Pachycrocuta
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''Pachycrocuta'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
hyena Hyenas or hyaenas ( ; from Ancient Greek , ) are feliform carnivoran mammals belonging to the family Hyaenidae (). With just four extant species (each in its own genus), it is the fifth-smallest family in the order Carnivora and one of the sma ...
. The largest and most well-researched
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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 one the largest known hyenas. It is often hypothesised to have been a specialised kleptoparasitic scavenger, using its imposing size to force other predators off of carcasses, though some authors have suggested they may have been effective pack hunters like living spotted hyenas. The precise time of the origin of the genus depends on what species are included, though the only unquestioned species of the genus, ''P. brevirostris'', had emerged by the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
(around 2.6-2 million years ago). Around 800,000 years ago at the end of the Early Pleistocene, it became locally extinct in Europe, with it surviving in East Asia until at least 500,000 years ago, and possibly later elsewhere in Asia. Extinct animals of Indonesia


Taxonomy

The first identified fossil of the short-faced hyena was discovered in Le Puy,
Auvergne Auvergne (; ; or ) is a cultural region in central France. As of 2016 Auvergne is no longer an administrative division of France. It is generally regarded as conterminous with the land area of the historical Province of Auvergne, which was dis ...
, France, in 1845 by French paleontologist Auguste Aymard. 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 medic ...
made it the
holotype specimen A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was Species description, formally described. It is either the single such physical example (or illus ...
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. Car ...
mistakenly 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 (norm), convention in zoology that rules the formal scientific name, scientific naming of organisms treated as animals. It is also informally known as the I ...
convincingly falsified it in 2013. The short-faced hyena was usually relegated to the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Hyaena'' alongside the modern
striped hyena The striped hyena (''Hyaena hyaena'') is a species of hyena native to North and East Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent. It is the only extant species in the genus ''Hyaena''. It is listed by the IU ...
and
brown hyena The brown hyena (''Parahyaena brunnea''), also called the strandwolf, is a species of hyena found in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is the only extant species in the genus ''Parahyaena' ...
. In 1938, Hungarian paleontologist
Miklós Kretzoi Miklós Kretzoi (9 February 1907 – 15 March 2005) was a Hungarian people, Hungarian geologist, paleontologist and paleoanthropologist and Széchenyi Prize winner. Exhibition in the Hungarian National Museum, 9 February - 24 May 2004 Kr ...
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,
Tuscany Tuscany ( ; ) is a Regions of Italy, region in central Italy with an area of about and a population of 3,660,834 inhabitants as of 2025. The capital city is Florence. Tuscany is known for its landscapes, history, artistic legacy, and its in ...
, 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 Roussillon ( , , ; , ; ) was a historical province of France that largely corresponded to the County of Roussillon and French Cerdagne, part of the County of Cerdagne of the former Principality of Catalonia. It is part of the region of ' ...
. Short-faced hyenas were also being discovered in East Asia. In 1870, English naturalist
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
described a Chinese specimen as "''H.''" ''sinensis''. In 1908, French paleoanthropologist Eugène Dubois described a Javan one as "''H. bathygnatha''". In 1934, Chinese paleoanthropologist Pei Wenzhong described another Chinese one, "''H.''" ''licenti'', from the Nihewan Basin. In 1954, mammalogist R. F. Ewer described "''P.''" ''bellax''" from
Kromdraai Kromdraai Conservancy is a protected conservation park located to the south-west of Gauteng province in north-east South Africa. It is in the Muldersdrift area not far from Krugersdorp. Etymology Its name is derived from Afrikaans meaning "Cr ...
, South Africa. In 1956, Finnish paleontologist Björn Kurtén identified the subspecies "''H. b. neglecta''" from
Jammu Jammu () is a city in Indian-administered Jammu and Kashmir (union territory), Jammu and Kashmir in the disputed Kashmir region.The application of the term "administered" to the various regions of Kashmir and a mention of the Kashmir dispute ...
, 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, 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 In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
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,
Northeast China Northeast China () is a geographical region of China, consisting officially of three provinces Liaoning, Jilin and Heilongjiang. The heartland of the region is the Northeast China Plain, the largest plain in China with an area of over . The regi ...
, 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. The taxonomic position of ''Pachycrocuta'' relative to modern hyenas is debated. A 2008 study suggested that it was most closely related to the
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
(''Croctuta crocuta'') among living hyenas. Conversely, in a 2024 analysis of metric data for teeth, Pérez-Claros argued that the
brown hyena The brown hyena (''Parahyaena brunnea''), also called the strandwolf, is a species of hyena found in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is the only extant species in the genus ''Parahyaena' ...
(''Parahyaena brunnea'') was its closest living relative, and along with the extinct '' Pliocrocuta'' and ''"Hyaena" prisca'' should be included within ''Pachycrocuta''. A later 2025 study continued to regard ''Pliocrocuta'' and the brown hyena as separate from ''Pachycrocuta,'' and ''P. brevirostris'' as the only valid Eurasian species of the genus.


