was a part of an [[adaptive radiation]] of [[great ape]]s in the expanding forests of Europe in the warm climates of the Miocene Climatic Optimum, possibly descending from [[early Miocene|early]] or [[middle Miocene|middle]] Miocene African apes which diversified in the proceeding [[Middle Miocene disruption]] (a cooling event). It is possible great apes first evolved in Europe or Asia, and then migrated down into Africa.<ref name=Begun1992>{{cite journal|first=D. R.|last=Begun|year=1992|title=''Dryopithecus crusafonti'' sp. nov., a new Miocene Hominoid species from Can Ponsic (northeastern Spain)|journal=American Journal of Physical Anthropology|volume=87|issue=3|pages=291–309|doi=10.1002/ajpa.1330870306|s2cid=41299778 }}</ref><ref name=Begun2018/><ref>{{cite journal|first1=D. R.|last1=Begun|first2=M. C.|last2=Nargolwolla|first3=M. P.|last3=Hutchinson|s2cid=189860406|year=2006|title=Primate evolution in the Pannionian Basin: In situ evolution, dispersals, or both?|journal=Beiträge zur Paläontologie|volume=30|pages=43–56}}</ref>

==Description==
Based on measurements of the [[femoral head]] of the Spanish IPS41724, the living weight for a male ''Dryopithecus'' was estimated to be {{cvt|44|kg|lbs}}.<ref name=MoyaSola2009/>

''Dryopithecus'' teeth are most similar to those of modern chimps. The teeth are small and have a thin [[tooth enamel|enamel]] layer. ''Dryopithecus'' has a slender jaw, indicating it was not well-suited for eating abrasive or hard food. Like modern apes, the males have pronounced canine teeth.<ref name=Alba2001/><ref name=Alba2012/> The [[molar tooth|molars]] are wide, and the [[premolar]]s wider. It has a wide [[palate|roof of the mouth]], a long [[muzzle (anatomy)|muzzle]] ([[prognathism]]), and a large nose which is oriented nearly vertically to the face. In total, the face shows many similarities to the [[gorilla]]; since early to middle Miocene African apes do not share such similarities, gorilla-like features likely evolved independently in ''Dryopithecus'' rather than as a result of close [[affinity (taxonomy)|affinities]].<ref name=MoyaSola2009/>

The humerus, measuring an approximate {{cvt|265|mm|in}}, is similar in size and form to the [[bonobo]]. Like in bonobos, the [[diaphysis|shaft]] bows outward, and the [[Anatomical terms of muscle#Insertion|insertion]] for the [[triceps]] and [[Deltoid muscle|deltoids]] was poorly developed, suggesting  ''Dryopithecus'' was not as adept to [[suspensory behaviour]] as orangutans.<ref name=Pilbeam1971>{{cite journal | last1 = Pilbeam | first1 = D.|authorlink=David Pilbeam| last2 = Simons | first2 = E. L.|authorlink2=Elwyn L. Simons| title = Biological sciences: humerus of ''Dryopithecus'' from Saint Gaudens, France| journal = Nature | volume = 229 | issue = 5,284 | pages = 406–407 |year= 1971| doi = 10.1038/229406a0 | pmid=4926991| bibcode = 1971Natur.229..406P| s2cid = 953761}}</ref> The [[femoral neck]], which connects the femoral head to the [[femoral shaft]], is not very long nor steep; the femoral head is positioned low to the [[greater trochanter]]; and the [[lesser trochanter]] is positioned more towards the backside. All these characteristics are important in the mobility of the [[hip]] joint, and indicate a [[quadrupedal]] mode of locomotion rather than suspensory.<ref name=MoyaSola2009/> However, fruit trees in the time and area of the Austrian ''Dryopithecus'' were typically {{cvt|5|to|12|m|0}} high and bore fruit on thinner terminal branches, suggesting suspensory behaviour to reach them.<ref name=Fuss2018/>

