View source for Hyena

{{Short description|Family of carnivoran mammals}}
{{About|the family of animals|other uses|Hyena (disambiguation)}}
{{Automatic taxobox
| name = Hyenas
| fossil_range = {{fossil range|22|0}} [[Early Miocene]] – recent
| image = Hyaenidae Diversity.jpg
| image_caption = The four extant species of hyena, clockwise from upper left: spotted hyena (''[[Crocuta crocuta]]''), brown hyena (''[[Parahyaena brunnea]]''), aardwolf (''[[Proteles cristata]]''), and striped hyena (''[[Hyaena hyaena]]'')
| taxon = Hyaenidae
| authority = [[John Edward Gray|Gray]], 1821
| type_genus = ''[[Hyaena]]''
| type_genus_authority = [[Mathurin Jacques Brisson|Brisson]], 1762
| subdivision_ranks = [[Genus|Genera]]
| subdivision = * '''''[[Incertae sedis]]'''''
** †''[[Tongxinictis]]''
* '''Hyaeninae'''
** †''[[Chasmaporthetes]]''
** †''[[Allohyaena]]''
** †''[[Leecyaena]]''
** †''[[Ikelohyaena]]''
** †''[[Belbus]]''
** †''[[Metahyaena]]''
** †''[[Werdelinus]]''
** '''Hyaenini'''
*** ''[[Crocuta]]''
*** ''[[Hyaena]]''
*** ''[[Parahyaena]]''
*** †''[[Adcrocuta]]''
*** †''[[Pachycrocuta]]''
*** †''[[Pliocrocuta]]''
*** †''[[Palinhyaena]]''
*** †''[[Hyaenictis]]''
* '''Protelinae'''
** ''[[Proteles]]''
** †''[[Gansuyaena]]''
* †'''Ictitheriinae'''
** †''[[Ictitherium]]''
** †''[[Protictitherium]]''
** †''[[Thalassictis]]''
** †''[[Lycyaena]]''
** †''[[Plioviverrops]]''
** †''[[Tungurictis]]''
** †'''Hyaenotheriini'''
*** †''[[Hyaenotherium]]''
*** †''[[Miohyaenotherium]]''
* †'''Percrocutinae'''?
** †''[[Percrocuta]]''?
** †''[[Dinocrocuta]]''?
| range_map = Hyaenidae_range.png
| range_map_alt = [[Striped hyena]] in blue, [[Brown hyena]] in purple, [[Spotted hyena]] in green, [[Aardwolf]] in red
| range_map_caption = Range of [[striped hyena]] in blue, [[brown hyena]] in purple, [[spotted hyena]] in green, and [[aardwolf]] in red
| synonyms = * Protelidae <small>[[William Henry Flower|Flower]], 1869</small>
* [[Percrocutidae]]? <small>Werdelin & Solounias, 1991</small>
}}

[[File: Aardwolf Skeleton.jpg|thumb|Aardwolf, smallest member of the hyena family, skeleton ([[Museum of Osteology]])]]

'''Hyenas''' or '''hyaenas''' ({{IPAc-en|h|aɪ|ˈ|iː|n|ə|z}} {{Respell|hi|EE|nəz}}; from [[Ancient Greek]] {{lang|grc|ὕαινα}}, {{Transliteration|grc|hýaina}})<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Du%28%2Faina^ ὕαινα], Henry George Liddell, Robert Scott, ''A Greek-English Lexicon'', on Perseus. Etymologically, this is a feminine of ὕς "swine", [https://www.etymonline.com/word/hyaena possibly due to its bristles].</ref> are [[feliformia|feliform]] [[carnivora]]n mammals belonging to the [[Family (taxonomy)|family]] '''Hyaenidae''' ({{IPAc-en|h|aɪ|ˈ|ɛ|n|ᵻ|d|iː}} {{IPA|la|i.ɛː.ni.dɛː}}). With just four extant [[species]] (each in its own [[genus]]), it is the fifth-smallest family in the order Carnivora and one of the smallest in the class [[Mammalia]].<ref name=MoW1>{{cite book|editor1=Wilson, D.E.|editor2=Mittermeier, R.A.|year=2009|title=Handbook of the Mammals of the World, Volume 1: Carnivora|publisher=Lynx Edicions|location=Barcelona|pages=50–658|isbn=978-84-96553-49-1}}</ref> Despite their low diversity, hyenas are unique and vital components of most [[Africa]]n ecosystems.<ref name="miv">{{Harvnb|Mills|Hofer|1998|p=iv}}</ref>

Although [[phylogenetics|phylogenetically]] closer to [[Felidae|felines]] and [[Viverridae|viverrid]]s, hyenas are [[Ethology|behaviourally]] and [[Morphology (biology)|morphologically]] similar to [[canidae|canids]] in several elements due to [[convergent evolution]]: both hyenas and canines are non-[[arboreal]], [[cursorial]] hunters that catch prey with their teeth rather than claws. Both eat food quickly and may store it, and their calloused feet with large, blunt, nonretractable claws are adapted for running and making sharp turns. However, hyenas' grooming, [[scent marking]], defecation habits, mating, and parental behavior are consistent with the behavior of other feliforms.<ref name="k274">{{Harvnb|Kruuk|1972|p=274}}</ref>

Hyenas feature prominently in the folklore and mythology of human cultures that live alongside them. Hyenas are commonly viewed as frightening and worthy of contempt. In some cultures, hyenas are thought to influence people's spirits, rob graves, and steal livestock and children.<ref name="m96">{{Harvnb|Mills|Hofer|1998|p=96}}</ref> Other cultures associate them with witchcraft, using their body parts in [[traditional medicine]].<ref>{{cite journal|title=A study on use of animals as traditional medicine by Sukuma Tribe of Busega District in North-western Tanzania|first1=Rajeev|last1=Vats|first2=Simion|last2=Thomas|date=7 May 2015|journal=Journal of Ethnobiology and Ethnomedicine|volume=11|article-number=38|doi=10.1186/s13002-015-0001-y|pmid=25947365|pmc=4472419 |doi-access=free }}</ref>

==Evolution==
<!-- Deleted image removed: [[File:Phylogeny and adaptive types of Hyaenidae according to Turner, Antón & Werdelin (2008).jpg|thumb|Phylogeny and adaptive types of Hyaenidae according to Alan Turner, Mauricio Antón & Lars Werdelin.<ref>Coca-Ortega C, Pérez-Claros JA. 2019. Characterizing ecomorphological patterns in hyenids: a multivariate approach using postcanine dentition. PeerJ 6:e6238 https://doi.org/10.7717/peerj.6238</ref>]] -->
===Origins===
Hyenas originated in the jungles of [[Miocene]] Eurasia 22&nbsp;million years ago, when most early feliform species were still largely [[arboreal]]. The first ancestral hyenas were likely similar to the modern [[African civet]]; one of the earliest hyena species described, ''[[Plioviverrops]]'', was a lithe, civet-like animal that inhabited Eurasia 20–22&nbsp;million years ago, and is identifiable as a hyaenid by the structure of the [[middle ear]] and dentition. The lineage of ''Plioviverrops'' prospered and gave rise to descendants with longer legs and more pointed jaws, a direction similar to that taken by canids in [[North America]].<ref name="mac119">{{Harvnb|Macdonald|1992|pp=119–144}}</ref>

Hyenas then diversified into two distinct types: lightly built dog-like hyenas and robust bone-crushing hyenas. Although the dog-like hyenas thrived 15&nbsp;million years ago (with one [[taxon]] having colonised North America), most became extinct after a change in climate, along with the arrival of canids into Eurasia. Of the dog-like hyena lineage, only the insectivorous [[aardwolf]] survived, while the bone-crushing hyenas (including the [[Extant taxon|extant]] spotted, [[brown hyena|brown]], and striped hyenas) became the undisputed top scavengers of Eurasia and Africa.<ref name="mac119" />

===Rise and fall of the dog-like hyenas===
[[File:Ictitherium viverrinum.JPG|thumb|Skull of ''Ictitherium viverrinum'', one of the "dog-like" hyenas. [[American Museum of Natural History]]]]

The descendants of ''Plioviverrops'' reached their peak 15&nbsp;million years ago, with more than 30 species having been identified. Unlike most modern hyena species, which are specialised bone-crushers, these dog-like hyenas were nimble-bodied, wolfish animals; one species among them was ''[[Ictitherium]] viverrinum'', which was similar to a [[jackal]]. The dog-like hyenas were numerous; in some Miocene fossil sites, the remains of ''Ictitherium'' and other dog-like hyenas outnumber those of all other carnivores combined. The decline of the dog-like hyenas began 5–7&nbsp;million years ago during a period of climate change, exacerbated by [[canid]]s crossing the [[Bering land bridge]] to Eurasia. One species, ''[[Chasmaporthetes|Chasmaporthetes ossifragus]]'', managed to cross the land bridge into North America, being the only hyena to do so. ''Chasmaporthetes'' managed to survive for some time in North America by deviating from the endurance-running and bone-crushing niches monopolized by canids and developing into a [[cheetah]]-like sprinter. Most of the dog-like hyenas had died off by 1.5&nbsp;million years ago.<ref name="mac119" />

