Editing Zebra (section)

==Behaviour and ecology==
[[File:Dust bathing mountain zebra, Namibia.jpg|thumb|right|alt=Mountain zebra dust bathing |Mountain zebra dustbathing in [[Namibia]]]]
Zebras may travel or [[animal migration|migrate]] to wetter areas during the dry season.<ref name="Grub 1981" /><ref name="Estes 1991" /> Plains zebras have been recorded travelling {{cvt|500|km|mi}} between Namibia and Botswana, the longest land migration of mammals in Africa.<ref>{{cite journal |last1=Naidoo|first1= R. |last2=Chase|first2= M. J. |last3=Beytall|first3= P. |last4=Du Preez|first4= P. |year=2016 |title=A newly discovered wildlife migration in Namibia and Botswana is the longest in Africa |journal=Oryx |volume=50 |issue=1 |pages=138–146 |doi=10.1017/S0030605314000222|doi-access=free }}</ref> When migrating, they appear to rely on some memory of the locations where foraging conditions were best and may predict conditions months after their arrival.<ref>{{cite journal |author=Bracis|first1= C. |last2=Mueller|first2= T. |year=2017 |title=Memory, not just perception, plays an important role in terrestrial mammalian migration |journal=Proceedings of the Royal Society B: Biological Sciences |volume=284 |issue=1855 |page=20170449 |doi=10.1098/rspb.2017.0449 |pmid=28539516 |pmc=5454266}}</ref> Plains zebras are more water-dependent and live in [[Mesic habitat|moister]] environments than other species. They usually can be found {{cvt|10|–|12|km}} from a water source.<ref name="Grub 1981" /><ref name="Estes 1991" /><ref name="Skinner" /> Grévy's zebras can survive almost a week without water but will drink it every day when given the chance, and their bodies maintain water better than cattle.<ref>{{cite journal |author=Youth|first= H. |title=Thin stripes on a thin line |url=http://nationalzoo.si.edu/Publications/ZooGoer/2004/6/grevys.cfm |url-status=dead |journal=Zoogoer |volume=33 |date=November–December 2004 |archive-date=26 October 2005  |archive-url=https://web.archive.org/web/20051026202556/http://nationalzoo.si.edu/Publications/ZooGoer/2004/6/grevys.cfm}}</ref><ref name="Churcher 1993" /> Mountain zebras can be found at elevations of up to {{cvt|2000|m}}.<ref>{{cite book |author=Woodward|first= Susan L. |year=2008 |title=Grassland Biomes |publisher=[[Greenwood Press]] |page=49 |isbn=978-0-313-33999-8}}</ref> Zebras sleep for seven hours a day, standing up during the day and lying down during the night. They regularly use various objects as rubbing posts and will [[dust bathing|roll on the ground]].<ref name="Estes 1991" />
[[File:Cebras de Burchell (Equus quagga burchellii), vista aérea del delta del Okavango, Botsuana, 2018-08-01, DD 30.jpg|thumb|left|alt=Plains zebras drinking at a river |Plains zebras at [[Okavango Delta]], [[Botswana]]]]
A zebra's diet is mostly [[grass]]es and [[sedge]]s, but they will opportunistically consume [[bark (botany)|bark]], leaves, buds, fruits, and roots. Compared to [[Ruminantia|ruminants]], zebras have a simpler and less efficient digestive system. Nevertheless, they can subsist on lower-quality vegetation. Zebras may spend 60–80% of their time feeding, depending on the availability of vegetation.<ref name="MacDonald" /><ref name="Estes 1991" /> The plains zebra is a pioneer grazer, mowing down the upper, less nutritious grass canopy and preparing the way for more specialised grazers like [[wildebeest]], which depend on shorter and more nutritious grasses below.<ref>{{cite book |last1=Pastor|first1= J.|last2= Cohen|first2= U.|last3= Hobbs|first3= T. |year=2006 |contribution=The roles of large herbivores in ecosystem nutrient cycles |editor=Danell|editor-first= K. |title=Large Herbivore Ecology, Ecosystem Dynamics and Conservation |url=https://archive.org/details/largeherbivoreec00dane_862 |url-access=limited |publisher=[[Cambridge University Press]] |page=[https://archive.org/details/largeherbivoreec00dane_862/page/n312 295] |isbn=978-0-521-53687-5}}</ref>

