Pachycrocuta
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''Pachycrocuta'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of prehistoric hyenas. The largest and most well-researched
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated to have averaged in weight, approaching the size of a lioness, making it the largest known hyena. ''Pachycrocuta'' first appeared during the late
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
( Messinian, 7.2 to 5.3 million years agoA. Hill, G. Curtis, and R. Drake. 1986. Sedimentary stratigraphy of the Tugen Hills, Baringo, Kenya. Geological Society of America Special Publication 25:285-295). By 800,000 years ago, it became locally extinct in Europe and became completely extinct during the middle
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
, 400,000 years ago.


Taxonomy

The first identified fossil of the short-faced hyena was discovered in Le Puy,
Auvergne Auvergne (; ; oc, label= Occitan, Auvèrnhe or ) is a former administrative region in central France, comprising the four departments of Allier, Puy-de-Dôme, Cantal and Haute-Loire. Since 1 January 2016, it has been part of the new region Au ...
, France, in 1845 by French paleontologist
Auguste Aymard Auguste Aymard (5 December 1808 – 26 June 1889) was a French prehistorian and palaeontologist who lived and died in Puy-en-Velay Le Puy-en-Velay (, literally ''Le Puy in Velay''; oc, Lo Puèi de Velai ) is the prefecture of the Haute-Loire ...
. In 1850, French paleontologist
Paul Gervais Paul Gervais full name François Louis Paul Gervais (26 September 1816 – 10 February 1879) was a French palaeontologist and entomologist. Biography Gervais was born in Paris, where he obtained the diplomas of doctor of science and of medicine ...
made it the
holotype specimen A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
of a new species, '' Hyaena brevirostris''. But, in 1893, while writing a much more detailed description, French paleontologist
Marcellin Boule Pierre-Marcellin Boule (1 January 1861 – 4 July 1942), better known as merely Marcellin Boule, was a French palaeontologist, geologist, and anthropologist. Early life and education Pierre-Marcellin Boule was born in Montsalvy, France. Care ...
mistakingly listed Aymard as the species authority instead of Gervais, citing volume 12 of Aymard's ''Annales de la Société d'Agriculture, Sciences, Arts et Commerce du Puy'' which does not mention the species at all. Boule further gave the annal's publication date as 1846 instead of the correct 1848. The fallacious authority Aymard, 1846, was reprinted for over a century until Spanish paleontologist David M. Alba and colleagues on behalf of the
International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the ...
convincingly falsified it in 2013. The short-faced hyena was usually relegated to the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Hyaena'' alongside the modern striped hyena and brown hyena. In 1938, Hungarian paleontologist Miklós Kretzoi suggested erecting a new genus for it, ''Pachycrocuta'', but this only became popular after Giovanni Ficcarelli and Danilo Torres' review of hyena classification in 1970. They, like many priors, placed ''Pachycrocuta'' as ancestral to '' Crocuta'' (the modern spotted hyena). Dozens more short-faced hyena remains have been found across Europe. In 1828, Jean-Baptiste Croizet and Antoine Claude Gabriel Jobert created the species "''H. perrieri''" for a specimen from Montagne de Perrier, France. In 1889, German paleontologist Karl Weithofer described "''H. robusta''" based on a specimen from
Olivola Olivola ( pms, Aulìvola) is a ''comune'' (municipality) in the Province of Alessandria in the Italian region Piedmont, about east of Turin and about northwest of Alessandria. Olivola borders the municipalities of Casorzo, Frassinello Monferrat ...
,
Tuscany it, Toscano (man) it, Toscana (woman) , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = Citizenship , demographics1_footnotes = , demographics1_title1 = Italian , demogra ...
, Italy, but Boule quickly synonymized it with "''H.''" ''brevirostris'' in 1893. In 1890, French paleontologist
Charles Depéret Charles Jean Julien Depéret (25 June 1854 – 18 May 1929) was a French geologist and paleontologist. He was a member of the French Academy of Sciences, the Société géologique de France
erected "''H. pyrenaica''" based on a specimen from Roussillon. Short-faced hyenas were also being discovered in East Asia. In 1870, English naturalist Richard Owen described a Chinese specimen as "''H.''" ''sinensis''. In 1908, French paleoanthropologist
Eugène Dubois Marie Eugène François Thomas Dubois (; 28 January 1858 – 16 December 1940) was a Dutch paleoanthropologist and geologist. He earned worldwide fame for his discovery of ''Pithecanthropus erectus'' (later redesignated ''Homo erectus''), or "Java ...
described a Javan one as "''H. bathygnatha''". In 1934, Chinese paleoanthropologist
Pei Wenzhong PEI or Pei may refer to: Places *Matecaña International Airport, Pereira, Colombia, IATA code PEI * Pei County (沛县), Jiangsu, China *Pei Commandery (沛郡), a commandery in Chinese history *Prince Edward Island, a province of Canada *Pei, ...
described another Chinese one, "''H.''" ''licenti'', from the Nihewan Basin. In 1954, mammalogist R. F. Ewer described "''P.''" ''bellax''" from Kromdraai, South Africa. In 1956, Finnish paleontologist Björn Kurtén identified the subspecies "''H. b. neglecta''" from
Jammu Jammu is the winter capital of the Indian union territory of Jammu and Kashmir. It is the headquarters and the largest city in Jammu district of the union territory. Lying on the banks of the river Tawi, the city of Jammu, with an area of ...
, India (he also chose to classify several other short-faced hyenas as subspecies of ''brevirostris''.) In 1970 Ficcarelli and Torres relegated these to ''Pachycrocuta'', though "''P. perrieri''" is sometimes split off into a different genus, '' Pliocrocuta'', erected by Kretzoi in 1938. In 2001, ''P. brevirostris'' was identified in
Gladysvale Cave Gladysvale Cave is a fossil-bearing breccia filled cave located about northeast of the well-known South African hominid-bearing sites of Sterkfontein and Swartkrans and about north-northwest of Johannesburg, South Africa. It is situated within ...
, South Africa. Usually, no more than one or two Asian short-faced hyenas were considered distinct from the European ''P. brevirostris''. The two species convention was especially popular among Chinese scientists. As the 20th century progressed, they were often classified as regional subspecies of ''P. brevirostris'', with ''P. b. brevirostris'' endemic to Europe, and ''P. b. licenti'' and ''P. b. sinensis'' to China. In 2021, Chinese paleontologist Liu Jinyi and colleagues reported the largest ever short-faced hyena skull from
Jinniushan Jinniushan () is a Middle Pleistocene paleoanthropological site, dating to around 260,000 BP, most famous for its archaic hominin fossils. The site is located near Yingkou, Liaoning, China. Several new species of extinct birds were also discovere ...
,
Northeast China Northeast China or Northeastern China () is a geographical region of China, which is often referred to as "Manchuria" or "Inner Manchuria" by surrounding countries and the West. It usually corresponds specifically to the three provinces east of ...
, belonging to ''P. b. brevirostris'', demonstrating the subspecies is not endemic to Europe. They suggested ''P. b. licenti'' (Middle Villafranchian) evolved into ''P. b. brevirostris'' (Late Villafranchian), which evolved into ''P. b. sinensis'' ( Galerian). Relict populations of ''P. b. licenti'' seem to have persisted for some time in southern China while ''P. b. brevirostris'' had replaced most other populations. Liu and colleagues were unsure how other supposed subspecies fit into this paradigm.