===Bone-crushing hyenas===
By 10–14&nbsp;million years ago, the hyena family had split into two distinct groups: dog-like hyenas and bone-crushing hyenas. The arrival of the ancestral bone-crushing hyenas coincided with the decline of the similarly built family [[Percrocutidae]]. The bone-crushing hyenas survived the changes in climate and the arrival of canids, which wiped out the dog-like hyenas, though they never crossed into North America, as their niche there had already been taken by the dog subfamily [[Borophaginae]]. By 5&nbsp;million years ago, the bone-crushing hyenas had become the dominant scavengers of Eurasia, primarily feeding on large herbivore carcasses felled by [[sabre-toothed cat]]s. One genus, ''[[Pachycrocuta]]'', was a {{cvt|110|kg}} mega-scavenger that could splinter the bones of [[elephant]]s.<ref name=Palmqvist_et_al_2011>{{cite journal|last1=Palmqvist|first1=P.|last2=Martinez-Navarro|first2=B.|last3=Pérez-Claros|first3=J. A.|last4=Torregrosa|first4=V.|last5=Figueiridio|first5=B.|last6=Jiménez-Arenas|first6=J. M.|last7=Patrocinio Espigares|first7=M.|last8=Ros-Montoya|first8=Sergio|last9=De Renzi|first9=M.|title=The giant hyena Pachycrocuta brevirostris: Modelling the bone-cracking behavior of an extinct carnivore|journal=Quaternary International|date=2011|volume=243|issue=#1|page=61|doi=10.1016/j.quaint.2010.12.035|bibcode=2011QuInt.243...61P |hdl=10630/33571|hdl-access=free}}</ref><ref name="mac119" /> Starting in the early [[Chibanian|Middle Pleistocene]], ''Pachycrocuta'' was replaced by the smaller ''Crocuta'' and ''Hyena'', which corresponds to a general faunal change, perhaps in connection to the [[Mid-Pleistocene Transition|Mid-Pleistocene transition]].<ref>{{Cite journal |last1=Iannucci |first1=Alessio |last2=Mecozzi |first2=Beniamino |last3=Sardella |first3=Raffaele |last4=Iurino |first4=Dawid Adam |date=2021-11-15 |title=The extinction of the giant hyena Pachycrocuta brevirostris and a reappraisal of the Epivillafranchian and Galerian Hyaenidae in Europe: Faunal turnover during the Early–Middle Pleistocene Transition |url=https://www.sciencedirect.com/science/article/pii/S0277379121004479 |journal=Quaternary Science Reviews |volume=272 |article-number=107240 |doi=10.1016/j.quascirev.2021.107240 |bibcode=2021QSRv..27207240I |issn=0277-3791|url-access=subscription }}</ref>

===Rise of modern hyenas===
[[File:Stripedspottedhyenas.jpg|thumb|right|upright|Skeletons of a [[striped hyena]] (left) and a [[spotted hyena]] (right), two species of the "bone-crushing" hyenas]]
The four extant species are the striped hyena (''Hyaena hyaena''), the brown hyena (''Parahyaena brunnea''), the spotted hyena (''Crocuta crocuta''), and the aardwolf (''Proteles cristata'').

The [[aardwolf]] can trace its lineage directly back to ''Plioviverrops'' 15&nbsp;million years ago, and is the only survivor of the dog-like hyena lineage. Its success is partly attributed to its insectivorous diet, for which it faced no competition from canids crossing from North America. It is likely that its unrivaled ability to digest the [[terpene]] excretions from soldier [[termite]]s is a modification of the strong digestive system its ancestors used to consume fetid carrion.<ref name="mac119" />

The [[striped hyena]] may have evolved from ''Hyaenictitherium namaquensis'' of [[Pliocene]] [[Africa]]. Striped hyena fossils are common in Africa, with records going back as far as the [[Villafranchian]]. As fossil striped hyenas are absent from the [[Mediterranean]] region, it is likely that the species is a relatively late invader to Eurasia, having likely spread outside Africa only after the extinction of [[spotted hyena]]s in Asia at the end of the [[Last Glacial Period|Ice Age]]. The striped hyena occurred for some time in [[Europe]] during the Pleistocene, having been particularly widespread in [[France]] and [[Germany]]. It also occurred in [[Montmaurin]], [[Hollabrunn]] in [[Austria]], the [[Furninha|Furninha Cave]] in [[Portugal]] and the Genista Caves in [[Gibraltar]]. The European form was similar in appearance to modern populations, but was larger, being comparable in size to the [[brown hyena]].<ref name="k66">{{Harvnb|Kurtén|1968|pp=66–68}}</ref>

The [[spotted hyena]] (''Crocuta crocuta'') diverged from the striped and brown hyena 10&nbsp;million years ago.<ref name="m1">{{Harvnb|Mills|Hofer|1998|p=1}}</ref> Its direct ancestor was the Indian ''Crocuta sivalensis'', which lived during the Villafranchian.<ref name="k69" /> Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp [[carnassial]]s behind their crushing premolars, therefore they did not need to wait for their prey to die, and thus became pack hunters as well as scavengers. They began forming increasingly larger [[territory (animal)|territories]], necessitated by the fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another [[Hyena clan|clan's]] turf.<ref name="mac119" /> Spotted hyenas spread from their original homeland during the [[Middle Pleistocene]], and quickly colonised a very wide area from Europe, to southern Africa and [[China]].<ref name="k69">{{Harvnb|Kurtén|1968|pp=69–72}}</ref> 

The eventual disappearance of the spotted hyena from Europe has traditionally been attributed to the end of the [[Last Glacial Period|last glacial period]] and a subsequent displacement of open grassland by closed forests, which favoured wolves and humans instead.<ref name="hyenawolf">{{cite web| url=http://www.u.arizona.edu/~mstiner/pdf/Stiner2004a.pdf| title=Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy| work=C. Stiner, Mary| publisher=Revue de Paléobiologie, Genève| access-date=2008-09-16| archive-date=2019-05-08| archive-url=https://web.archive.org/web/20190508135925/http://www.u.arizona.edu/~mstiner/pdf/Stiner2004a.pdf}}</ref> However, analyses have shown that [[climate change]] alone is insufficient to explain the spotted hyena's disappearance from Europe, suggesting that other factors – such as human pressure – must have played a role.<ref>{{Cite journal |last1=Varela |first1=Sara |last2=Lobo |first2=Jorge M. |last3=Rodríguez |first3=Jesús |last4=Batra |first4=Persaram |date=2010-08-01 |title=Were the Late Pleistocene climatic changes responsible for the disappearance of the European spotted hyena populations? Hindcasting a species geographic distribution across time |url=https://www.sciencedirect.com/science/article/pii/S0277379110001265 |journal=Quaternary Science Reviews |volume=29 |issue=17 |pages=2027–2035 |doi=10.1016/j.quascirev.2010.04.017 |bibcode=2010QSRv...29.2027V |issn=0277-3791|url-access=subscription }}</ref> This suggests that the events must be seen within the broader context of [[Late Pleistocene extinctions|late-Quaternary extinctions]], as the late Pleistocene and early Holocene saw the disappearance of many primarily large mammals from Europe and the world.

Expansion or [[Gene duplication|duplication]] of the [[Olfactory receptor|olfatory receptor]] gene family has been found in all 4 extant species, which would have led to the evolution of the more specialised feeding habits of hyenas.<ref name="Westbury-2021">{{Cite journal |last1=Westbury |first1=Michael V |last2=Le Duc |first2=Diana |last3=Duchêne |first3=David A |last4=Krishnan |first4=Arunkumar |last5=Prost |first5=Stefan |last6=Rutschmann |first6=Sereina |last7=Grau |first7=Jose H |last8=Dalén |first8=Love |last9=Weyrich |first9=Alexandra |last10=Norén |first10=Karin |last11=Werdelin |first11=Lars |last12=Dalerum |first12=Fredrik |last13=Schöneberg |first13=Torsten |last14=Hofreiter |first14=Michael |date=2021 |title=Ecological Specialization and Evolutionary Reticulation in Extant Hyaenidae |url=https://academic.oup.com/mbe/article/38/9/3884/6149117 |journal=Molecular Biology and Evolution |volume=38 |issue=9 |pages=3884–3897 |doi=10.1093/molbev/msab055 |issn=1537-1719 |pmc=8382907 |pmid=34426844}}</ref>

Expansion in immune-related gene families was also found in the spotted hyena, striped hyena and brown hyena, which would have led to the evolution of the [[scavenging]] in these species. Mutations and variants were also found in digestion-related genes (''ASH1L'', ''PTPN5'', ''PKP3'', ''AQP10''). One of these digestion-related genes has variants also related to enhanced bone mineralisation (''PTPN5''), while other have also a role in inflammatory skin responses (''PKP3'').<ref name="Westbury-2021" />

In aardwolves, expansion of genes related to toxin response were found ([[Lipocalin]] and [[Glucuronosyltransferase|UDP Glucuronosyltransferase]] gene families), which would have led to the evolution of the feeding of termites [[Trinervitermes trinervoides|Trinervitermes]] in this species. Mutations and variants in genes related to craniofacial shape were also found (''GARS'', ''GMPR'', ''STIP1'', ''SMO'' and ''PAPSS2''). Another gene is related to protective epidermis function (''DSC1'').<ref name="Westbury-2021" />