Zebras are preyed on mainly by lions. [[Leopards]], [[cheetahs]], [[spotted hyenas]], [[brown hyena]]s and [[African wild dog|wild dogs]] pose less of a threat to adults.{{sfn|Caro|2016|pp=61–63}} Biting and kicking are a zebra's defense tactics. When threatened by lions, zebras flee, and when caught they are rarely effective in fighting off the big cats.{{sfn|Caro|2016|p=61–62}} In one study, the maximum speed of a zebra was found to be {{cvt|50|km/h}} while a lion was measured at {{cvt|74|km/h}}. Zebras do not escape lions by speed alone but by sideways turning, especially when the cat is close behind.<ref>{{cite journal |author=Wilson|first1= A.|last2= Hubel|first2= T.|last3= Wilshin|first3= S. |display-authors=etal |year=2018 |title=Biomechanics of predator–prey arms race in lion, zebra, cheetah and impala |journal=Nature |volume=554 |issue= 7691|pages=183–188 |doi=10.1038/nature25479|pmid= 29364874|bibcode= 2018Natur.554..183W|s2cid= 4405091|url= https://researchonline.rvc.ac.uk/id/eprint/11143/1/11143.pdf}}</ref> With smaller predators like hyenas and dogs, zebras may act more aggressively, especially in defense of their young.{{sfn|Caro|2016|pp=62–63}}

===Social behaviour===
{{See also|Horse behaviour}}
[[File:Zebra Botswana edit02.jpg|thumb|upright|A plains zebra group |alt=A group of six plains zebra]]
Zebra species have two basic social structures. Plains and mountain zebras live in stable, closed family groups or [[Harem (zoology)|harems]] consisting of one [[stallion]], several [[mares]], and their offspring. These groups have their own [[home ranges]], which overlap, and they tend to be nomadic. Stallions form and expand their harems by herding young mares away from their birth harems. The stability of the group remains even when the family stallion is displaced. Plains zebras groups gather into large herds and may create temporarily stable subgroups within a herd, allowing individuals to interact with those outside their group. Females in harems can spend more time feeding, and gain protection both for them and their young. They have a linear [[Dominance (ethology)|dominance hierarchy]] with the high-ranking females having lived in the group longest. While traveling, the most dominant females and their offspring lead the group, followed by the next most dominant; the family stallion trails behind. Young of both sexes leave their natal groups as they mature; females are usually herded by outside males to become part of their harems.<ref name="MacDonald" /><ref name="Estes 1991" /><ref name="Rubenstein 1986"/>

In the more arid-living Grévy's zebras, adults have more fluid associations and adult males establish large [[Territory (animal)|territories]], marked by dung piles, and mate with the females that enter them.<ref name="Estes 1991" /><ref name="MacDonald" /> Grazing and drinking areas tend to be separated in these environments and the most dominant males establish territories near watering holes, which attract females with dependent foals and those who simply want a drink, while less dominant males control territories away from water with more vegetation, and only attract mares without foals.<ref name="Rubenstein 2010"/> Mares may travel through several territories but remain in one when they have young. Staying in a territory offers a female protection from harassment by outside males, as well as access to resources.<ref name="Rubenstein 1986">{{cite book |last=Rubenstein|first= D. I. |year=1986 |url=http://www.princeton.edu/~dir/pdf_dir/1986_Rubenstein_bookChapt.pdf |contribution=Ecology and sociality in horses and zebras |pages=282–302 |title=Ecological Aspects of Social Evolution |editor=Rubenstein|editor-first= D. I.|editor2= Wrangham|editor-first2= R. W. |publisher=[[Princeton University Press]] |isbn=978-0-691-08439-8}}</ref><ref>{{cite journal |last1=Sundaresan|first1=S. R.|last2=Fischhoff|first2=I. R.|last3=Rubenstein|first3=D.|year=2007|title= Male harassment influences female movements and associations in Grevy's zebra (''Equus grevyi'')|journal= Behavioral Ecology|volume= 18|issue= 5|pages= 860–65|doi= 10.1093/beheco/arm055 |url= http://www.princeton.edu/~equids/images/sundaresan_grevys_harassment.pdf|doi-access=free}}</ref>
[[File:Grevy's zebra group.jpg|thumb|left|alt=Three Grévy's zebras grazing |Group of Grévy's zebras grazing]]
In all species, excess males gather in [[bachelor herd|bachelor group]]s. These are typically young males that are not yet ready to establish a harem or territory.<ref name="MacDonald" /><ref name="Estes 1991" /> With the plains zebra, the oldest males are the most dominant and group membership is stable.<ref name="Estes 1991" /> Bachelor groups tend to be at the boundaries of herds and during group movements, the bachelors follow behind or along the sides.<ref name="Skinner">{{cite book |last1=Skinner |first1=J. D. |title=The Mammals of the Southern African Subregion |year=2005 |publisher=[[Cambridge University Press]] |isbn=978-0-521-84418-5 |pages=541–546 |edition=3rd |last2=Chimimba |first2=C. T. |chapter=Equidae}}</ref> Mountain zebra bachelor groups may also include young females that have left their natal group early, as well as old, former harem males. A territorial Grévy's zebra stallion may allow non-territorial bachelors in their territory, however when a mare in [[oestrous]] is present the territorial stallion keeps other stallions at bay. Bachelors prepare for their future harem roles with play fights and greeting/challenge rituals, which make up most of their activities.<ref name="Estes 1991" />

