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{{Short description|Extinct genus of mammals}}
{{Italic title}}
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'''''Pliohyrax'''''
[[File:PliohyraxDB22.jpg|thumb|left|[[Life restoration]]]]
Fossils of this [[Miocene]]-[[Pliocene]], [[Arboreal locomotion|scansorial]] [[herbivore]] have been found
In Spain, ''Pliohyrax graecus'' is among the large mammals species found in the [[Almenara, Castellón|Almenara]] site, deposited during the [[Messinian salinity crisis]], together with ''[[Macaque|Macaca]]'' sp., [[Bovidae]] indet., cf. ''[[Nyctereutes]]'' sp., and [[Felidae]] indet.<ref>{{Cite journal | last1 = Agustí | first1 = Jordi | last2 = Garcés | first2 = Miguel | last3 = Krijgsman | first3 = Wout | title = Evidence for African–Iberian exchanges during the Messinian in the Spanish mammalian record | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | year = 2006 | volume = 238 | issue = 1–4 | pages = 5–14 | url = http://www.geo.uu.nl/~forth/publications/Agusti_2006.pdf | doi = 10.1016/j.palaeo.2006.03.013| bibcode = 2006PPP...238....5A }}</ref>
==History==
[[File:Leptodon Gaudry 1867.png|thumb|left|
In 1853, following a field trip in [[Cyprus]], French palaeontologist and geologist [[Albert Gaudry]] stopped in [[Greece]] on his way back to [[France]], and, on the invitation of the [[List of ambassadors of France to Greece|ambassador of France to Greece]] [[Alexandre de Forth-Rouen]], visited a local fossil site at [[Pikermi]], in the [[Attica]] peninsula, near [[Athens]], first visited by German scientists. Gaudry immediatly recognized the importance of the deposits and led, in 1855 and in 1860, two excavation campaigns in Attica under the commission of the [[French Academy of Science]]. In Pikermi, Gaudry excavated the fossils of a diverse fauna whose preservation was almost unprecedented in the Neogene of Europe. Among those discoveries figured two large but isolated mandibles belonging to the same adult individual of a yet unknown large species of mammal. In a seminal book published in 1862, Gaudry described these incomplete remains under the name ''Leptodon graecus'', after the slender shape of its molars. At the time of Gaudry, [[Hyracoid]]s, [[Proboscidean]]s, [[Perissodactyl]]s and [[Artiodactyl]]s were classified together within the order [[Pachydermata]]. Gaudry speculated that ''Leptodon'' was an extinct member of a clade including the modern [[rhinoceros]] and [[hyrax]]es.<ref name=Gaudry1862>{{cite book |last1=Gaudry|first1=A. |date=1862-1867|title=Animaux fossiles et géologie de l'Attique, d'après les recherches faites en 1855, 1856 et 1860 par Albert Gaudry|publisher=F. Savy (Paris) |orig-date= |pages=474 p.}}
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===The 2000s===
In 2002, [[Qiu Zhanxiang]], [[Wei Qi]], [[Pei Shuwen]] and [[Chen Zheying]] reexamined the heavily worn holotype remains of ''Sogdohyrax'' and determined that it represented a juvenile form of a larger species of ''Pliohyrax'', that they renamed ''P. soricus''.<ref name=Qiu2002>{{cite journal|first1=Z.|last1=Qiu|first2=Q.|last2=Wei|first3=S.|last3=Pei|first4=Z.|last4=Chen|title=Preliminary report on Postschizotherium (Mammalia: Hyarcoidea) material from Tianzhen, Shanxi, China|date=2002|journal=Vertebrata PalAsiatica|volume=40(2)|pages=146-160}}</ref> In 2008, [[Bernard Ménouret]] and [[Pierre Mein]] reviewed the dentition attributed to ''P. rossignoli'' from Soblay.<ref name=Ménouret2008>{{cite journal|first1=B.|last1=Ménouret|first2=P.|last2=Mein|title=Les vertébrés du Miocène supérieur de Soblay (Ain, France)|date=2008|journal=Documents des Laboratoires de Géologie, Lyon|volume=165|pages=10-93}}</ref>