{{See also|Cave hyena}}

==Genera of the Hyaenidae (extinct and recent)==
[[File:Crocuta crocuta.jpg|thumb|A [[spotted hyena]] of subfamily Hyaeninae]]
The list follows McKenna and Bell's ''Classification of Mammals'' for prehistoric genera (1997)<ref name="McKenna & Bell">Malcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'', Columbia University Press, New York 1997, 631 Seiten, {{ISBN|0-231-11013-8}}</ref> and Wozencraft (2005) in Wilson and Reeders ''[[Mammal Species of the World]]'' for extant genera.<ref name="MSW3">{{MSW3 Wozencraft|pages=532–548}}</ref> The percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family [[Percrocutidae]], although they are generally grouped as sister-taxa to hyenas.<ref>{{cite journal| year=2013| title=Skull Shape Evolution In Durophagous Carnivorans| journal=Evolution| volume=67| issue=7| pages=1975–93| doi=10.1111/evo.12059 | last1=Figueirido| first1=Borja| last2=Jack Tseng| first2=Zhijie| last3=Martín-Serra| first3=Alberto| pmid=23815654| bibcode=2013Evolu..67.1975F| s2cid=23918004 | doi-access=free| hdl=10630/32909| hdl-access=free}}</ref> However, {{as of|2024|lc=y}} many experts consider them members of Hyaenidae.<ref>{{Cite journal |last=Pérez-Claros |first=Juan A. |date=26 April 2023 |title=An ecomorphological characterization of the percrocutoid hyaenids: a multivariate approach using postcanine dentition |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2023.2197972 |journal=Journal of Vertebrate Paleontology |volume=42 |issue=5 |article-number=e2197972 |doi=10.1080/02724634.2023.2197972 }}</ref><ref name="Xiong 2022">{{Cite journal |last=Xiong |first=Wuyang |date=1 May 2022 |title=New species of Percrocuta (Carnivora, Hyaenidae) from the early middle Miocene of Tongxin, China |journal=Historical Biology |volume=35 |issue=5 |pages=799–820 |doi=10.1080/08912963.2022.2067757 |issn=0891-2963 |s2cid=248627038 }}</ref><ref>{{Cite journal |last=Wuyang |first=Xiong |date=October 2019 |title=Basicranial morphology of Late Miocene Dinocrocuta gigantea (Carnivora: Hyaenidae) from Fugu, Shaanxi |url=https://www.researchgate.net/publication/336578352 |journal=Vertebrata PalAsiatica |volume=57 |issue=4 |pages=274–307}}</ref><ref>{{Cite journal |last1=Barret |first1=Paul Z. |last2=Hopkins |first2=Samatha S. B. |date=2024 |title=Mosaic evolution underlies feliform morphological disparity |journal=Proc. R. Soc. B |volume=291 |issue=2028 |article-number=20240756 |doi=10.1098/rspb.2024.0756 |pmid=39137889 |pmc=11321862 }}</ref> Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus ''Pachycrocuta'', but in the genus ''Parahyaena''. However, some research has suggested ''Parahyaena'' may be synonymous with ''Pachycrocuta'', making the brown hyena the only extant member of this genus.<ref>{{cite journal|first1=J. A|last1=Pérez-Claros|year=2024|title= Unravelling the origin of the brown hyena (Parahyena brunnea) and its evolutionary and paleoecological implications for the Pachycrocuta lineage |journal=Palaeontologia Electronica|doi=10.26879/1372 |doi-access=free}}</ref>

* '''Family Hyaenidae'''
** '''Subfamily ''Incertae sedis'''''
*** †''[[Tongxinictis]]''<ref name=CocaOrtega2019>{{cite journal|last1=Coca-Ortega|first1=C.|last2=Pérez-Claros|first2=J.A.|date=January 2019|title=Characterizing ecomorphological patterns in hyenids: a multivariate approach using postcanine dentition|pages=6:e6238|journal=PeerJ|volume=6|article-number=e6238 |doi=10.7717/peerj.6238|pmid=30648005|pmc=6330948|doi-access=free }}</ref> (Middle Miocene of Asia)
** †'''Subfamily Ictitheriinae'''
*** †''[[Herpestides]]'' (Early Miocene of Africa and Eurasia)
*** †''[[Plioviverrops]]'' (including ''Jordanictis'', ''Protoviverrops'', ''Mesoviverrops''; Early Miocene to Early Pliocene of Europe, Late Miocene of Asia)
*** †''[[Ictitherium]]'' (=''Galeotherium''; including ''Lepthyaena'', ''Sinictitherium'', ''Paraictitherium''; Middle Miocene of Africa, Late Miocene to Early Pliocene of Eurasia)
*** †''[[Thalassictis]]'' (including ''Palhyaena'', ''Miohyaena'', ''Hyaenictitherium'', ''Hyaenalopex''; Middle to Late Miocene of Asia, Late Miocene of Africa and Europe)
*** †''[[Hyaenotherium]]'' (Late Miocene to Early Pliocene of Eurasia)
*** †''[[Miohyaenotherium]]''<ref name=Werdelin1996>{{cite journal|last1=Werdelin|first1=Lars|date=March 1996|title=Community-wide character displacement in Miocene hyaenas|journal=Lethaia|volume=29|issue=1|pages=97–106|doi=10.1111/j.1502-3931.1996.tb01843.x|bibcode=1996Letha..29...97W }}</ref> (Late Miocene of Europe)
*** †''[[Lycyaena]]'' (Late Miocene of Eurasia)
*** †''[[Tungurictis]]''<ref name=Wang2020>{{cite journal|last1=Wang|first1=X.|last2=Tsang|first2=Z.J.|display-authors=etal|date=February 2020|title=A new species of Tungurictis Colbert, 1939 (Carnivora, Hyaenidae) from the middle Miocene of Junggar Basin, northwestern China and the early divergence of basal hyaenids in East Asia|journal=Geodiversitas|volume=42|issue=3|pages=29–45|doi=10.5252/geodiversitas2020v42a3|bibcode=2020Geodv..42...29W |doi-access=free }}</ref> (Middle Miocene of Africa and Eurasia)
*** †''[[Protictitherium]]'' (Middle Miocene of Africa and Asia, Middle to Late Miocene of Europe)
** '''Subfamily Hyaeninae'''
*** †''[[Palinhyaena]]''<ref name=Turner2008>{{cite journal|last1=Turner|first1=A.|last2=Antón|first2=M.|last3=Werdelin|first3=L.|date=September 2008|title=Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe|journal=Geobios|volume=41|issue=5|pages=677–687|doi=10.1016/j.geobios.2008.01.001|bibcode=2008Geobi..41..677T }}</ref> (Late Miocene of Asia)
*** †''[[Ikelohyaena]]''<ref name=Tseng2011>{{cite journal|last1=Tseng|first1=Z.J.|last2=Stynder|first2=D.|date=March 2011|title=Mosaic functionality in a transitional ecomorphology: skull biomechanics in stem Hyaeninae compared to modern South African carnivorans|journal=Biological Journal of the Linnean Society|volume=102|issue=3|pages=540–559|doi=10.1111/j.1095-8312.2010.01602.x|doi-access=free }}</ref> (Early Pliocene of Africa)
*** ''[[Hyaena]]'' (=''Euhyaena'',=''Parahyaena''; including [[striped hyena]], ''Pliohyaena'', ''Pliocrocuta'', ''Anomalopithecus'') Early Pliocene (?Middle Miocene) to Recent of Africa, Late Pliocene (?Late Miocene) to Late Pleistocene of Europe, Late Pliocene to recent in Asia
*** ''[[Parahyaena]]'' (=''Hyaena''; [[brown hyena]] Pliocene to recent of Africa)
*** †''[[Hyaenictis]]''<ref name=Vinuesa2016>{{cite journal|last1=Vinuesa|first1=V.|last2=Madurell-Malapeira|first2=J.|display-authors=etal|date=April 2016|title=A new skull of ''Hyaenictis'' Gaudry, 1861 (Carnivora, Hyaenidae) shows incipient adaptations to durophagy|journal=Journal of Mammalian Evolution|volume=24|issue=2|pages=207–219|doi=10.1007/s10914-016-9334-0|s2cid=23453043|url=http://nrm.diva-portal.org/smash/get/diva2:926393/FULLTEXT02 }}</ref> (Late Miocene of Asia?, Late Miocene of Europe, Early Pliocene (?Early Pleistocene) of Africa)
*** †''[[Leecyaena]]''<ref name=Turner2008/> (Late Miocene and/or Early Pliocene of Asia)
*** †''[[Chasmaporthetes]]'' (=''Ailuriaena''; including ''Lycaenops'', ''Euryboas''; Late Miocene to Early Pleistocene of Eurasia, Early Pliocene to Late Pliocene or Early Pleistocene of Africa, Late Pliocene to Early Pleistocene of North America)
*** †''[[Pachycrocuta]]'' (Pliocene and Pleistocene of Eurasia and Africa)
*** †''[[Adcrocuta]]'' (Late Miocene of Eurasia)
*** ''[[Spotted hyena|Crocuta]]'' (=''Crocotta''; including ''Eucrocuta''; [[spotted hyena]] and [[cave hyena]]. Late Pliocene to recent of Africa, Late Pliocene to Late Pleistocene of Eurasia)
** '''Subfamily Protelinae'''
*** †''[[Gansuyaena]]''<ref>{{cite journal |last1=Tseng |first1=Z. Jack |date=2022 |title=A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China |url=https://www.researchgate.net/publication/361162806 |journal=Vertebrata PalAsiatica |volume=60 |issue=2 |pages=81–116 |doi=10.19615/j.cnki.2096-9899.211025 }}</ref>
*** ''[[Proteles]]'' (=''Geocyon''; [[aardwolf]]. Pleistocene to Recent of Africa)