==Paleobiology==
[[File:500px photo (55874722).jpeg|thumb|[[Arbutus unedo|Strawberry tree]]s (above) may have been a plentiful and thus important food source in St. Stefan.<ref name=Fuss2018/>]]
''Dryopithecus'' likely predominantly ate fruit ([[frugivory]]), and evidence of [[tooth decay|cavities]] on the teeth of the Austrian ''Dryopithecus'' indicates a high-sugar diet, likely deriving from ripe fruits and honey.<ref name=Fuss2018>{{cite journal|first1=J.|last1=Fuss|first2=G.|last2=Uhlig|first3=M.|last3=Böhme|year=2018|title=Earliest evidence of caries lesion in hominids reveal sugar-rich diet for a Middle Miocene dryopithecine from Europe|journal=PLOS ONE|volume=13|issue=8|pages=e0203307|doi=10.1371/journal.pone.0203307|pmc=6117023|pmid=30161214|bibcode=2018PLoSO..1303307F|doi-access=free}}</ref> Dental wearing indicates ''Dryopithecus'' ate both soft and hard food, which could either indicate they consumed a wide array of different foods, or they ate harder foods as a fallback.<ref>{{cite journal|first1=D.|last1=DeMiguel|first2=D. M.|last2=Alba|first3=S.|last3=Moyà-Solà|authorlink3=Salvador Moyà-Solà|year=2014|title=Dietary Specialization during the Evolution of Western Eurasian Hominoids and the Extinction of European Great Apes|journal=PLOS ONE|volume=9|issue=5|pages=e97442|doi=10.1371/journal.pone.0097442|pmc=4029579|pmid=24848272|bibcode=2014PLoSO...997442D|doi-access=free}}</ref> Nonetheless, its unspecialized teeth indicate it had a flexible diet, and large body size would have permitted a large gut to aid in the processing of less-digestible food, perhaps stretching to include foods such as leaves ([[folivory]]) in times of famine like in modern apes. Unlike modern apes, ''Dryopithecus'' likely had a high [[carbohydrate]], low [[fibre]] diet.<ref name=Fuss2018/>

A high-[[fructose]] diet is associated with elevated levels of [[uric acid]], which is neutralized by [[uricase]] in most animals except great apes. It is thought they stopped producing it by 15 mya, resulting in increased [[blood pressure]], which in turn led to increased activity, and a greater ability to build up fat reserves. The palaeoenvironment of late Miocene Austria indicates an abundance of fruiting trees and [[honey]] for nine or ten months out of the year, and ''Dryopithecus'' may have relied on these fat reserves during the late winter. High uric acid levels in the blood are also associated with increased intelligence.<ref name=Fuss2018/>

''Dryopithecus'' males had larger canines than females, which is associated with high levels of aggression in modern primates.<ref name=Alba2001>{{cite journal|first1=D. M.|last1=Alba|first2=S.|last2=Mayà-Solà|first3=M.|last3=Köhler|year=2001|title=Canine reduction in the miocene hominoid ''Oreopithecus bambolii'': behavioural and evolutionary implications|journal=Journal of Human Evolution|volume=40|issue=1|pages=1–16|doi=10.1006/jhev.2000.0439|pmid=11139357|bibcode=2001JHumE..40....1A }}</ref>