===Citations ===
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In 1995, a study by [[Gary T. Schwartz]], [[David Tab Rasmussen]] and [[Richard J. Smith]] estimated the size of extinct hyracoids based on the of their dentition and proposed that ''P. chardini'' weighed between 900 and 1432 kg.<ref name=Schwartz1995>{{cite journal|first1=G.T.|last1=Schwartz|first2=D.|last2=Tab Rasmussen|first3=R.J.|last3=Smith|title=Body-size diversity and community structure of fossil hyracoids|date=1995|journal=Journal of Mammalogy|volume=76(4)|pages=1088-1099}}</ref> In 1996, [[Zong Guanfu]] described a new species of ''Postschizotherium'', ''P. tibetensis'', from the [[Hengduan Mountains]] of [[Sichuan]], China, based on a maxilla and two mandibles, and referred to the genus an additional maxilla from the same site.<ref name=Zong1996>{{cite book |last=Zong|first=G.F.|date=1996|editor-last1=Zong|editor-first1=G.F.|editor-last2=Chen|editor-first2=W.Y.|editor-last3=Huang|editor-first3=X.S.|title=Cenozoic Mammals and Environment of Hengduan Mountains Region|pages=61-80|chapter=Hyracoidea Huxley|isbn=}}</ref>
In 2002, Qiu, [[Wei Qi]], [[Pei Shuwen]] and [[Chen Zheying]] redescribed ''P. intermedium'' on the basis of two new crania associated with their mandibles collected by Wei and Chen between 1981 and 1983 in the locality 81018, in [[Tianzhen County]], [[Shanxi]]. Additionaly, they considered that the holotype of ''P. licenti'' had been misidentified and that it was therefore a junior synonym of ''P. chardini'', and that most of the material referred to ''Postschizotherium'' by Zong in 1996 belonged in reality to [[Schizotheriinae|schizotheres]], although the maxilla designated as type for ''P. tibetensis'' was genuine.<ref name=Qiu2002>{{cite journal|first1=Z.|last1=Qiu|first2=Q.|last2=Wei|first3=S.|last3=Pei|first4=Z.|last4=Chen|title=Preliminary report on Postschizotherium (Mammalia: Hyarcoidea) material from Tianzhen, Shanxi, China|date=2002|journal=Vertebrata PalAsiatica|volume=40(2)|pages=146-160}}</ref> In 2003, [[Chen Guanfang]] erected ''P. tibetensis'' as its own genus, ''[[Hengduanshanhyrax]]''.<ref name=Chen2003>{{cite journal|first1=G.|last1=Chen|title=A new genus of Pliohyracinae (Hyracoidea, Mammalia) from the Late Pliocene of Dege, Sichuan, China|date=2003|journal=Vertebrata PalAsiatica|volume=41(3)|pages=240-248}}</ref>
In 2009, [[Nakaya Hideo]], [[Takai Masanaru]], [[Fukuchi Akira]] and [[Ogino Shintaro]] reported the first fossil remains of ''Postschizotherium'' outside of China. The material, consisting in several associated teeth, were collected near [[Udunga]], [[Buryatia]], in [[Transbaikal|Transbaikalian]] [[Russia]], and was originally described as ''Postschizotherium'' cf. ''chardini''.<ref name=Nakata2009>{{cite journal|first1=H.|last1=Nakaya|first2=M.|last2=Takai|first3=A.|last3=Fukuchi|first4=S.|last4=Ogino|title=A preliminary report on some fossil mammals (Equidae, Perissodactyla and Hyracoidea) from the Pliocene Udunga fauna, Transbaikalia, Russia|date=2009|journal=Asian Paleoprimatology|volume=5|pages=99-104}}</ref>
=Prohyrax=
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During the [[First World War]], German [[geologist]] [[Erich Kaiser (geologist)|Erich Kaiser]] led the first paleontological expedition into the Cenozoic diamond-bearing beds of [[German South West Africa]], in modern day [[Namibia]]. Subsequent researches on the area were led by [[Werner Beetz]] under Kaiser's directions. Remains coming from three distinct localities were uncovered during these expeditions and were assembled in three collections, to which [[Ernst Stromer]], well-versed in African palaeontology and geology, was