===Phylogeny===
The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991),<ref name="Werdelin1991">{{cite journal|last1=Werdelin|first1=L.|last2=Solounias|first2=Nikos|title=The Hyaenidae: taxonomy, systematics and evolution.|url=http://foreninger.uio.no/ngf/FOS/pdfs/F&S_30.pdf|journal=Fossils and Strata|volume=30|pages=1–104|year=1991|doi=10.18261/8200374815-1991-01 |isbn=82-00-37481-5 }}</ref> as updated by Turner et al. (2008).<ref name=Turner2008/>
{{Clade
 |style=font-size:80%; line-height:85%;background-color:#eeeeff
 |caption=Evolution of hyaenas
 |captionstyle=background-color:#8899ff;padding:10px;font-weight:bold;
 |footer=Phylogenic relationships based on morphological characteristics, after Werdelin & Solounias (1991) and Turner et al (2008)
 |footerstyle=background-color:#eeeeee;text-align:center;
 |1={{clade
  |style=background-color:#ccffcc;
  |label1=
  |1=''[[Protictitherium|Protictitherium crassum]]''
  |2=''"Protictitherium" cingulatum''
  |3=''"Protictitherium" intermedium''
  |4=''"Protictitherium" llopisi''
  |5=''"Protictitherium" punicum''
  |6=''"[[Protictitherium|Protictitherium" gaillardi]]''
  |state7=dotted
  |7=''"Protictitherium" sumegense''
  |state8=dotted
  |8=''"Protictitherium" csakvarense''
  |style9=background-color:#eeeeff;
  |9={{clade
    |label1=
    |1={{clade
        |label1=''[[Plioviverrops]]''
        |style1=background-color:#ccffcc;
        |1={{clade
           |grouplabel1=Civet/mongoose-like insectivore-omnivores 
           |grouplabelstyle1=font-size:12px;font-weight:bold;color:darkgreen;text-align:right;
           |label1=
           |1=''Plioviverrops gervaisi''
           |2={{clade
              |1=''Plioviverrops orbignyi''
              |2={{Clade
                 |label1=
                 |1=''Plioviverrops guerini''
                 |2=''Plioviverrops faventinus''
                 |3=''Plioviverrops gaudryi''
                  }} 
               }} 
            }}
         }}
     |2={{clade
        |label1=
        |1=''Tungurictis spocki''
        |2={{clade
           |label1=
           |1=''[[Thalassictis|Thalassictis robusta]]''
           |2=''"Thalassictis" certa''
           |3=''"Thalassictis" montadai''
           |4=''"Thalassictis" proava''
           |5=''"Thalassictis" sarmatica''
           |6=''"Thalassictis" spelaea''
           |7={{clade
              |label1=
              |1=''Tongxinictis primordialis''
              |2={{clade
                 |label1=
                 |1={{clade
                    |label1=''[[Proteles]]''
                    |style1=background-color:#ddeeff;
                    |1={{clade
                       |label1=
                       |1='''''[[Proteles cristatus]]''''' (aardwolf) [[File:The life of animals (Colored Plate 4) (proteles cristatus).jpg|50 px]]
                       |2=''Proteles amplidentus''
                        }}
                     }}
                 |2={{clade
                    |label1=
                    |1={{clade
                       |label1=''[[Ictitherium]]''
                       |style=background-color:#ddddff;
                       |1={{clade
                          |label1=
                          |1=''[[Ictitherium|Ictitherium viverrinum]]''
                          |2=''Ictitherium ebu''
                          |grouplabel2=Jackal-like hyaenas 
                          |grouplabelstyle2=font-size:12px;font-weight:bold;color:darkcyan;text-align:right;
                          |3={{clade
                             |label1=
                             |1={{clade
                                |label1=
                                |1=''Ictitherium  tauricum''
                                |2=''Ictitherium ibericum''
                                 }}
                             |2={{clade
                                |label1=
                                |1=''Ictitherium kurteni''
                                |2={{clade
                                   |1=''Ictitherium intuberculatum''
                                   |2=''Ictitherium pannonicum''
                                    }}
                                 }} 
                              }} 
                           }} 
                        }}
                    |2={{clade 
                       |label1=
                       |1=''Miohyaenotherium bessarabicum''
                       |2={{clade
                          |1={{clade
                             |label1=''[[Hyaenotherium]]''
                             |1={{clade
                                |1=''Hyaenotherium wongii''
                                |2=''Hyaenictitherium hyaenoides''
                                |3=''"Hyaenictitherium" pilgrimi''
                                |4=''"Hyaenictitherium" parvum''
                                |5=''"Hyaenictitherium" namaquensis''
                                |6=''"Hyaenictitherium" minimum''
                                 }}
                              }}
                          |style2=background-color:#ccccff;
                          |2={{clade
                             |1={{clade
                                |label1=''[[Lycyaena]]''
                                |1={{clade
                                   |grouplabel2=Cursorial hunting hyaenas 
                                   |grouplabelstyle2=font-size:12px;font-weight:bold;color:darkblue;text-align:right;
                                   |1=''Lycyaena chaeretis''
                                   |2=''Lycyaena dubia''
                                   |3={{clade
                                      |1=''Lycyaena macrostoma''
                                      |2=''Lycyaena crusafonti''
                                       }}
                                    }} 
                                 }}
                             |2={{clade
                                |1={{clade
                                   |label1=''[[Hyaenictis]]''
                                   |1={{clade
                                      |1=''Hyaenictis graeca''
                                      |2=''Hyaenictis almerai''
                                      |3=''Hyaenictis hendeyi''
                                       }}
                                    }}
                                 |2={{clade
                                    |1={{clade
                                       |label1=''[[Lycyaenops]]''
                                       |1={{clade
                                          |1=''Lycyaenops rhomboideae''
                                          |2=''Lycyaenops silberbergi''
                                           }}
                                        }}
                                    |2={{clade
                                       |label1=''[[Chasmaporthetes]]''
                                       |sublabel1=(running hyaenas)
                                       |1={{clade
                                          |1={{clade
                                             |1=''Chasmaporthetes exitelus''
                                             |2=''Chasmaporthetes bonisi''
                                              }}
                                          |2={{clade
                                             |1=''Chasmaporthetes borissiaki''
                                             |2={{clade
                                                |1={{clade
                                                   |1=''Chasmaporthetes lunensis''
                                                   |2=''Chasmaporthetes melei''
                                                    }}
                                                |2={{clade
                                                   |1={{clade
                                                      |1=''Chasmaporthetes ossifragus''
                                                      |2=''Chasmaporthetes'' sp. Florida
                                                       }}
                                                   |2={{clade
                                                      |1=''Chasmaporthetes nitidula''
                                                      |2=''Chasmaporthetes australis''
                                                       }} }} }} }} }}
                                       |style2=background-color:#ddccff
                                       |label2=[[Hyaeninae]]
                                       |sublabel2=(bone-cracking hyenas)
                                       |2={HYAENINAE}
                                         }} }} }} }} }} }} }} }} }} }} }} }} }} 
|targetA={HYAENINAE}
|subcladeA={{clade
  |caption=Bone-cracking hyaenas
  |captionstyle=font-size:12px;font-weight:bold;color:darkred;text-align:right;padding:5px 20px 0px 0px;
  |1=''Metahyaena confector''
  |2=''Palinhyaena reperta''
  |3={{clade
     |1=Hyaenid sp. E Langebaar
     |2={{clade
        |1=''Belbus beaumonti''
    <!--|1=''Leecyaena lycyaenoides''
        |2=''"Leecyaena" bosei'' -->
        |style2=background-color:#eeccff;
        |2={{clade
           |1={{clade
              |1={{clade
                 |1=''Hyaena abronia''
                 |2='''''[[Hyaena hyaena]]''''' (striped hyaena) [[File:Hyaena striata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|50 px]]
                  }}
              |2={{clade
                 |1=''Parahyaena howelli''
                 |2='''''[[Parahyaena brunnea]]''''' (brown hyaena) [[File:Hyaena fusca (white background).jpg|50 px]]
                  }} }}
           |2={{clade
              |1=''Pliocrocuta perrieri''
              |2={{clade
                 |1=''[[Pachycrocuta|Pachycrocuta brevirostris]]'' (giant hyaena) [[File:Pachycrocuta brevirostris restoration.jpg|70 px]]
             <!--|2=''"Pachycrocuta"'' bellax''-->
                 |2={{clade
                    |1=''[[Adcrocuta|Adcrocuta eximia]]'' [[File:Adcrocuta eximia restoration.jpg|60 px]]
                    |2=''Allohyaena kadici''
                    |3={{clade
                       |1='''''[[Crocuta crocuta]]''''' (spotted hyaena) [[File:Spotted hyena (Crocuta crocuta) by Connor Ashbridge.png|50 px]]
                       |2=''Crocuta eturono''
                   <!--|2=''Crocuta sivalensis''
                       |3=''Crocuta dietrichi'' -->
}} }} }} }} }} }} }} }} }}

A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli ''et al'', 2006<ref name="Koepfli2006">{{cite journal|last1=Koepfli|first1=K.-P.|last2=Jenks|first2=S. M.|last3=Eizirik|first3=E.|last4=Zahirpour|first4=T.|last5=Van Valkenburgh|first5=B.|last6=Wayne|first6=R. K.|title=Molecular systematics of the Hyaenidae: Relationships of a relictual lineage resolved by a molecular supermatrix|journal=Molecular Phylogenetics and Evolution|volume=38|issue=3|date=2006|pages=603–620|doi=10.1016/j.ympev.2005.10.017|pmid=16503281|bibcode=2006MolPE..38..603K }}</ref>).

