Editing Dhole (section)

==Ecology and behaviour==
Dholes produce whistles resembling the calls of red foxes, sometimes rendered as ''coo-coo''. How this sound is produced is unknown, though it is thought to help in coordinating the pack when travelling through thick brush. When attacking prey, they emit screaming ''KaKaKaKAA'' sounds.<ref name="f93">{{Harvnb|Fox|1984|p=93}}</ref> Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.<ref name="f95">{{Harvnb|Fox|1984|p=95}}</ref> In contrast to wolves, dholes do not [[Howling|howl]] or bark.<ref name="heptner1998"/>

Dholes have a complex [[body language]]. Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes pucker their lips forward in a snarl and raise the hairs on their backs, as well as keep their tails horizontal or vertical. When afraid, they pull their lips back horizontally with their tails tucked and their ears flat against the skull.<ref name="f97">{{Harvnb|Fox|1984|p=97}}</ref>

===Social and territorial behaviour===
Dholes are more social than [[gray wolves]],<ref name="heptner1998"/> and have less of a dominance hierarchy, as seasonal scarcity of food is not a serious concern for them. In this manner, they closely resemble African wild dogs in social structure.<ref name="f85">{{Harvnb|Fox|1984|p=85}}</ref> They live in [[clan]]s rather than [[pack (canine)|pack]]s, as the latter term refers to a group of animals that always hunt together. In contrast, dhole clans frequently break into small packs of three to five animals, particularly during the spring season, as this is the optimal number for catching fawns.<ref name="f81">{{Harvnb|Fox|1984|pp=81–2}}</ref> Dominant dholes are hard to identify, as they do not engage in dominance displays as wolves do, though other clan members will show submissive behaviour toward them.<ref name="f86"/> Intragroup fighting is rarely observed.<ref name="nowak1983"/>

[[File:20140303 7687 Pench Dhole.jpg|thumb|Dholes playing, [[Pench National Park]]]]
[[File:Asiatic Wild Dogs (43518944044).jpg|thumb|A pack of Asiatic wild dog]]

Dholes are far less [[territory (animal)|territorial]] than wolves, with pups from one clan often joining another without trouble once they mature sexually.<ref name="f92">{{Harvnb|Fox|1984|p=92}}</ref> Clans typically number 5 to 12 individuals in India, though clans of 40 have been reported. In [[Thailand]], clans rarely exceed three individuals.<ref name="cohen1978"/> Unlike other canids, there is no evidence of dholes using [[urine]] to mark their territories or travel routes. When urinating, dholes, especially males, may [[Raised-leg urination|raise one hind leg]] or both to result in a handstand. Handstand urination is also seen in [[bush dog]]s (''Speothos venaticus'')<ref>{{cite journal |title=Einige beobachtungen zum verhalten des Dekkan-Rothundes (Cuon alpinus dukhunensis Sykes) im Kanha National Park |trans-title=Some observations on the behavior of the Deccan Red Dog (Cuon alpinus dukhunensis Sykes) in Kanha National Park |last1=Keller |first1=R. |journal= Vierteljahresschrift. Naturf. Ges. Zürich |volume=118 |year=1973|pages=129–135 |language=de |url= https://www.ngzh.ch/archiv/1973_118/118_1/118_13.pdf |archive-url=https://web.archive.org/web/20190812095239/https://www.ngzh.ch/archiv/1973_118/118_1/118_13.pdf |archive-date=2019-08-12 |url-status=live }}</ref> and domestic dogs.<ref>{{cite journal |last1=Wirant |first1=Sharon Cudd |first2=Betty |last2=McGuire |url=https://www.researchgate.net/publication/248335168 |title=Urinary behavior of female domestic dogs (''Canis familiaris''): influence of reproductive status, location, and age |journal=Applied Animal Behaviour Science |volume=85 |issue=3–4 |year=2004 |pages=335–348|doi=10.1016/j.applanim.2003.09.012 }}</ref> They may defecate in conspicuous places, though a territorial function is unlikely, as [[faeces]] are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal [[Animal latrine|latrines]]. They do not scrape the earth with their feet, as other canids do, to mark their territories.<ref name="f97"/>