given access for further studies. In 1926, Stromer published the result of these studies. Among the heavily worn fossil remains collected by Beetz in the [[Langental]] locality, in the heart of the [[Sperrgebiet]], between the [[Bogenfels]] and [[Pomona, Namibia|Pomona]] mining settlements, still in activity at the time, were a left [[maxilla]] and a single fragmentary tooth tip probably belonging to the same individual. Stromer determined that it belonged to an immature individual belonging to a new genus, short-snouted and substantially larger than any modern hyraxes, that he named ''Prohyrax tertiarius''.<ref name=Stromer1926>{{cite journal|first1=E.|last1=Stromer|title=Reste Land- und Süsswasser-Bewohnender wirbeltiere aus den Diamantfeldern Deutsch-Südwestafrikas|date=1926|journal=Die Diamantenwüste Südwest-Afrikas|volume=2|pages=107-153}}</ref> These remains were destroyed, like many others, during the widespread destructions of the [[Second World War]] ; however, casts survived.<ref name=Pickford1997>{{cite journal|first1=M.|last1=Pickford|first2=S.|last2=Moyà-Solà|first3=P.|last3=Mein|title=A revised phylogeny of Hyracoidea (Mammalia) based on new specimens of Pliohyracidae from Africa and Europe|date=1997|journal=Neues Jahrbuch für Geologie und Paläontologie|volume=205(2)|pages=265-288}}</ref> In 1954, [[Thomas Whitworth]] tentatively assigned the genus as an early-diverging member of the subfamily [[Saghatheriinae]], along with ''[[Saghatherium]]'' and ''[[Meroehyrax]]''.<ref name=Whitworth1954>{{cite journal|first1=T.|last1=Whitworth|title=The Miocene Hyracoids of East Africa with some observations on the order Hyracoidea|date=1954|journal=Fossil mammals of Africa|volume=7|pages=1-59}}</ref> In 1956, [[Charles S. Churcher]] speculated that ''P. tertiarius'' was a direct ancestor of the extant genus ''[[Procavia]]'', based on the similarities of its dentition with that of the extinct ''[[P. transvaalensis]]'' of similar age, and proposed to assign the species to the modern genus, as ''Procavia tertiaria''.<ref name=Churcher1956>{{cite journal|first1=C.S.|last1=Churcher|title=The fossil Hyracoidea of the Transvaal and Taungs deposits|date=1956|journal=Annals of the Transvaal Museum|volume=22(4)|pages=478-501}}</ref>
In 1972, [[Cary T. Madden]] reported the presence of remains tentatively assigned to ''P. tertiarius'' among Late Miocene-aged fossils collected in 1948 at [[Muruarot Hill]], near [[Lake Turkana]], in [[Kenya]] during a field expedition from the [[University of California]] led by [[H. Basil S. Cooke]].<ref name=Madden1972>{{cite journal|first1=C.T.|last1=Madden|title=Miocene mammals, stratigraphy and environment of Muruarot Hill, Kenya|date=1972|journal=Paleobios|volume=14|pages=1-12}}</ref> In 1973, [[R. J. G. Savage]] and [[William Roger Hamilton]] reported the presence of the genera in Miocene deposits of the [[Gebel Zelten]] locality.<ref name=Savage1973>{{cite journal|first1=R.J.G.|last1=Savage|first2=W.R.|last2=Hamilton|title=Introduction to the Miocene mammal faunas of Gebel Zelten, Libya|date=1973|journal=Bulletin of the British Museum (Natural History) Geology|volume=22(8)|pages=513-527}}</ref>
In 1978, [[Grant E. Meyer]] recovered it as belonging to the family [[Procaviidae]], being potentially closely related with extant hyraxes. He referred to the genus a specimen collected by [[Bryan Patterson]] in 1964 in Early Miocene deposits of the [[Turkana Grits]] near [[Loperot]], and noted that [[William Roger Hamilton]] reported the presence of the genus in [[Gebel Zelten]], in [[Cyrenaica]]. While Meyer believed that those specimens and that reported in 1972 by Madden in Muruarot likely represented at least a new species, the lack of overlapping material and the fragmentary nature of the type species limiting further speculation.<ref name=Meyer1978>{{cite book |last=Meyer|first=G.E.|date=1978|editor-last1=Maglio|editor-first1=V.J.|editor-last2=Cooke|editor-first2=H.B.S.|title=Evolution of African Mammals|publisher=Harvard University Press|pages=284-314|chapter=Hyracoidea|isbn=}}</ref>
