| Pachycrocuta | |
|---|---|
 | |
| Cast of theholotype skull at theMusée Crozatier | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Carnivora |
| Family: | Hyaenidae |
| Genus: | †Pachycrocuta Kretzoi 1938 |
| Species: | †P. brevirostris |
| Binomial name | |
| †Pachycrocuta brevirostris (Gervais, 1850) | |
| Possible species | |
| |
Pachycrocuta is anextinctgenus ofhyena. The largest and most well-researchedspecies isPachycrocuta brevirostris, colloquially known as thegiant short-faced hyena as it stood about 90–100 cm (35–39 in) at the shoulder[1] and it is estimated to have averaged 110 kg (240 lb) in weight,[2] approaching the size of a lioness, making it the largest known undisputed hyena, only exceeded in size by the possible hyenaDinocrocuta. It is often hypothesised to have been a specialisedkleptoparasitic 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 livingspotted 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 theEarly Pleistocene (around 2.6-2 million years ago).[3] 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.[4]
The first identified fossil of the short-faced hyena was discovered in LePuy,Auvergne, France, in 1845 by French paleontologistAuguste Aymard. In 1850, French paleontologistPaul Gervais made it theholotype specimen of a new species,Hyaena brevirostris. But, in 1893, while writing a much more detailed description, French paleontologistMarcellin Boule mistakenly listed Aymard as thespecies authority instead of Gervais, citing volume 12 of Aymard'sAnnales 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 theInternational Code of Zoological Nomenclature convincingly falsified it in 2013.[5]
The short-faced hyena was usually relegated to thegenusHyaena alongside the modernstriped hyena andbrown hyena. In 1938, Hungarian paleontologistMiklós 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, placedPachycrocuta as ancestral toCrocuta (the modern spotted hyena).[6]
Dozens more short-faced hyena remains have been found across Europe. In 1828,Jean-Baptiste Croizet andAntoine 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 fromOlivola,Tuscany, Italy, but Boule quicklysynonymized it with "H."brevirostris in 1893. In 1890, French paleontologistCharles Depéret erected "H. pyrenaica" based on a specimen fromRoussillon. Short-faced hyenas were also being discovered in East Asia. In 1870, English naturalistRichard Owen described a Chinese specimen as "H."sinensis. In 1908, French paleoanthropologistEugène Dubois described a Javan one as "H. bathygnatha". In 1934, Chinese paleoanthropologistPei Wenzhong described another Chinese one, "H."licenti, from theNihewan Basin. In 1954, mammalogist R. F. Ewer described "P."bellax" fromKromdraai, South Africa. In 1956, Finnish paleontologistBjörn Kurtén identified the subspecies "H. b. neglecta" fromJammu, India (he also chose to classify several other short-faced hyenas as subspecies ofbrevirostris.) In 1970 Ficcarelli and Torres relegated these toPachycrocuta, though "P. perrieri" is sometimes split off into a different genus,Pliocrocuta, erected by Kretzoi in 1938.[6][7] In 2001,P. brevirostris was identified inGladysvale Cave, South Africa.[8]
Usually, no more than one or two Asian short-faced hyenas were considered distinct from the EuropeanP. brevirostris. The two species convention was especially popular among Chinese scientists. As the 20th century progressed, they were often classified as regionalsubspecies ofP. brevirostris, withP. b. brevirostris endemic to Europe, andP. b. licenti andP. b. sinensis to China.[6] In 2021, Chinese paleontologist Liu Jinyi and colleagues reported the largest ever short-faced hyena skull fromJinniushan,Northeast China, belonging toP. b. brevirostris, demonstrating the subspecies is not endemic to Europe. They suggestedP. b. licenti (MiddleVillafranchian) evolved intoP. b. brevirostris (Late Villafranchian), which evolved intoP. b. sinensis (Galerian). Relict populations ofP. b. licenti seem to have persisted for some time in southern China whileP. b. brevirostris had replaced most other populations. Liu and colleagues were unsure how other supposed subspecies fit into this paradigm.[6]
The taxonomic position ofPachycrocuta relative to modern hyenas is debated. A 2008 study suggested that it was most closely related to thespotted hyena (Croctuta crocuta) among living hyenas.[9] Conversely, in a 2024 analysis of metric data for teeth, Pérez-Claros argued that thebrown hyena (Parahyaena brunnea) was its closest living relative, and along with the extinctPliocrocuta and"Hyaena" prisca should be included withinPachycrocuta.[3] A later 2025 study continued to regardPliocrocuta and the brown hyena as separate fromPachycrocuta, andP. brevirostris as the only valid Eurasian species of the genus.[10]