Description

''Pachycroctua brevirostris'' was one of the largest hyenas, only the percrocutid hyena, '' Dinocrocuta gigantea'' grew larger. Two adult individuals from Zhoukoudian were estimated to be around at the shoulder respectively, though some European individuals may have been somewhat larger. The average individual is suggested to have been around , while very large individuals may have reached . The limb bones are massively built, though the limbs bones towards the ends (distal parts) of the limbs like the
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
are particularly short, suggesting an adaption for dismembering and/or carrying carcasses. The shortening of the distal limb bones results in ''P. brevirostris'' not being much taller than modern hyenas despite being considerably larger. The skull is large and has a well developed
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are excepti ...
, indicating well developed and large
temporal muscles In anatomy Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It i ...
in life. The teeth, particularly the large premolars, and the powerfully built mandible show a strong adaption to bone cracking.


Behaviour and ecology

It has been suggested that ''Pachycrocuta'' lived in packs, similar to living
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
s. A cache of very comprehensive bone material was unearthed at the famous Zhoukoudian cave site in Northern China, 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. Yet another example exists in the Pabbi Hills of
Pakistan Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...
, where remains of animals scavenged or killed by ''Pachycrocuta'' were accumulated. The morphology of its limbs suggests that it was less adapted to running than living spotted hyenas. Similar to the modern day
striped hyena The striped hyena (''Hyaena hyaena'') is a species of hyena native to North and East Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent. It is the only extant species in the genus ''Hyaena''. It is listed by the IU ...
, ''Pachycrocuta'' is often suggested to have been a kleptoparasitic
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
of the kills of other predators, such as the
sabertooth cat Machairodontinae (from Ancient Greek μάχαιρα '' machaira,'' a type of Ancient Greek sword and ὀδόντος ''odontos'' meaning tooth) is an extinct subfamily of carnivoran mammals of the cat family Felidae, representing the earliest ...
s ''
Homotherium ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
'' and ''
Megantereon ''Megantereon'' is an extinct genus of prehistoric machairodontine saber-toothed cat that lived in Eurasia, Africa and possibly North America from the late Pliocene to the Middle Pleistocene. It is a member of the tribe Smilodontini, and close ...
'', as well as the wild dog '' Xenocyon lycaonoides.'' However, other authors have argued that while ''P. brevirostris'' likely engaged in kleptoparasitism, it was likely equally capable of hunting medium-large sized prey in packs, similar to living spotted hyenas. Its powerfully built limbs would likely have been effective in subduing prey. ''P. brevirostris'' is known from fossil evidence found at Tsiotra Vryssi in
Greece Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...
to have preferentially consumed certain bones due depending on their nutrient value. Species preyed upon and/or scavenged by ''Pachycrocuta'' during the Early Pleistocene in Europe include the mammoth species '' Mammuthus meridionalis'', equines (likely including '' Equus altidens''),
rhinoceros A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
es, deer (likely including '' Praemegaceros'') and bovids (likely including
bison A bison (: bison) is a large bovine in the genus ''Bison'' (from Greek, meaning 'wild ox') within the tribe Bovini. Two extant taxon, extant and numerous extinction, extinct species are recognised. Of the two surviving species, the American ...
). At the site of Cueva Victoria in southeast Spain, evidence for the consumption of monk seals ('' Monachus''), has also been found. Eggs found in coprolites (fossilised feces) from Crimea indicate that ''Pachycrocuta brevirostris'' harboured a number of parasites, including flukes,
tapeworms Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, commonly known as tapeworms. Their bodies con ...
of the genus '' Taenia'' and parasitic
nematodes The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (he ...
of the genera '' Toxocara'' and '' Capillaria.''