Around 1976, numerous hyracoid remains were collected in the Miocene-aged [[Arrisdrift]] locality, near [[Oranjemund]], in the [[ǁKaras Region]] of [[Namibia]]. These specimens, including a relatively complete skull, numerous fragmentary maxilla, mandibles, isolated teeth, limb bones, belonging to all sexes and ages, were described in 1994 by [[Martin Pickford]] and assigned to a new species of ''Prohyrax'', ''P. hendeyi''. He also refered a fragmentary juvenile maxilla from Langental to ''P. tertiarius''.<ref name=Pickford1994>{{cite journal|first1=M.|last1=Pickford|title=A new species of Prohyrax (Mammalia, Hyracoidea) from the middle Miocene of Arrisdrift, Namibia|date=1994|journal=Communications of the Geological Survey of Namibia|volume=9|pages=43-62}}</ref><ref name=Pickford1996>{{cite journal|first1=M.|last1=Pickford|first2=B.|last2=Senut|first3=P.|last3=Mein|first4=D.|last4=Gommery|first5=J.|last5=Morales|first6=D.|last6=Soria|first7=M.|last7=Nieto|first8=J.|last8=Ward|title=Prelimînary results of new excavations at Arrisdrift, middle Miocène of southern Namibia|date=1996|journal=Comptes-rendus de l'Académie des Sciences de Paris|volume=322|pages=991-996}}</ref> In 1995, Pickford ''et al.'' reported the presence of ''Prohyrax'' sp. in the early Miocene AM 02 quarry of the [[Auchas]] locality, in southern [[Namibia]].<ref name=Pickford1995>{{cite journal|first1=M.|last1=Pickford|first2=B.|last2=Senut|first3=P.|last3=Mein|first4=J.|last4=Morales|first5=D.|last5=Soria|first6=M.|last6=Nieto|first7=J.|last7=Ward|first8=M.|last8=Bamford|title=The discovery of lower and middle Miocene vertebrates at Auchas, southern Namibia|date=1995|journal=Comptes-rendus de l'Académie des Sciences de Paris|volume=322|pages=901-906}}</ref> The following year, Pickford described teeth from the Middle Miocene [[Berg Aukas]] locality in northern Namibia that he tentatively referred to ''Prohyrax'' or ''[[Parapliohyrax]]''.
In 1996, [[Jan van der Made]] assigned a mandible of large hyracoid, collected in the Miocene [[Melabes|Melambes]] locality of the [[Pandánassa Formation]] in [[Crete]], and initially considered in 1973 by [[Siegfried E. Kuss]] as a new species belonging to the genus ''[[Pliohyrax]]''<ref name=Kuss1976>{{cite journal|first1=S.E.|last1=Kuss|title=Ein erster Fund von Pliohyrax aus dem Vallesian von Kreta/Griechenland|date=1976|journal=Neues Jahrbuch für Geologie und Paläontologie|volume=1|pages=1-64}}</ref>, to cf. ''P. hendeyi''.<ref name=vanderMade1996>{{cite book |last1=van der Made|first1=J.|date=1996|editor-last1=Reese|editor-first1=D.S.|title=Pleistocene and Holocene Fauna of Crete and Its First Settlers|publisher=Prehistory Press|pages=70-79|chapter=6 - Pre-Pleistocene Land Mammals from Crete|isbn=}}</ref>
In 1997, Pickford, [[Salvador Moyà-Solà]] and [[Pierre Mein]] described additional remains attributed to ''P. tertiarius'', including several maxilla, mandibles and isolated teeth from Langental and [[Elizabethfeld]]. The authors noticed the great similarities with those of ''P. hendeyi'', the only difference being the size. This additional material was instrumental to resolve its affinities with ''[[Meroehyrax]]'' and with modern [[Procaviidae]].<ref name=Pickford1997/>