==Paleoecology==
[[File:Late Miocene Europe.jpg|thumb|300px|A map of late Miocene Europe]]
The remains of ''Dryopithecus'' are often associated with several large mammals, such as proboscideans (e. g., though not limited to, ''[[Gomphotherium]]''), rhinoceroses (e. g., ''[[Lartetotherium]]''), suids (e. g., ''[[Listriodon]]''), bovids (e. g., ''[[Miotragocerus]]''), equids (e. g., ''[[Anchitherium]]''), [[hyaenids]] (e. g., ''[[Protictitherium]]''), and felids (e. g., ''[[Pseudaelurus]]''). Other associated primates are the great apes ''Hispanopithecus'', ''Anoiapithecus'', and ''Pierolapithecus''; and the [[Pliopithecidae|pliopithecid ape]] ''[[Pliopithecus]]''.<ref name=Casanovas2016>{{cite journal|first1=I.|last1=Casanovas-Vilar|first2=A.|last2=Madern|first3=D. M.|last3=Alba|year=2016|title=The Miocene mammal record of the Vallès-Penedès Basin (Catalonia)|journal=Comptes Rendus Palevol|volume=15|issue=7|pages=791–812|doi=10.1016/j.crpv.2015.07.004|bibcode=2016CRPal..15..791C |doi-access=free|hdl=2445/164290|hdl-access=free}}</ref><ref>{{cite journal|first1=P.|last1=Mein|first2=L.|last2=Ginsburg|year=2002|title=Sur l'âge relatif des différents dépôts karstiques miocènes de La Grive-Saint-Alban (Isère)|journal=Cahiers scientifiques|volume=5|issue=2|pages=7–47|doi=10.3406/mhnly.2002.1328|s2cid=192653053|language=French|trans-title=On the relative age of the different karstic Miocene deposits of La Grive-Saint-Alban (Isère)|url=https://www.persee.fr/doc/mhnly_1627-3516_2002_num_5_2_1328?pageid=t1_32}}</ref> These fauna are consistent with a warm, forested, [[tropical rainforest|paratropical]] wetland environment,<ref name=Casanovas2016/><ref>{{cite journal|first=M.|last=Gross|year=2008|title=A limnic ostracod fauna from the surroundings of the Central Paratethys (Late Middle Miocene/Early Late Miocene; Styrian Basin; Austria)|journal= Palaeogeography, Palaeoclimatology, Palaeoecology|volume=264|issue=3|pages=263–276|doi=10.1016/j.palaeo.2007.03.054|bibcode=2008PPP...264..263G }}</ref> and it may have lived in a seasonal climate. For the Austrian ''Dryopithecus'', plants such as ''[[Prunus]]'', [[grapevine]]s, [[black mulberry]], [[Arbutus unedo|strawberry tree]]s, [[hickory]], and [[chestnut]]s may have been important fruit sources; and the latter two, oak, [[beech]], [[elm]], and [[pine]] honey sources.<ref name=Fuss2018/>

The late Miocene was the beginning of a drying trend in Europe. Increasing seasonality and [[dry spell]]s in the Mediterranean region and the emergence of a [[Mediterranean climate]] likely caused the replacement of forestland and woodland by open [[shrubland]]; and the uplift of the [[Alps]] caused tropical and warm-climate vegetation in Central Europe to retreat in favor of mid-latitude and [[alpine climate|alpine]] flora. This likely led to the extinction of great apes in Europe.<ref>{{cite journal|first1=G.|last1=Merceron|first2=T. M.|last2=Kaiser|first3=D. S.|last3=Kostopoulos|first4=E.|last4=Schulz|year=2010|title=Ruminant diets and the Miocene extinction of European great apes|journal=Proceedings of the Royal Society B|volume=277|issue=1697|pages=3105–3112|doi=10.1098/rspb.2010.0523|pmid=20519220|doi-access=free|pmc=2982054}}</ref>

==See also==
{{Commons category}}
{{Wikispecies}}
{{columns-list|colwidth=30em|
* [[Timeline of human evolution]]
* ''[[Anoiapithecus]]''
* ''[[Chororapithecus]]''
* ''[[Hispanopithecus]]''
* ''[[Nakalipithecus]]''
* ''[[Oreopithecus]]''
* ''[[Pierolapithecus]]''
* ''[[Samburupithecus]]''
* ''[[Rangwapithecus]]''
* ''[[Danuvius guggenmosi|Danuvius]]''
}}

==References==
{{Reflist|30em}}

{{Haplorhini|Ho.}}
{{Portal bar|Paleontology}}
{{Taxonbar|from=Q131637}}
[[Category:Dryopithecini]]
[[Category:Prehistoric apes]]
[[Category:Miocene primates]]
[[Category:Fossil taxa described in 1856]]
[[Category:Prehistoric primate genera]]
[[Category:Miocene primates of Europe]]