===Denning===
Four kinds of den have been described; simple earth dens with one entrance (usually remodeled [[striped hyena]] or [[porcupine]] dens); complex cavernous earth dens with more than one entrance; simple cavernous dens excavated under or between rocks; and complex cavernous dens with several other dens in the vicinity, some of which are interconnected. Dens are typically located under dense scrub or on the banks of dry rivers or creeks. The entrance to a dhole den can be almost vertical, with a sharp turn three to four feet down. The tunnel opens into an antechamber, from which extends more than one passage. Some dens may have up to six entrances leading up to {{cvt|30|m}} of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together.<ref name="f43">{{Harvnb|Fox|1984|pp=43–49}}</ref> Like [[African wild dog]]s and [[dingo]]es, dholes will avoid killing prey close to their dens.<ref name="f80"/>

===Reproduction and development===
[[File:Cuon alpinus alpinus puppy.jpg|thumb|Dhole pup, [[Kolmården Wildlife Park]]]]
In India, the [[mating season]] occurs between mid-October and January, while captive dholes in the [[Moscow Zoo]] breed mostly in February.<ref name="cohen1978"/> Unlike wolf packs, dhole clans may contain more than one breeding female.<ref name="f86"/> More than one female dhole may den and rear their litters together in the same den.<ref name="nowak1983"/> During [[mating]], the female assumes a crouched, cat-like position. There is no [[copulatory tie]] characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation.<ref name="f79">{{Harvnb|Fox|1984|p=79}}</ref> The [[gestation period]] lasts 60–63 days, with litter sizes averaging four to six pups.<ref name="cohen1978"/> Their growth rate is much faster than that of wolves, being similar in rate to that of [[coyote]]s.

The [[hormone]] metabolites of five males and three females kept in Thai zoos was studied. The breeding males showed an increased level of [[testosterone]] from October to January. The [[oestrogen]] level of captive females increases for about two weeks in January, followed by an increase of [[progesterone]]. They displayed sexual behaviours during the oestrogen peak of the females.<ref>{{cite journal |author1=Khonmee, J. |name-list-style=amp |author2= Rojanasthien, S. |author3=Thitaram, C. |author4=Sumretprasong, J. |author5= Aunsusin, A. |author6=Chaisongkram, C. |author7=Songsasen, N. |title=Non-invasive endocrine monitoring indicates seasonal variations in gonadal hormone metabolites in dholes (''Cuon alpinus'') |date=2017 |journal=Conservation Physiology |volume=5 |issue=1 |pages=cox001 |doi= 10.1093/conphys/cox001 |pmid= 28852505|pmc=5570072 |doi-access=free }}</ref>

Pups are suckled at least 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use [[wikt:rendezvous|rendezvous]] sites to meet their pups as wolves do, though one or more adults will stay with the pups at the den while the rest of the pack hunts. Once [[weaning]] begins, the adults of the clan will regurgitate food for the pups until they are old enough to join in hunting. They remain at the den site for 70–80 days. By the age of six months, pups accompany the adults on hunts and will assist in killing large prey such as [[sambar deer|sambar]] by the age of eight months.<ref name="f80">{{Harvnb|Fox|1984|p=80}}</ref> Maximum longevity in captivity is 15–16 years.<ref name="nowak1983"/>

===Hunting behaviour===
Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and mounting.<ref name="f100">{{Harvnb|Fox|1984|pp=100–101}}</ref> Dholes are primarily [[Diurnality|diurnal]] hunters, hunting in the early hours of the morning. They rarely hunt at night, except on moonlit nights, indicating they greatly rely on [[Visual perception|sight]] when hunting.<ref name="f50">{{Harvnb|Fox|1984|p=50}}</ref> They can chase their prey for many hours.<ref name="heptner1998"/> During a pursuit, one or more dholes takes over chasing the prey, while the rest of the pack keeps up at a steadier pace behind, taking over once the other group tires. Most chases are short, lasting only {{cvt|500|m}}.<ref name="f73">{{Harvnb|Fox|1984|p=73}}</ref> When chasing fleet-footed prey, they run at a pace of {{cvt|30|mph|km/h|-1|order=flip}}.<ref name="heptner1998"/> Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.<ref name="f67">{{Harvnb|Fox|1984|p=67}}</ref>

[[File:2012-bandipur-dhole-sambar.jpg|thumb|Dholes attacking a [[sambar (deer)|sambar]], [[Bandipur National Park]]]]