In 2003, Pickford mentionned in passing that the specimen from Berg Aukas tentatively assigned to the genus belonged to its larger relative ''Parapliohyrax ngororaensis'', and estimated the size and weight of ''P. hendeyi''.<ref name=Pickford2003>{{cite journal|first1=M.|last1=Pickford|title=Giant dassie (Hyracoidea, Mammalia) from the Middle Miocene of South Africa|date=2003|journal=South African Journal of Science|volume=99|issue=7|pages=366–367}}</ref> In 2006, Pickford constated that the attribution of hyracoid remains from Gebel Zelten to the genus were unsubstantiated.<ref name=Pickford2006>{{cite journal|first1=M.|last1=Pickford|title=Giant hyracoid from basal Middle Miocene deposits at Gebel Zelten, Libya|date=2006|journal=Estudios Geológicos|volume=62|issue=1|pages=495–498}}</ref> In 2008, Pickford described additional material attributed to ''P. tertiarius'' found in the Northern Sperrgebiet, including several cranial remains from Langental, cranial and postcranial material from Elizabethfeld, and an isolated [[humerus]] from [[Grillental]] ; these new remains helped Pickford to confirm that, contrarily to Churcher's thesis, ''Prohyrax'' was not the direct ancestor of the modern ''Procavia''.<ref name=Pickford>{{cite journal|first1=M.|last1=Pickford|title=Hyracoidea from the Early Miocene of the Northern Sperrgebiet, Namibia|date=2008|journal=Memoir of the Geological Survey of Namibia|volume=20|pages=315-325}}</ref>
=Thyrohyrax=
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From 1961 to 1967, expeditions organized by [[Yale University]] to the [[Fayoum]] uncovered a rich diversity of fossil taxa from the [[Jebel Qatrani Formation]]. Among the remains were a vast array of fragmentary dentaries and associated postcranial remains belonging to a new type of hyracoid, collected between 1962 and 1967 in the quarries G, M and I. In 1973, [[Grant E. Meyer]], who had participated in the expeditions, described formally the remains as the new genus and species that he named ''Thyrohyrax domorictus'', based on the only fragmentary right mandible collected in Quarry M during the last year of the expeditions by Meyer himself. Meyer noted the great resemblance between ''Thyrohyrax'' and ''[[Meroehyrax]]'', then placed among the [[Pliohyracinae]], and proposed that ''T. domorictus'', while a [[Sagatheriinae]], was closely related to the last common ancestor of the more derived pliohyracins.<ref name=Meyer1973>{{cite journal|first1=G.E.|last1=Meyer|title=A new Oligocene hyrax from the Jebel el Qatrani Formation, Fayum, Egypt|date=1973|journal=Postilla|volume=1653|pages=1-11}}</ref> In 1978, in an overview of the fossil African hyraxes, Meyer still excluded ''Thyrohyrax'' from Pliohyracinae, but speculated that it was at least closely related in its dental anatomy to the earliest stage of a lineage leading directly to ''Pliohyrax'' through ''Meroehyrax''. He also transfered in the same chapter ''M. pygmaeus'' to the genus ''[[Pachyhyrax]]''.<ref name=Meyer1978>{{cite book |last=Meyer|first=G.E.|date=1978|editor-last1=Maglio|editor-first1=V.J.|editor-last2=Cooke|editor-first2=H.B.S.|title=Evolution of African Mammals|publisher=Harvard University Press|pages=284-314|chapter=Hyracoidea|isbn=}}</ref>