{{clade|style=font-size:90%; line-height:100%;
|label1='''Hyaenidae'''
|1={{clade
  |label1=
  |1={{clade
     |label1=
     |1=''[[Aardwolf|Proteles cristatus]]'' (aardwolf) [[File:The life of animals (Colored Plate 4) (proteles cristatus).jpg|60 px]]
     }}
  |2={{clade
     |label1=
     |1={{clade |label1=|1=''[[Spotted hyena|Crocuta crocuta]]'' (spotted hyena) [[File:Spotted hyena (Crocuta crocuta) by Connor Ashbridge.png|80 px]] }}
     |2={{clade
        |1=''[[Striped hyena|Hyaena hyaena]]'' (striped hyena) [[File:Hyaena striata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|70 px]]
        |2=''[[Brown hyena|Parahyaena brunnea]]'' (brown hyena) [[File:Hyaena fusca (white background).jpg|65 px]] 
         }} }} }} 
         
}}

==Characteristics==

===Build===
{{further|Spotted hyena#Description|Striped hyena#Description|Aardwolf#Description|Brown hyena#Description}}
[[File:StripedHyenaSkull.jpg|thumb|right|220px|Striped hyena skull. Note the disproportionately large carnassials and premolars adapted for bone consumption.]]
[[File:Aardwolfskull.jpg|thumb|right|220px|Aardwolf skull. Note the greatly reduced molars and carnassials, rendered redundant from insectivory.]]
Hyenas have relatively short torsos and are fairly massive and [[wolf]]-like in build, but have lower hind quarters, high withers and their backs slope noticeably downward towards their rumps. The forelegs are high, while the hind legs are very short and their necks are thick and short. Their skulls superficially resemble those of large canids, but are much larger and heavier, with shorter facial portions. Hyenas are [[digitigrade]], with the fore and hind paws having four digits each and sporting bulging pawpads.<ref name="h3" /> Like canids, hyenas have short, blunt, non-retractable claws.<ref name="p62" /> Their [[Fur|pelage]] is sparse and coarse with poorly developed or absent underfur. Most species have a rich mane of long hair running from the withers or from the head.<ref name="h3">{{Harvnb|Heptner|Sludskii|1992|p=3}}</ref> With the exception of the spotted hyena, hyaenids have striped coats, which they likely inherited from their [[viverrid]] ancestors.<ref name="mac119" /> Their ears are large and have simple basal ridges and no marginal bursa.<ref name="p62" /> Their [[vertebral columns]], including the [[Cervical vertebrae|cervical region]], are of limited mobility. Hyenas have no [[baculum]].<ref name="h4" /> Hyenas have one more pair of ribs than canids do, and their tongues are rough like those of felids and viverrids.<ref>Holl, William & Wood, Neville ''The Analyst: a quarterly journal of science, literature, natural history, and the fine arts, Volume 10'', p. 59, Simpkin & Marshall, 1840</ref> Males in most hyena species are larger than females,<ref name="m21">{{Harvnb|Mills|Hofer|1998|p=21}}</ref> though the spotted hyena is an exception, as it is the female of the species that outweighs and dominates the male. Also, unlike other hyenas, the [[Female genitalia of spotted hyenas|female spotted hyena's external genitalia]] closely resembles that of the male.<ref name="k210">{{Harvnb|Kruuk|1972|pp=210–211}}</ref>

Their [[dentition]] is similar to that of the [[canid]], but is more specialised for consuming coarse food and crushing bones. The [[carnassial]]s, especially the upper, are very powerful and are shifted far back to the point of exertion of peak pressure on the jaws. The other teeth, save for the underdeveloped upper [[molar (tooth)|molars]], are powerful, with broad bases and cutting edges. The [[canine teeth|canines]] are short, but thick and robust.<ref name="h4">{{Harvnb|Heptner|Sludskii|1992|pp=4–5}}</ref> [[Anatomical terms of location#Teeth|Labiolingually]], their [[mandible]]s are much stronger at the canine teeth than in canids, reflecting the fact that hyenas crack bones with both their anterior dentition and [[premolars]], unlike canids, which do so with their post-carnassial molars.<ref>{{cite journal|title=Mandibular force profiles of extant carnivorans and implications for the feeding behavior of extinct predators|journal=Journal of Zoology|year=2005|first=François|last=Therrien|volume=267|issue=3|pages=249–270|doi=10.1017/S0952836905007430 |bibcode=2005JZoo..267..249T }}</ref> The strength of their jaws is such that both striped and spotted hyenas have been recorded to kill dogs with a single bite to the neck without breaking the skin.<ref name="johnson">[[Daniel Johnson (surgeon)|Daniel Johnson]] (1827) ''[https://archive.org/stream/sketchesofindian00johnrich#page/46/mode/2up/search/hyena Sketches of Indian Field Sports: With Observations on the Animals; Also an Account of Some of the Customs of the Inhabitants; with a Description of the Art of Catching Serpents, as Practiced by the Conjoors and Their Method of Curing Themselves when Bitten: with Remarks on Hydrophobia and Rabid Animals]'' p. 45-46, R. Jennings, 1827</ref><ref name="hamilton">Stevenson-Hamilton, James (1917) ''[https://archive.org/details/animallifeinafri01steviala Animal life in Africa], Vol. 1'', p.95, London : William Heinemann</ref> The spotted hyena is renowned for its strong bite proportional to its size, but a number of other animals (including the [[Tasmanian devil]]) are proportionately stronger.<ref>{{cite web|last=Salleh|first=Anna|title=Marsupial has the deadliest bite|url=http://www.abc.net.au/science/articles/2005/04/04/1336220.htm|publisher=abc.net.au|access-date=24 January 2013|date=4 April 2005}}</ref><ref name="Wroe2005">{{cite journal|last1=Wroe|first1=S.|last2=McHenry|first2=C.|last3=Thomason|first3=J.|year=2005|title=Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa|journal=Proceedings of the Royal Society B: Biological Sciences|pmid=15817436|volume=272|issue=1563|pmc=1564077|pages=619–625|doi=10.1098/rspb.2004.2986}}</ref> The aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dental formula as the other three species.<ref name="EoM">{{cite book|editor1-last=Macdonald|editor1-first=D.|last1=Richardson|first1=Philip K.R.|last2=Bearder|first2=Simon|year=1984|title=The Encyclopedia of Mammals|publisher=Facts on File|location=New York|pages=[https://archive.org/details/encyclopediaofma00mals_0/page/154 154–159]|isbn=0-87196-871-1|url=https://archive.org/details/encyclopediaofma00mals_0/page/154 }}</ref> The [[dentition|dental formula]] for all hyena species is: {{DentalFormula|upper=3.1.4.1|lower=3.1.3.1}}

Although hyenas lack [[perineum|perineal]] scent glands, they have a large pouch of naked skin located at the anal opening. Large [[anal gland]]s above the anus open into this pouch. Several [[sebaceous gland]]s are present between the openings of the anal glands and above them.<ref name="p62">{{Harvnb|Pocock|1941|pp=62–63}}</ref> These glands produce a white, creamy secretion that the hyenas paste onto grass stalks. The odor of [[hyena butter|this secretion]] is very strong, smelling of boiling cheap [[soap]] or burning, and can be detected by humans several meters downwind.<ref name="k222">{{Harvnb|Kruuk|1972|p=222}}</ref> The secretions are primarily used for [[territorial marking]], though both the aardwolf<ref name="mac119" /> and the striped hyena<ref name="h38">{{Harvnb|Heptner|Sludskii|1992|p=38}}</ref> will [[Spraying (animal behavior)|spray]] them when attacked.