Fossils

The oldest specimens of ''Pachycrocuta'' were found in the late Miocene of
Baringo County Baringo County is one of the 47 counties in Kenya. It is located in the former Rift Valley Province. Its headquarters and largest town is Kabarnet. The county is home to Lake Baringo. Geographical location Baringo bounded by Turkana County an ...
(
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi ...
).
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
remains have been found broadly in
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago ...
and southern and eastern
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
. Most material consists of fragmented remains, usually of the skull, but a cache of very comprehensive bone material was unearthed at the famous
Zhoukoudian Zhoukoudian Area () is a town and an area located on the east Fangshan District, Beijing, China. It borders Nanjiao and Fozizhuang Townships to its north, Xiangyang, Chengguan and Yingfeng Subdistricts to its east, Shilou and Hangcunhe Towns t ...
site, which probably represents the remains of animals using these caves as lairs for many millennia. At the western end of their former range, at Venta Micena in southeastern Spain, a huge assemblage of Pleistocene fossils also represents a den. Other proposed species, ''P. robusta'' and ''P. pyrenaica'', are less well researched; the former may simply be an exceptionally large European paleosubspecies of the brown hyena (''Parahyaena brunnea''). Sometimes included in this genus (as ''Pachycrocuta bellax'') is the extinct giant striped hyaena, ''Hyaena bellax''.


Behaviour

Similar to the modern day striped hyena, it was probably primarily a
kleptoparasitic Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when f ...
scavenger of the kills of other predators, such as sabertooth cats. ''Pachycrocuta'' scavenged for food, probably preferentially so, because it was a heavyset animal not built for chasing prey over long distances. In this respect it would have differed from the spotted hyena of today, which is a more nimble animal that, contrary to its image as a scavenger, usually kills its own food, but often gets displaced by lions. Apparently it was ecologically close enough to its smaller (but still large) relative '' Pliocrocuta perrieri'' that they are never found as contemporary fossils in the same region. Research by anthropologists Noel Boaz and Russell Ciochon on remains of '' Homo erectus'' unearthed alongside ''Pachycrocuta'' at the Zhoukoudian site attributed scoring and puncture patterns observed on
hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The ...
long bones and skulls—originally thought to be signs of cannibalism—to predation by ''Pachycrocuta''. It has been proposed that ''Pachycrocuta'' was outcompeted and driven to extinction by the spotted hyena, which was formerly present in Eurasia as well as Africa.Kurtén, Björn (1988) On evolution and fossil mammals, Columbia University Press, pp. 238–242, Other predators, such as lions, cave lions,
tiger The tiger (''Panthera tigris'') is the largest living cat species and a member of the genus ''Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, it primarily preys on ...
s, and
wolves The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
, could have put pressure on ''Pachycrocuta''.


See also

* Cave hyena (''Crocuta crocuta spelaea'')


Notes


References

* Raoul J. Mutter, Lee R. Berger, Peter Schmid. 2001
New evidence of the giant hyena, Pachycrocuta brevirostris (Carnivora, Hyaenidaae), from the Gladyslave cave deposit (Plio-Pleistocene, John Nash Nature Reserve, Gauteng, South Africa)
Palaeont. afr., 37, 103-113 {{Taxonbar, from=Q135094 Prehistoric hyenas Pliocene carnivorans Pleistocene carnivorans Pleistocene genus extinctions Cenozoic mammals of Africa Fossils of China Cenozoic mammals of Asia Cenozoic mammals of Europe Prehistoric carnivoran genera Pliocene first appearances