Once large prey is caught, one dhole grabs the prey's nose, while the rest of the pack pulls the animal down by the flanks and hindquarters. They do not use a killing bite to the throat.<ref name="f61">{{Harvnb|Fox|1984|p=61}}</ref> They occasionally blind their prey by attacking the eyes.<ref>{{cite journal |author=Grassman, L. I. Jr. |author2=M. E. Tewes |author3=N. J. Silvy |author4=K. Kreetiyutanont |year=2005 |title=Spatial ecology and diet of the dhole ''Cuon alpinus'' (Canidae, Carnivora) in north central Thailand |name-list-style= amp| journal=Mammalia |volume=69 |issue=1 |pages=11–20 |s2cid= 85198149 |doi= 10.1515/mamm.2005.002 |url= http://www.mnhn.fr/museum/front/medias/publication/6555_m05n1a2 |url-status=dead |archive-url=https://web.archive.org/web/20061123090326/http://www.mnhn.fr/museum/front/medias/publication/6555_m05n1a2 |archive-date=23 November 2006}}</ref> [[Serow]]s are among the only ungulate species capable of effectively defending themselves against dhole attacks, due to their thick, protective coats and short, sharp horns capable of easily impaling dholes.<ref name="lydekker1907"/> Dholes tear open their prey's flanks and [[Disembowelment|disembowel]] it, eating the [[heart]], [[liver]], [[lungs]] and some sections of the [[intestines]]. The [[stomach]] and [[rumen]] are usually left untouched.<ref name="f63">{{Harvnb|Fox|1984|p=63}}</ref> Prey weighing less than {{cvt|50|kg|lb}} is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes tear off pieces of the carcass and eat in seclusion.<ref name="f70">{{Harvnb|Fox|1984|p=70}}</ref> They give the pups access to a kill.<ref name="f86">{{Harvnb|Fox|1984|pp=86–87}}</ref> They are generally tolerant of [[scavenger]]s at their kills.<ref name="f51">{{Harvnb|Fox|1984|p=51}}</ref> Both mother and young are provided with regurgitated food by other pack members.<ref name="nowak1983"/>
{{clear}}

===Feeding ecology===
Prey animals in India include [[chital]], [[sambar deer]], [[muntjac]], [[mouse deer]], [[barasingha]], [[wild boar]], [[gaur]], [[water buffalo]]es, [[banteng]], [[cattle]], [[nilgai]], [[goat]]s, [[Indian hare]]s, [[Himalayan field rat]]s and [[langur]]s.<ref name="cohen1978"/><ref name=Pocock1941/><ref name="f58">{{Harvnb|Fox|1984|pp=58–60}}</ref> There is one record of a pack bringing down an [[Indian elephant]] calf in [[Assam]], despite desperate defense of the mother, resulting in numerous losses to the pack.<ref name="perry1965"/> In Kashmir, they prey on [[markhor]],<ref name=Pocock1941/> and [[thamin]] in Myanmar,<ref name="cohen1978"/> [[Malayan tapir]], [[Sumatran serow]] in [[Sumatra]] and the [[Malay Peninsula]] and [[Javan rusa]] in Java.<ref name=Durbin2004/> In the [[Tian Shan]] and [[Tarbagatai Mountains]], dholes prey on [[Siberian ibex]]es, [[arkhar]], [[roe deer]], [[Caspian red deer]] and [[wild boar]]. In the [[Altai Mountains|Altai]] and [[Sayan Mountains]], they prey on [[musk deer]] and [[reindeer]]. In eastern Siberia, they prey on roe deer, [[Manchurian wapiti]], wild pig, musk deer and reindeer, while in [[Primorye]] they feed on [[sika deer]] and [[goral]]. In Mongolia, they prey on [[argali]] and rarely Siberian ibex.<ref name="heptner1998"/>

[[File:2012-bandipur-dhole-chital.jpg|thumb|Dholes feeding on a [[chital]], Bandipur National Park]]
[[File:Dhole feeding Khao Yai NP.jpg|thumb|Dhole feeding on [[sambar deer]] carcass, [[Khao Yai National Park]]]]