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Pachycroctua brevirostris was one of the largest hyenas, onlyDinocrocuta gigantea (which may be a true hyena or a member of the closely related extinct familyPercrocutidae) grew larger.[4][1][11] Two adult individuals from Zhoukoudian were estimated to be around 90 and 100 cm (3.0 and 3.3 ft) at the shoulder respectively, though some European individuals may have been somewhat larger.[1] The average individual is suggested to have been around 100 kilograms (220 lb), while very large individuals may have reached 150 kilograms (330 lb).[6] The limb bones are massively built, though the limbs bones towards the ends (distal parts) of the limbs like thetibia are particularly short, suggesting an adaption for dismembering and/or carrying carcasses.[1][2] The shortening of the distal limb bones results inP. brevirostris not being much taller than modern hyenas despite being considerably larger.[1] The skull is large and has a well developedsagittal crest, indicating well developed and largetemporal muscles in life.[1] The teeth, particularly the large premolars, and the powerfully built mandible show a strong adaption to bone cracking.[2] The fourthpremolars and firstmolars ofP. brevirostris had zigzagHunter-Schreger bands in theirenamel, indicating they were very well adapted fordurophagy.[12]
It has been suggested thatPachycrocuta lived in packs, similar to livingspotted hyenas.[1] A cache of very comprehensive bone material was unearthed at the famousZhoukoudian cave site in Northern China, which probably represents the remains of animals using these caves as lairs for many millennia.[1] At the western end of their former range, at Venta Micena in southeastern Spain, a huge assemblage of Pleistocene fossils also represents a den.[13] Yet another example exists in the Pabbi Hills ofPakistan, where remains of animals scavenged or killed byPachycrocuta were accumulated.[14] The morphology of its limbs suggests that it was less adapted to running than living spotted hyenas.[1]Pachycrocuta is often suggested to have been akleptoparasiticscavenger, using its imposing size to force other predators, such as thesabertooth catsHomotherium andMegantereon, as well as the wild dogXenocyon lycaonoides off of carcasses.[1][2] However, other authors have argued that whileP. brevirostris likely engaged in kleptoparasitism, it was likely equally capable of hunting medium-large sized prey in packs, similar to living spotted hyenas.[4] Its powerfully built limbs would likely have been effective in subduing prey.[1]P. brevirostris is known from fossil evidence found at Tsiotra Vryssi inGreece to have preferentially consumed certain bones due depending on their nutrient value.[15]
Species preyed upon and/or scavenged byPachycrocuta during the Early Pleistocene in Europe include the mammoth speciesMammuthus meridionalis,[16][17] equines (likely includingEquus altidens[2]),rhinoceroses, deer (likely includingPraemegaceros[2]) and bovids (likely includingbison[2]).[18] At the site of Cueva Victoria in southeast Spain, evidence for the consumption ofmonk seals (Monachus), has also been found.[18]
Eggs found incoprolites (fossilised feces) from Crimea indicate thatPachycrocuta brevirostris harboured a number of parasites, includingflukes,tapeworms of the genusTaenia and parasiticnematodes of the generaToxocara andCapillaria.[19]
P. brevirostris likely competed with early representatives ofHomo (archaic humans) in Early Pleistocene Europe likeHomo antecessor for carrion,[16][20] with one mammoth carcass from the Fuente Nueva-3 site in Spain showing evidence of having been consumed by both archaic humans andPachycrocuta.[17] A skull ofHomo erectus georgicus from Dmanisi, Georgia in the Caucasus displays bite marks that may be attributable toPachycrocuta.[21] Remains of "Peking Man" (an East Asian form ofHomo erectus) in the Zhoukoudian cave site display evidence of having been consumed byPachycrocuta 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.[22][23]
The oldest fossils usually considered to belong to the genus are known from thePliocene of East Africa,[4] the speciesP. bellax, known from the Early Pleistocene of South Africa, falls within the morphological variation ofP. brevirostris, though its geographical separation from the that species renders its validity equivocal.[4] It has been proposed thatP. brevirostris ultimately evolved in Asia fromPliocrocutaperrieri, which is only distinguishable fromP. brevirostris by the presence of ametaconid on the first molar. The earliest fossils ofP. brevirostris in Europe date to around 1.8-2 million years ago, with the earliest fossils in East Asia probably being slightly older.[4] The arrival ofPachycrocuta in Europe was associated with afaunal 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 theEburonian glaciation.[24][25] During its existence in Europe,Pachycrocuta brevirostris represented the only species of hyena present in the region, and largest carnivore alongside the lion-sized sabertoothHomotherium.[1]P. brevirostris became extinct in Europe around 800,000 years ago, around the time of the arrival of spotted hyenas (Crocuta crocuta) to Europe,[4] which some authors have suggested may have outcompetedPachycrocuta,[26] though there is no evidence of temporal overlap between the two species.[4] The extinction of the sabertooth catMegantereon, whose killsPachycrocuta likely scavenged, has also suggested by some authors as a factor.[27]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 tochanges in the length of glacial cycles.[4] 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.[4]
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