Relationship with humans

''P. brevirostris'' likely competed with early representatives of ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'' (archaic humans) in Early Pleistocene Europe like ''
Homo antecessor ''Homo antecessor'' (Latin "pioneer man") is an extinct species of archaic human recorded in the Spanish Archaeological Site of Atapuerca, Sierra de Atapuerca, a productive archaeological site, from 1.2 to 0.8 million years ago during the Early ...
'' for carrion, with one mammoth carcass from the Fuente Nueva-3 site in Spain showing evidence of having been consumed by both archaic humans and ''Pachycrocuta''. A skull of '' Homo erectus georgicus'' from Dmanisi, Georgia in the Caucasus displays bite marks that may be attributable to ''Pachycrocuta''. Remains of "
Peking Man Peking Man (''Homo erectus pekinensis'', originally "''Sinanthropus pekinensis''") is a subspecies of '' H. erectus'' which inhabited what is now northern China during the Middle Pleistocene. Its fossils have been found in a cave some southw ...
" (an East Asian form of ''
Homo erectus ''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
'') in the Zhoukoudian cave site display evidence of having been consumed by ''Pachycrocuta'' which includes distinctive fracturing of the skull, likely the result of biting on the facial region likely to expose the brain, as well as bones displaying signs of having been swallowed and subject to erosion by stomach acid. It is unclear whether this consumption indicates predation or scavenging.


Evolution and extinction

The oldest fossils usually considered to belong to the genus are known from the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58 the species ''P. bellax'', known from the Early Pleistocene of South Africa, falls within the morphological variation of ''P. brevirostris'', though its geographical separation from the that species renders its validity equivocal. It has been proposed that ''P. brevirostris'' ultimately evolved in Asia from ''Pliocrocuta'' ''perrieri'''','' which is only distinguishable from ''P. brevirostris'' by the presence of a
metaconid Many different terms have been proposed for features of the tooth crown in mammals. The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...
on the first molar. The earliest fossils of ''P. brevirostris'' in Europe date to around 1.8-2 million years ago, with the earliest fossils in East Asia probably being slightly older. The arrival of ''Pachycrocuta'' in Europe was associated with a faunal turnover event called the "''Pachycrocuta'' event", coinciding with considerable extinctions among the herbivore and carnivore guild and their replacement by new arrivals from elsewhere, coinciding with the onset of cold conditions of the Eburonian glaciation. During its existence in Europe, ''Pachycrocuta brevirostris'' represented the only species of hyena present in the region, and largest carnivore alongside the lion-sized sabertooth ''
Homotherium ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
''. ''P. brevirostris'' became extinct in Europe around 800,000 years ago, around the time of the arrival of spotted hyenas (''Crocuta crocuta'') to Europe, which some authors have suggested may have outcompeted ''Pachycrocuta'', though there is no evidence of temporal overlap between the two species. The extinction of the sabertooth cat ''
Megantereon ''Megantereon'' is an extinct genus of prehistoric machairodontine saber-toothed cat that lived in Eurasia, Africa and possibly North America from the late Pliocene to the Middle Pleistocene. It is a member of the tribe Smilodontini, and close ...
'', whose kills ''Pachycrocuta'' likely scavenged, has also suggested by some authors as a factor. ''Pachycrocuta'' became extinct in Europe as part of a major faunal turnover event at the Early-Middle Pleistocene ( Villafranchian-Galerian) transition in which many European megafauna species became extinct, likely as a result of a more unstable climate as a consequence to changes in the length of glacial cycles. The latest remains from East Asia are from the Zhoukoudian site, dating to around 500,000 years ago, and the species may have persisted later elsewhere in Asia.


See also

* Cave hyena (''Crocuta crocuta spelaea'') an extinct subspecies of spotted hyena alternatively considered a distinct species native to Eurasia during the Pleistocene. *'' Dinocrocuta'' an extinct genus of giant hyena-like carnivore known from the late Miocene.


External links


Blog post by paleontologist Mauricio Anton featuring a skeleton diagram and life restoration of ''Pachycrocuta''


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 Pleistocene mammals of Europe