Like African wild dogs, but unlike wolves, dholes are not known to actively hunt people.<ref name="heptner1998"/><ref name=Pocock1941/> They are known to eat [[insect]]s and [[lizard]]s.<ref>{{Cite web |last=Chacon |first=Raquel |title=Cuon alpinus (dhole) |url=https://animaldiversity.org/accounts/Cuon_alpinus/ |access-date=2023-04-04 |website=Animal Diversity Web |language=en |archive-date=27 January 2023 |archive-url=https://web.archive.org/web/20230127081104/https://animaldiversity.org/accounts/cuon_alpinus/ |url-status=live }}</ref> Dholes eat [[fruit]] and [[vegetable]] matter more readily than other canids. In captivity, they eat various kinds of grasses, herbs and leaves, seemingly for pleasure rather than just when ill.<ref name="mivart1890">{{cite book |last=Mivart |first=George |title=Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ |publisher=R.H. Porter |year=1890 |location=London |pages=177–188 |chapter=Genus ''Cyon'', Hodgson (1838) |chapter-url=https://archive.org/stream/dogsjackalswolve00mivauoft#page/n307/mode/2up}}</ref> In summertime in the Tian Shan Mountains, dholes eat large quantities of mountain [[rhubarb]].<ref name="heptner1998"/> Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.<ref name="f71">{{Harvnb|Fox|1984|p=71}}</ref> [[Livestock]] predation by dholes has been a problem in Bhutan since the late 1990s, as domestic animals are often left outside to graze in the forest, sometimes for weeks at a time. Livestock stall-fed at night and grazed near homes are never attacked. [[Oxen]] are killed more often than [[cow]]s, probably because they are given less protection.<ref>{{Cite journal |last1=Johnsingh |first1=A.J.T. |last2=Yonten |first2=D. |last3=Wangchuck |first3=S. |year=2007 |title=Livestock-Dhole Conflict in Western Bhutan |url=https://www.biodiversitylibrary.org/page/48380954 |journal=Journal of the Bombay Natural History Society |volume=104 |issue=2 |pages=201–202 |via=Biodiversity Heritage Library}}</ref>

===Enemies and competitors===
[[File:Dhole killed by leopard.jpg|thumb|Dhole killed and cached in a tree by a [[leopard]], India]]
In some areas, dholes are [[wiktionary:sympatric|sympatric]] to [[tiger]]s and [[leopard]]s. Competition between these species is mostly avoided through differences in prey selection, although there is still substantial dietary overlap. Along with leopards, dholes typically target animals in the {{cvt|30|–|175|kg}} range (mean weights of {{cvt|35.3|kg}} for dhole and {{cvt|23.4|kg}} for leopard), while tigers selected for prey animals heavier than {{cvt|176|kg}} (but their mean prey weight was {{cvt|65.5|kg}}). Also, other characteristics of the prey, such as sex, arboreality and aggressiveness, may play a role in prey selection. For example, dholes preferentially select male chital, whereas leopards kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopards' greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight.<ref name="jstor.org">{{cite journal |author1=Karanth, K. U. |author2=Sunquist, M. E. |jstor=5647 |title=Prey selection by tiger, leopard and dhole in tropical forests |journal=[[Journal of Animal Ecology]] |year=1995 |volume=64 |issue=4 |pages=439–450|doi=10.2307/5647 |bibcode=1995JAnEc..64..439K |name-list-style=amp}}</ref>

Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike.<ref name="perry1965">{{cite book |author=Perry, R. |year=1964 |title=The World of the Tiger |publisher=Cassell |location=London}}</ref> Dhole packs are smaller in areas with higher tiger densities due to tigers directly killing dholes and stealing kills they made. The kleptoparasitism causes dholes to prefer hunting smaller animals because they can eat more of a smaller carcass before a tiger arrives to steal it. Direct predation can lead to lower reproductive and recruitment rates, lower hunting success rates and less food for the pups when a helper is killed, and potentially pack destabilization if one member of the breeding pair is killed.<ref>{{cite journal |last1=Bhandari |first1=Aishwarya |last2=Ghaskadbi |first2=Pallavi |last3=Nigam |first3=Parag |last4=Habib |first4=Bilal |name-list-style=and |year=2021 |title=Dhole pack size variation: Assessing the effect of prey availability and apex predator |url= |journal=Ecology and Evolution |volume=11 |issue=9 |pages=4774–4785 |bibcode=2021EcoEv..11.4774B |doi=10.1002/ece3.7380 |pmc=8093734 |pmid=33976847}}</ref>

Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.<ref name=Pocock1941/> There are numerous records of leopards being treed by dholes.<ref name="nowak1983">{{cite book |author1=Walker, E. P. |author2=Nowak, R. M. |author3=Warnick, F. |year=1983 |title=Walker's Mammals of the World |edition=Fourth |publisher=Johns Hopkins University Press |location=Baltimore |name-list-style=amp}}</ref> Dholes were once thought to be a major factor in reducing [[Asiatic cheetah]] populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes.<ref name="fin1929">{{cite book |author=Finn, F. |year=1929 |title=Sterndale's Mammalia of India |publisher=Thacker, Spink & Co. |location=London}}</ref> Since leopards are smaller than tigers and are more likely to hunt dholes, dhole packs tend to react more aggressively toward them than they do towards tigers.<ref name="venkataraman">{{cite journal |author=Venkataraman, A. |year=1995 |title=Do dholes (''Cuon alpinus'') live in packs in response to competition with or predation by large cats? |journal=Current Science |volume=69 |issue=11 |pages=934–936 |url=http://eprints.iisc.ac.in/25085/1/article_id_069_11_0934_0936_0.pdf |archive-url=https://web.archive.org/web/20190713030406/http://eprints.iisc.ac.in/25085/1/article_id_069_11_0934_0936_0.pdf |archive-date=2019-07-13 |url-status=live}}</ref>

Dhole packs occasionally attack [[Asiatic black bear]]s, [[snow leopard]]s and [[sloth bear]]s. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves and lacerate their hindquarters.<ref name=Pocock1941/>
Although usually antagonistic toward [[wolves]],<ref name=heptner1998/> they may hunt and feed alongside one another.<ref name=shretha1997>{{cite book |author=Shrestha, T. J. |year=1997 |title=Mammals of Nepal: (with reference to those of India, Bangladesh, Bhutan and Pakistan) |publisher=Bimala Shrestha |location=Kathmandu |isbn=978-0-9524390-6-6}}</ref>

The dhole is also sympatric with the [[Indian wolf]] (''Canis lupus pallipes'') in parts of its range.<ref>{{cite journal |author=Mukherjee, S. |name-list-style=amp |author2=Zelcer, M. |author3=Kotler, B.P. |year=2009 |title=Patch use in time and space for a meso-predator in a risky world |journal=Oecologia |volume=159 |issue=3| pages=661–668 |doi=10.1007/s00442-008-1243-3 |pmid=19082629 |bibcode=2009Oecol.159..661M |s2cid=24051254}}</ref><ref>{{cite journal |author=Afik, D. |author2=Pinshow, B. |year=1993 |title=Temperature regulation and water economy in desert wolves |journal=Journal of Arid Environments |volume=24 |issue=2 |pages=197–209 |doi=10.1006/jare.1993.1017 |bibcode=1993JArEn..24..197A}}</ref> There is at least one record of a lone wolf associating with a pair of dholes in [[Debrigarh Wildlife Sanctuary]],<ref>{{cite journal |author1=Nair M. V. |name-list-style=amp |author2=Panda S. K. |year=2013 |title=Just Friends |journal=Sanctuary Asia |volume=XXXIII |page=3 |url=http://www.sanctuaryasia.com/magazines/features/9390-just-friends.html |access-date=23 December 2014 |archive-date=24 July 2019 |archive-url=https://web.archive.org/web/20190724195645/https://www.sanctuaryasia.com/magazines/features/9390-just-friends.html |url-status=dead }}</ref> and two observations in [[Satpura Tiger Reserve]].<ref>{{cite journal |author1=Ghaskadbi, P. |name-list-style=amp |author2=Das, J. |author3=Mahadev, V. |author4=Habib, B. |year=2021 |title=First record of mixed species association between dholes and a wolf from Satpura Tiger Reserve, India |journal=Canid Biology & Conservation |volume=23 |issue=4 |pages=15–17 |url=https://www.canids.org/CBC/23/Dhole_wolf_association.pdf |archive-url=https://web.archive.org/web/20211017142640/https://www.canids.org/CBC/23/Dhole_wolf_association.pdf |archive-date=2021-10-17 |url-status=live}}</ref> They infrequently associate in mixed groups with [[golden jackal]]s. Domestic dogs may kill dholes, though they will feed alongside them on occasion.<ref name=humphrey1990>{{cite book |author1=Humphrey, S. R. |author2=Bain, J. R. |name-list-style=amp |year=1990 |title=Endangered Animals of Thailand |publisher=Sandhill Crane Press |location=Gainesville |isbn=978-1-877743-07-8}}</ref>

===Diseases and parasites===
Dholes are vulnerable to a number of different diseases, particularly in areas where they are [[wiktionary:Sympatric|sympatric]] with other canid species. Infectious pathogens such as ''[[Toxocara canis]]'' are present in their faeces. They may suffer from [[rabies]], [[canine distemper]], [[mange]], [[trypanosomiasis]], [[canine parvovirus]] and [[endoparasite]]s such as [[cestode]]s and [[roundworm]]s.<ref name=Durbin2004/>