===Behavior===
[[File:Brown hyena in Ghanzi, BW-GH, BW 2.jpg|thumb|Brown hyena marking its territory with its anal glands]]
[[File:2667 Spotted Hyena Cubs.JPG|thumb|200px|right|Spotted hyena cubs at their den]]
Hyenas [[personal grooming|groom themselves]] often like [[Felidae|felids]] and [[Viverridae|viverrids]], and their way of licking their genitals is very cat-like (sitting on the lower back, legs spread with one leg pointing vertically upward). They defecate in the same manner as other Carnivora, though they never [[Raised-leg urination|raise their legs as canids do when urinating]], as urination serves no territorial function for them. Instead, hyenas [[territorial marking|mark their territories]] using their anal glands, a trait found also in viverrids and [[Mustelidae|mustelid]]s, but not canids and felids.<ref name="k271">{{Harvnb|Kruuk|1972|pp=271–73}}</ref> When attacked by lions or dogs, striped<ref name="p72"/> and brown hyenas<ref name="mills60">{{Harvnb|Mills|Mills|2010|pp=60–61}}</ref> will [[Apparent death|feign death]], though the spotted hyena will defend itself ferociously.<ref name="hamilton" /> The spotted hyena is very vocal, producing a number of different sounds consisting of whoops, grunts, groans, lows, giggles, yells, growls, laughs and whines.<ref name="k220">{{Harvnb|Kruuk|1972|p=220}}</ref> The striped hyena is comparatively silent, its vocalizations being limited to a chattering laugh and howling.<ref name="p73">{{Harvnb|Pocock|1941|p=73}}</ref>
[[file:Spotted Hyaena (Crocuta crocuta) (W1CDR0000381 BD12).ogg|left|thumb|Whoop of a spotted hyena in Umfolosi Game Park, South Africa.]]
{{Anchor|Mating}}
[[Animal sexual behavior|Mating]] between hyenas involves a number of short copulations with brief intervals, unlike canids, who generally [[canine tying|engage in a single, drawn out copulation]].<ref name="k271" /> Spotted hyena cubs are born almost fully developed, with their eyes open and erupting incisors and canines, though lacking adult markings.<ref name="k247">{{Harvnb|Kruuk|1972|pp=247–249}}</ref> In contrast, striped hyena cubs are born with adult markings, closed eyes and small ears.<ref name="r350">{{Harvnb|Rosevear|1974|p=350}}</ref> Hyenas do not [[regurgitation (digestion)|regurgitate]] food for their young and male spotted hyenas play no part in raising their cubs,<ref name="k271" /> though male striped hyenas do so.<ref name="h40">{{Harvnb|Heptner|Sludskii|1992|pp=40–42}}</ref>

The striped hyena is primarily a scavenger, though it will also attack and kill any animals it can overcome,<ref name="p72">{{Harvnb|Pocock|1941|p=72}}</ref> and will supplement its diet with fruit.<ref name="h31">{{Harvnb|Heptner|Sludskii|1992|pp=31–33}}</ref> The spotted hyena, though it also scavenges occasionally, is an active pack hunter of medium to large sized [[ungulates]], which it catches by wearing them down in long chases and dismembering them in a canid-like manner. Spotted hyenas may kill as many as 95% of the animals they eat.<ref>[http://www.hyaenidae.org/the-hyaenidae/spotted-hyena-crocuta-crocuta/crocuta-diet-and-foraging.html Hyaena Specialist Group - Spotted Hyena: Diet and Foraging] {{webarchive|url=https://web.archive.org/web/20110204092003/http://www.hyaenidae.org/the-hyaenidae/spotted-hyena-crocuta-crocuta/crocuta-diet-and-foraging.html|date=2011-02-04 }}. Hyaenidae.org. Retrieved on 2015-11-06.</ref>

The aardwolf is primarily an insectivore, specialised for feeding on [[termite]]s of the genus ''[[Trinervitermes]]'' and ''[[Hodotermes]]'', which it consumes by licking them up with its long, broad tongue. An aardwolf can eat 300,000 ''Trinervitermes'' on a single outing.<ref name="mac119" />

Except for the aardwolf, hyenas are known to drive off larger predators, like lions, from their kills, despite having a reputation in popular culture for being cowardly.<ref name="p72"/> Hyenas are primarily nocturnal animals, but sometimes venture from their lairs in the early-morning hours. With the exception of the highly social spotted hyena, hyenas are generally not gregarious animals, though the striped and brown hyenas may live in family groups and congregate at kills.<ref name="r343">{{Harvnb|Rosevear|1974|pp=343–344}}</ref>

Spotted hyenas are one of the few mammals other than bats known to survive infection with rabies virus<ref>{{cite journal|date=18 December 2001|title=Regular exposure to rabies virus and lack of symptomatic disease in Serengeti spotted hyenas|journal=PNAS|volume=98|issue=26|doi=10.1073/pnas.261411898|pmid=11742089| last1=East| first1=M.L.|pages=15026–31|pmc=64977|bibcode=2001PNAS...9815026E|doi-access=free}}</ref> and have shown little or no disease-induced mortality during outbreaks in sympatric carnivores, in part due to the high concentration of [[Antibody|antibodies]] present in their saliva.<ref>{{cite journal|date=7 October 2015|title=Markedly Elevated Antibody Responses in Wild versus Captive Spotted Hyenas Show that Environmental and Ecological Factors Are Important Modulators of Immunity|journal=PLOS ONE|doi=10.1371/journal.pone.0137679|pmid=26444876| last1=Flies| first1=A.S.|display-authors=etal|volume=10|issue=10|article-number=e0137679|pmc=4621877|bibcode=2015PLoSO..1037679F|doi-access=free}}</ref> Despite this perceived unique disease resistance, little is known about the immune system of spotted hyenas,<ref>{{cite journal|date=29 February 2016|title=Socioecological predictors of immune defenses in a wild spotted hyenas|journal=Functional Ecology|volume=30|issue=9|doi=10.1111/1365-2435.12638|pmid=27833242| last1=Flies| first1=A.S.|display-authors=etal|pages=1549–1557|pmc=5098940|bibcode=2016FuEco..30.1549F }}</ref><ref>{{cite journal|date=15 January 2012|title=Development of a hyena immunology toolbox|journal=Veterinary Immunology and Immunopathology|volume=145|issue=1–2|doi=10.1016/j.vetimm.2011.10.016|pmid=22173276| last1=Flies| first1=A.S.|display-authors=etal|pages=110–9|pmc=3273618}}</ref><ref>{{cite journal|date=2 February 2014|title=Characterization of toll-like receptors 1-10 in spotted hyenas|journal=Veterinary Research Communications|volume=38|doi=10.1007/s11259-014-9592-3|pmid=24488231| last1=Flies| first1=A.S.|display-authors=etal|issue=2|pages=165–70|pmc=4112752}}</ref> and even less is known about other Hyaenidae species.

==Relationships with humans==
===Folklore, mythology and literature===
[[File:20,000 Year Old Cave Paintings Hyena.png|thumb|[[Cave hyena]] (''Crocuta crocuta spelaea'') painting found in the [[Chauvet Cave]] in 1994]]
[[File:Hyenamosaic.jpg|thumb| A depiction of the legendary striped hyena, Krokottas of Kytheros Island, from the [[Nile mosaic of Palestrina]]]]

Spotted hyenas vary in their folkloric and mythological depictions, depending on the ethnic group from which the tales originate. It is often difficult to know whether spotted hyenas are the specific hyena species featured in such stories, particularly in West Africa, as both spotted and striped hyenas are often given the same names. In West African tales, spotted hyenas are sometimes depicted as bad [[Muslim]]s who challenge the local [[animism]] that exists among the [[Beng language|Beng]] in [[Côte d'Ivoire]].<ref name="magic"/> 

In [[East Africa]], [[Tabwa]] mythology portrays the spotted hyena as a solar animal that first brought the sun to warm the cold earth, while West African folklore generally shows the hyena as symbolizing immorality, dirty habits, the reversal of normal activities, and other negative traits. In [[Tanzania]], there is a belief that [[witch]]es use spotted hyenas as [[Riding animal|mounts]].<ref name="magic"/> In the [[Mtwara Region]] of Tanzania, it is believed that a child born at night while a hyena is crying will likely grow up to be a thief. In the same area, hyena feces are believed to enable a child to walk at an early age, thus it is not uncommon in that area to see children with hyena dung wrapped in their clothes.<ref name="m97">{{Harvnb|Mills|Hofer|1998|p=97}}</ref> The [[Kaguru]] of Tanzania and the [[Kujamaat]] of southern [[Senegal]] view hyenas as [[Food and drink prohibitions|inedible]] and greedy hermaphrodites. A mythical African tribe called the [[Bouda]] is reputed to have members able to transform into hyenas.<ref name="AP">{{Cite journal | url=http://www.highbeam.com/doc/1G1-17909878.html| archive-url=https://web.archive.org/web/20110811101045/http://www.highbeam.com/doc/1G1-17909878.html| archive-date=2011-08-11| title=The spotted hyena from Aristotle to the Lion King: reputation is everything - In the Company of Animals| journal=Stephen E. Glickman| date=22 September 1995| access-date=2007-05-22}}</ref> A similar myth occurs in [[Mansôa]]. These "[[werehyena]]s" are killed when discovered, and do not revert to human form once dead.<ref name="m97"/>

Striped hyenas are often referred to in [[Middle East]]ern literature and folklore, typically as symbols of treachery and stupidity.<ref name="ABI">Mounir R. Abi-Said (2006) ''Reviled as a grave robber: The ecology and conservation of striped hyaenas in the human dominated landscapes of Lebanon'' Ph.D. thesis, University of Kent (Biodiversity management)</ref> In the Near and Middle East, striped hyenas are generally regarded as physical incarnations of [[genie|jinn]]s.<ref name="magic">Frembgen, Jürgen W. ''[http://nirc.nanzan-u.ac.jp/nfile/364 The Magicality of the Hyena: Beliefs and Practices in West and South Asia] {{Webarchive|url=https://web.archive.org/web/20160412175143/http://nirc.nanzan-u.ac.jp/nfile/364 |date=2016-04-12 }}'', Asian Folklore Studies, Volume 57, 1998: 331–344</ref> Arab writer [[Abu Yahya Zakariya' ibn Muhammad al-Qazwini|al-Qazwīnī]] (1204–1283) spoke of a tribe of people called ''al-Ḍabyūn'' meaning "hyena people". In his book ''<nowiki />'Ajā'ib Al-Makhlūqāt'' he wrote that should one of this tribe be in a group of 1,000 people, a hyena could pick him out and eat him.<ref name="ABI" /> A [[Persia]]n medical treatise written in 1376 tells how to cure cannibalistic people known as ''kaftar'', who are said to be "half-man, half-hyena".<ref name="magic" /> [[Al-Damiri|al-Damīrī]] in his writings in ''Ḥawayān al-Kubrā'' (1406) wrote that striped hyenas were [[vampiric]] creatures that attacked people at night and sucked the blood from their necks. He also wrote that hyenas only attacked brave people. Arab folklore tells of how hyenas can mesmerise victims with their eyes or sometimes with their pheromones.<ref name="ABI" /> 