In 1988, [[David Tab Rasmussen]] and [[Elwyn L. Simons]], describing the hyracoid remains collected in the [[Jebel Qatrani Formation]] of the Fayum between 1977 and 1984, finally transfered to ''Thyrohyrax'' ''P. pygmaeus'', and referred another isolated molar to the species ; he also referred a great number of specimens to the more abundant type species ''T. domorictus'', and mentioned the presence of a third smaller species, ''T.'' sp., known from a few isolated teeth and a fragmentary mandible.<ref name=TabRasmussen1988>{{cite journal|first1=D.|last1=Tab Rasmussen|first2=E.L.|last2=Simons|title=New Oligocene hyracoids from Egypt|date=1988|journal=Journal of Vertebrate Paleontology|volume=8(1)|pages=67-83}}</ref> In 1990, Tab Rasmussen, [[Mario Gagnon]] and Simons signaled the collection of several isolated hyracoid [[astragalus (bone)|astragali]], including one of small size, found in the newly exploited quarry L-41 of the Jebel Qatrani Formation, and tentatively attributable to ''Thyrohyrax'' or ''Saghatherium''.<ref name=TabRasmussen1990>{{cite journal|first1=D.|last1=Tab Rasmussen|first2=M.|last2=Gagnon|first3=E.L.|last3=Simons|title=Taxeopody in the carpus and tarsus of Oligocene Pliohyracidae (Mammalia: Hyracoidea) and the phyletic position of Hyraxes|date=1990|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=87|pages=4688-4691}}</ref> The next year, 1991, Tab Rasmussen and Simons described, among several new hyracoid taxa, two new species of ''Thyrohyrax'' from Quarry L-41 : the smaller ''T. meyeri'', based on CGM 41882, a partial mandible, to which they referred several fragmentary mandibles, a partial skull, and speculated that some postcranial elements found in the same quarry belonged to the new species, the name honouring Grant E. Meyer, the founder of the genus ; and the larger ''T. litholagus'', based on CGM 41880, a partial mandible, to which they referred four other mandibles, two maxilla and two cranial fragments, the species name, based on the greek prefix "'lithos'" ("rock"), and the suffix "'lagos'", ("rabbit"), meaning "rock rabbit", a translation of the common name used for hyraxes by modern Egyptians.<ref name=TabRasmussen1991>{{cite journal|first1=D.|last1=Tab Rasmussen|first2=E.L.|last2=Simons|title=The Oldest Egyptian Hyracoids (Mammalia: Pliohyracidae): New Species of Saghatherium and Thyrohyrax from the Fayum|date=1991|journal=Neues Jahrbuch für Geologie und Paläontologie|volume=182(2)|pages=187-209}}</ref> In 1995, a study by [[Gary T. Schwartz]], Tab Rasmussen and [[Richard J. Smith]] estimated the size of extinct hyracoids based on the of their dentition and proposed that the smaller ''T. meyeri'', weighing between 4.6 and 7.1 kg, was barely larger than modern hyraxes ; ''T. domorictus'' weighed between 6.9 and 12.6 kg, and the larger ''T. litholagus'' weighed between 13.7 and 23.1 kg.<ref name=Schwartz1995>{{cite journal|first1=G.T.|last1=Schwartz|first2=D.|last2=Tab Rasmussen|first3=R.J.|last3=Smith|title=Body-size diversity and community structure of fossil hyracoids|date=1995|journal=Journal of Mammalogy|volume=76(4)|pages=1088-1099}}</ref>
The genus name, ''Thyrohyrax'', is formed from the [[Ancient Greek|Greek]] prefix "''thyra-''", meaning "window", and the suffix "''-hyrax''", referring to the internal fenestra in the mandible of the presumed females. The type species name is formed from the Greek "''domos-''", meaning "house" and the [[Latin]] ''-ramus'', meaning "jaws", referring to the chambered jaws of the type species.<ref name=Meyer1973/>
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'''''Naranbulagornis''''' is an
==History and Etymology==
The [[holotype]] and [[paratype]] remains of ''Narambulagornis'' were collected in 1970 - 1971 during the [[Joint Russian-Mongolian Paleontological Expedition]] in the
The genus name is formed from the prefix "''Naranbulag''", designing the name of the type locality, meaning "sunshine spring" in [[Mongolian language|Mongolian]], and the suffix "-''ornis''", meaning "bird". The species name, "''khun''", means "[[swan]]" in Mongolian.
==Description==
''Naranbulagornis'' is a quite large
==Paleontology==
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