In a similar vein to al-Damīrī, the [[Greek people|Greeks]] until the end of the 19th century believed that the bodies of werewolves, if not destroyed, would haunt battlefields as vampiric hyenas that drank the blood of dying soldiers.<ref name="Woodward">{{cite book| last=Woodward|first=Ian| title=The Werewolf Delusion| year=1979| page=256|publisher=Paddington Press | isbn=0-448-23170-0 }}</ref> The image of striped hyenas in [[Afghanistan]], India and [[Palestine (region)|Palestine]] is more varied. Though feared, striped hyenas were also symbolic of love and fertility, leading to numerous varieties of love medicine derived from hyena body parts. Among the [[Baluch people|Baluch]] and in [[northern India]], witches or magicians are said to ride striped hyenas at night.<ref name="magic" />

The striped hyena is mentioned in the [[Bible]]. The [[Arabic]] word for the hyena, {{Lang|ar-Latn|ḍab`}} or {{Lang|ar-Latn|ḍabu`}} (plural {{Lang|ar-Latn|ḍibā`}}), is alluded to in a valley in [[Israel]] known as Shaqq-ud-Diba` (meaning "cleft of the hyenas") and Wadi-Abu-Diba` (meaning "valley of the hyenas"). Both places have been interpreted by some scholars as being the Biblical Valley of [[Zeboim (Hebrew Bible)|Tsebo`im]] mentioned in 1 [[Book of Samuel|Samuel]] 13:18. In modern [[Hebrew language|Hebrew]], the word for hyena and hypocrite are both the same: [[wikt:צבוע|''tsavua'']]. Though the [[Authorized King James Version]] of the Bible interprets the term "''`ayit tsavua`''" (found in [[Book of Jeremiah|Jeremiah]] 12:9) as "speckled bird", [[Henry Baker Tristram]] argued that it was most likely a hyena being mentioned.<ref name="BOTF">{{cite book| last=Bright|first=Michael| title=Beasts of the Field: The Revealing Natural History of Animals in the Bible| year=2006| pages=[https://archive.org/details/beastsoffieldrev0000brig/page/127 127–129]|publisher=Pavilion Books | isbn=1-86105-831-4| url=https://archive.org/details/beastsoffieldrev0000brig/page/127 }}</ref>

The vocalization of the spotted hyena resembling hysterical human [[laughter]] has been alluded to in numerous works of literature: "to laugh like a hyæna" or a "hyen" was a common [[simile]], and is featured in ''The Cobbler's Prophecy'' (1594), Webster's ''[[The Duchess of Malfi|Duchess of Malfi]]'' (1623) and [[Shakespeare]]'s ''[[As You Like It]]'', Act IV. Sc.1.<ref>{{Cite web |title=The Cobler's Prophecy, by Robert Wilson |url=https://archive.org/details/coblersprophecy100wilsrich/page/n3/mode/2up |access-date=2025-10-11 |website=archive.org}}</ref><ref>{{Cite web |title=The Duchess of Malfi, by John Webster |url=https://www.gutenberg.org/files/2232/2232-h/2232-h.htm |access-date=2025-01-25 |website=www.gutenberg.org}}</ref><ref>{{Cite web |title=As You Like It: Entire Play |url=https://shakespeare.mit.edu/asyoulikeit/full.html |access-date=2025-01-25 |website=shakespeare.mit.edu}}</ref>

''Die Strandjutwolf'' (The [[brown hyena]]) is an [[allegory|allegorical]] poem by the renowned South African poet, [[N. P. van Wyk Louw]], which evokes a sinister and ominous presence.<ref>{{cite journal |doi=10.17159/2309-9070/tvl.v.55i3.5506 |title=Symbolic values of the dog in Afrikaans literature |date=2018 |last1=Taljaard-Gilson |first1=Gerda |journal=Tydskrif vir Letterkunde |volume=55 |issue=3 }}</ref>

===Attacks on humans===
Ordinarily, striped hyenas are extremely timid around humans, though they may show bold behaviors towards people at night.<ref name="h36">{{Harvnb|Heptner|Sludskii|1992|p=36}}</ref> On rare occasions, striped hyenas have preyed on humans.

Among hyenas, only the spotted and striped hyenas have been known to become [[Man-eating animal|man-eater]]s. Hyenas are known to have preyed on humans in prehistory: human hair has been found in fossilized hyena dung dating back 195,000 to 257,000 years.<ref>{{cite web|url=http://dsc.discovery.com/news/2009/02/10/oldest-human-hair.html|title=Oldest Human Hair Found in Fossilized Dung|first=Jennifer|last=Viegas|website=Discovery News|access-date=29 November 2020|date=10 February 2009|archive-url=https://web.archive.org/web/20100107023507/http://dsc.discovery.com/news/2009/02/10/oldest-human-hair.html|archive-date=7 January 2010}}</ref> Some paleontologists believe that competition and predation by [[cave hyena]]s (''Crocuta crocuta spelaea'') in [[Siberia]] was a significant factor in delaying human colonization of [[Alaska]]. Hyenas may have occasionally stolen human kills, or entered campsites to drag off the young and weak, much as modern spotted hyenas do in Africa. The oldest Alaskan human remains coincide with roughly the same time cave hyenas became extinct, leading some paleontologists to infer that hyena predation prevented humans from crossing the [[Bering Strait]] earlier.<ref name="Siberia">{{cite web|url=http://www.asu.edu/provost/emerituscollege/EVoice1/Articles%20section/n1%20Turner%206x9.pdf|title=Hyenas and Humans in Ice Age Siberia|work=Christy G. Turner II|publisher=School of Human Evolution and Social Change, Arizona State University|access-date=2008-08-02 }}{{dead link|date=May 2016|bot=medic}}{{cbignore|bot=medic}}</ref> 

Hyenas readily scavenge from human corpses; in Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the [[1960 Ethiopian coup attempt|1960 attempted coup]]<ref>[[Ryszard Kapuściński|Kapuściński, Ryszard]], ''[[The Emperor (book)|The Emperor: Downfall of an Autocrat]]''. 1978. {{ISBN|0-679-72203-3}}</ref> and the [[Red Terror (Ethiopia)|Red Terror]].<ref>Donham, Donald Lewis (1999) ''Marxist modern: an ethnographic history of the Ethiopian revolution'', University of California Press, page 135, {{ISBN|0-520-21329-7}}</ref> Hyenas habituated to scavenging on human corpses may develop bold behaviors towards living people: hyena attacks on people in southern Sudan increased during the [[Second Sudanese Civil War]], when human corpses were readily available to them.<ref>Copson, Raymond W. (1994) ''Africa's wars and prospects for peace'', M.E. Sharpe, page 6, {{ISBN|1-56324-300-8}}</ref>

Spotted hyenas have been known to prey on humans in modern times, but such incidents are rare. However, attacks on humans by spotted hyenas are likely to be underreported.<ref name="SGDRN">Begg, Colleen, Begg, Keith & Muemedi, Oscar (2007) ''[http://www.selous-niassa-corridor.org/fileadmin/publications/2007-Human-Carnivore_Crocodile_Conflict_-NNR.pdf Preliminary data on human - carnivore conflict in Niassa National Reserve, Mozambique, particularly fatalities due to lion, spotted hyaena and crocodile] {{Webarchive|url=https://web.archive.org/web/20111226170051/http://www.selous-niassa-corridor.org/fileadmin/publications/2007-Human-Carnivore_Crocodile_Conflict_-NNR.pdf|date=2011-12-26 }}'', SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique)</ref> Man-eating spotted hyenas tend to be very large specimens; a pair of man-eating hyenas, responsible for killing 27 people in [[Mulanje]], [[Malawi]] in 1962, weighed in at 72&nbsp;kg (159&nbsp;lb) and 77&nbsp;kg (170&nbsp;lb) after being shot.<ref name="maneat">Kruuk, Hans (2002) ''Hunter and hunted: relationships between carnivores and people'' Cambridge University Press, {{ISBN|0-521-89109-4}}</ref> A 1903 report describes spotted hyenas in the [[Mzimba District|Mzimba district]] of [[Angoniland]] waiting at dawn outside people's huts to attack them when they opened their doors.<ref name="NE"/> Victims of spotted hyenas tend to be women, children and sick or infirm men; [[Theodore Roosevelt]] wrote in 1908–1909 in [[Uganda]] that spotted hyenas regularly killed sufferers of [[African sleeping sickness]] as they slept outside in camps.<ref>Roosevelt, Theodore (1910) ''African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist'', New York, C. Scribner's sons</ref> 

Spotted hyenas are widely feared in Malawi, where they have been known to attack people at night, particularly during the hot season when people sleep outside. A spate of hyena attacks was reported in Malawi's [[Phalombe]] plain, with five deaths recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961, when eight people were killed. Attacks occurred most commonly in September, when people slept outdoors and bush fires made the hunting of wild game difficult for the hyenas.<ref name="SGDRN"/><ref name="NE">{{cite book| last=Knight|first=John| title=Natural Enemies: People-Wildlife conflicts in Anthropological Perspective| year=2000|publisher=Psychology Press | isbn=0-415-22441-1 }}</ref> A 2004 news report stated that 35 people were killed by spotted hyenas in a 12-month period in [[Mozambique]] along a 20-km stretch of road near the [[Tanzania]]n border.<ref name="SGDRN"/>

In the 1880s, a hyena was reported to have attacked humans, especially sleeping children, over a three-year period in the [[Iğdır Province]] of Turkey, with 25 children and 3 adults being wounded in one year. Further attacks were reported later in some parts of the [[South Caucasus]], particularly in 1908. Instances are known in [[Azerbaijan]] of striped hyenas killing children sleeping in courtyards during the 1930s and 1940s. In 1942, a sleeping guard was mauled in his hut by a hyena in [[Qalıncaq]] (Golyndzhakh).<!-- Through Rissian: В 1942 г., в Исмаиллинском районе (с. Голынджах) гиена проникла в шалаш, где спал сторож, и нанесла ему ранения (Ф. Ф. Алиев) --> Cases of children being taken by hyenas by night are known in southeast Turkmenistan's [[Bathyz Nature Reserve]]. A further attack on a child was reported around [[Serakhs]] in 1948.<ref name="h46">{{Harvnb|Heptner|Sludskii|1992|p=46}}</ref> Several attacks have occurred in India; in 1962, 9 children were thought to have been taken by hyenas in the town of [[Bhagalpur]] in the [[Bihar]] State in a six-week period,<ref name="BOTF"/> and 19 children up to the age of four were killed by hyenas in [[Karnataka]] in 1974.<ref name="m25">{{Harvnb|Mills|Hofer|1998|p=25}}</ref> A survey of wild animal attacks during a five-year period in the Indian state of [[Madhya Pradesh]] reported that hyenas had attacked three people, causing fewer deaths than [[Indian wolf|wolves]], [[gaur]], [[boar]], [[Asian elephant|elephants]], [[Bengal tiger|tigers]], [[Indian leopard|leopards]] and [[sloth bear]]s.<ref name="Attacks">{{cite web| url=http://www.nina.no/archive/nina/Publikasjoner/oppdragsmelding/NINA-OM731.pdf| archive-url=https://web.archive.org/web/20070927173051/http://www.nina.no/archive/nina/Publikasjoner/oppdragsmelding/NINA-OM731.pdf| archive-date=2007-09-27| title=The Fear of Wolves: A Review of Wolf Attacks on Humans| publisher=Norsk Institutt for Naturforskning| access-date=2008-06-26}}</ref>

<gallery widths=250 heights=250>
File:Hyena attack (1842 woodcut).jpg|Illustration from ''Fraser's'' magazine showing an artist's impression of a "stag-hound" biting a spotted hyena attacking its master
File:Bodleian Libraries, Handbill of Merchant's Hall, 1739, announcing A lion, lionesses, tigers, etc..jpg|A 1739 advertisement by [[Charles Benjamin Incledon (advertiser)|Charles Benjamin Incledon]] featuring [[feliform]]s: the [[Mesopotamian lion]] from the vicinity of [[Bassorah]], [[Cape lion]], [[tiger]] from the [[East Indies]], [[jaguar|panther]] from [[Buenos Aires]], ''[[Hyaena hyaena]]'' from [[West Africa]], and [[leopard]] from [[Turkey]], besides a "[[Lampago|Man tyger]]" from [[Africa]]. The advertisement mentions that the 'hyaena' can mimic a human voice to lure humans.
</gallery>

===Hyenas as food and medicine===
Hyenas have occasionally been used for food and medicinal purposes in [[Somalia]].<ref>{{cite web|url=http://www.somalilandpress.com/islamists-authorise-hyena-meat-in-southern-somalia/|title=Islamists authorise hyena meat in Southern Somalia.|work=Somalilandpress|access-date=2017-01-03|archive-url=https://web.archive.org/web/20170103170353/http://www.somalilandpress.com/islamists-authorise-hyena-meat-in-southern-somalia/|archive-date=2017-01-03|date=August 12, 2012}}</ref> Some Muslims consider it [[halal]] in [[Islam]]<ref>{{Cite web|url=https://sunnah.com/tirmidhi:851|title=Jami' at-Tirmidhi 851 - The Book on Hajj - كتاب الحج عن رسول الله صلى الله عليه وسلم - Sunnah.com - Sayings and Teachings of Prophet Muhammad (صلى الله عليه و سلم)|website=sunnah.com}}</ref><ref>{{Cite web |date=April 25, 2022 |title=#368: Eating Hyena |url=https://maktabahalbakri.com/368-eating-hyena/ |access-date=February 8, 2025 |website=maktabahalbakri.com |archive-date=October 16, 2022 |archive-url=https://web.archive.org/web/20221016100722/https://maktabahalbakri.com/368-eating-hyena/ }}</ref><ref>{{Cite web |date=August 23, 2009 |title=Ruling on Eating the Meat of Hyena |url=https://www.aliftaa.jo/research-fatwa-english/321/Ruling-on-Eating-the-Meat-of-Hyena- |access-date=February 8, 2025 |website=Fatwa Department of the Hashemite Kingdom of Jordan}}</ref> although this is disputed by other Muslims.<ref>{{Cite web|url=https://islam.stackexchange.com/questions/62942/confusion-on-hyenas|title=Confusion on hyenas|website=Islam Stack Exchange}}</ref><ref>{{Cite web|url=https://qna.ilmhub.com/question/ruling-on-consuming-hyena-meat/|title=Ruling on consuming hyena meat}}</ref> This practice dates back to the times of the [[Ancient Greeks]] and [[Ancient Rome|Romans]], who believed that different parts of the hyena's body were effective means to ward off evil and to ensure love and fertility.<ref name="magic" />

==References==
=== Citations ===
{{Reflist}}

=== General and cited references ===
* {{Cite book|last1=Heptner|first1=V. G.|last2=Sludskii|first2=A. A.|url=https://archive.org/details/mammalsofsov221992gept|title=Mammals of the Soviet Union: Carnivora (hyaenas and cats), Volume 2|publisher=Smithsonian Institution Libraries and National Science Foundation|year=1992 }}
* {{Cite book|last=Kruuk|first=Hans|title=The Spotted Hyena: A Study of Predation and Social Behaviour|publisher=University of California Press|year=1972}}
* {{Cite book|last=Kurtén|first=Björn|title=Pleistocene mammals of Europe|publisher=Weidenfeld and Nicolson|year=1968}}
* {{Cite book|last=Macdonald|first=David|url=https://archive.org/details/velvetclawnatura00macd|title=The Velvet Claw: A Natural History of the Carnivores |location=New York |publisher=Parkwest |year=1992|isbn=0-563-20844-9}}
* {{Cite book|last1=Mills|first1=Gus|author-link1=Gus Mills|last2=Hofer|first2=Heribert|url=http://data.iucn.org/dbtw-wpd/edocs/1998-013.pdf|title=Hyaenas: status survey and conservation action plan|publisher=IUCN/SSC Hyena Specialist Group|year=1998|isbn=2-8317-0442-1|archive-url=https://web.archive.org/web/20130506084714/http://data.iucn.org/dbtw-wpd/edocs/1998-013.pdf|archive-date=May 6, 2013 }}
* {{Cite book|last1=Mills|first1=Gus|author-link1=Gus Mills|last2=Mills|first2=Margie|title=Hyena Nights and Kalahari Days|publisher=Jacana Education|year=2010|isbn=978-1-77009-811-4}}
* {{Cite book|last=Pocock|first=R. I.|url=https://archive.org/details/PocockMammalia2|title=Fauna of British India: Mammals Volume 2|publisher=Taylor and Francis|year=1941}}
* {{Cite book|last=Rosevear|first=Donovan Reginald|url=https://archive.org/details/carnivoresofwest00rose|title=The carnivores of West Africa |location=London |publisher=Trustees of the British Museum (Natural History)|year=1974|isbn=0-565-00723-8 }}

==Further reading==
* Funk, Holdger (2010). ''Hyaena: On the Naming and Localisation of an Enigmatic Animal''. GRIN Verlag. {{ISBN|3-640-69784-7}}.
* Lawick, Hugo & Goodall, Jane (1971). ''Innocent Killers''. Boston: Houghton Mifflin Company.
* Mills, M. G. L. (2003). ''Kalahari Hyenas: Comparative Behavioral Ecology of Two Species''. The Blackburn Press.

==External links==
{{Commons and category|Hyaenidae|Hyaenidae}}
{{Wiktionary}}
{{Wikispecies|Hyaenidae}}
{{Wikiquote|Hyena}}
* [https://web.archive.org/web/20110507123347/http://www.hyaenidae.org/ IUCN Conservation Union Hyaena Specialist Group]

{{Carnivora|F.}}
{{Taxonbar|from=Q42046}}
{{Authority control}}

{{DEFAULTSORT:Hyena}}
[[Category:Hyenas| ]]
[[Category:Extant Miocene first appearances]]
[[Category:Mammals of Africa]]
[[Category:Mammals of Asia]]
[[Category:Necrophages]]
[[Category:Taxa named by John Edward Gray]]

[[eo:Hieno]]