Fights between males usually occur over mates and involve biting and kicking. In plains zebra, stallions fight each other over recently matured mares to bring into their group and her father will fight off other males trying to abduct her. As long as a harem stallion is healthy, he is not usually challenged. Only unhealthy stallions have their harems taken over, and even then, the new stallion slowly takes over, peacefully displacing the old one. [[Agonistic behaviour]] between male Grévy's zebras occurs at the border of their territories.<ref name="Estes 1991" />

===Communication===
[[File:Cebras comunes (Equus quagga), parque nacional de Tarangire, Tanzania, 2024-05-25, DD 90.jpg|thumb|right|alt=A pair of Plains zebra facing each other and rubbing heads on the others body | Plains zebras mutually grooming]]
Zebras produce a number of vocalisations and noises. The plains zebra has a distinctive, [[Bark (sound)|barking]] contact call heard as "a-''ha'', a-''ha'', a-''ha''" or "kwa-ha, kaw-ha, ha, ha".<ref name="Grub 1981" /><ref name="Estes 1991" /> The mountain zebra may produce a similar sound while the call of Grévy's zebra has been described as "something like a [[hippo]]'s grunt combined with a donkey's wheeze". Loud snorting and rough "gasping" in zebras signals alarm. Squealing is usually made when in pain, but can also be heard in friendly interactions. Zebras also communicate with visual displays, and the flexibility of their lips allows them to make complex facial expressions. Visual displays also consist of head, ear, and tail postures. A zebra may signal an intention to kick by dropping back its ears and whipping its tail. Flattened ears, bared teeth and a waving head may be used as threatening gestures by stallions.<ref name="Estes 1991" />

Individuals may greet each other by mutually touching and rubbing, sniffing their genitals and resting their heads on their shoulders. They then may caress their shoulders against each other and lay their heads on one another. This greeting usually occurs between harem or territorial males or among bachelor males playing.<ref name="Estes 1991" /> Plains and mountain zebras strengthen their social bonds with [[Social grooming|grooming]]. Members of a harem nibble and rake along the neck, shoulder, and back with their teeth and lips. Grooming usually occurs between mothers and foals and between stallions and mares. Grooming establishes social rank and eases aggressive behaviour,<ref name="Estes 1991" />{{sfn|Caro|2016|p=143}} although Grévy's zebras generally do not perform social grooming.<ref name="Churcher 1993" />

===Reproduction and parenting===
{{See also|Horse breeding}}
[[File:Grévy's Zebra mating.jpg|thumb|left|alt=A pair of Grévy's zebras mating | Captive Grévy's zebras mating]]
Among plains and mountain zebras, the adult females mate only with their harem stallion, while in Grévy's zebras, mating is more [[Polygynandry|polygynandrous]] and the males have larger testes for [[sperm competition]].<ref name="Rubenstein 2010">{{cite book |last=Rubenstein|first= D. I. |year=2010 |url=https://dir.princeton.edu/include/pdf/2010_Rubenstein_ASB_vol42.pdf |contribution=Ecology, social behavior, and conservation in zebras |pages=231–258 |title=Behavioral Ecology of Tropical Animals |editor=Macedo|editor-first= R.|editor2= Wrangham|publisher=Academic Press|isbn=978-0123808943}}</ref><ref name="Ginsberg 1990">{{cite journal |last1=Ginsberg|first1= R|last2= Rubenstein|first2= D. I. |year=1990 |title=Sperm competition and variation in zebra mating behavior |journal=[[Behavioral Ecology and Sociobiology]] |volume=26 |issue=6 |pages=427–434 |doi=10.1007/BF00170901 |bibcode= 1990BEcoS..26..427G|s2cid= 206771095|url=http://www.princeton.edu/~dir/pdf_dir/1990_Ginsberg&dir_BehEcolSo.pdf}}</ref> Female zebras have five to ten day long oestrous cycles; physical signs include a swollen, everted (inside out) labia and copious flows of urine and mucus. Upon reaching peak oestrous, mares spread-out their legs, lift their tails and open their mouths when in the presence of a male. Males assess the female's reproductive state with a curled lip and bared teeth ([[flehmen response]]) and the female will solicit mating by backing in. Gestation is typically around a year. A few days to a month later, mares can return to oestrus.<ref name="Estes 1991" /> In harem-holding species, oestrus in a female becomes less noticeable to outside males as she gets older, hence competition for older females is virtually nonexistent.<ref name="Grub 1981" />
[[File:Cape Mountain Zebras (Equus zebra) mare and foal suckling ... (31281408687).jpg|thumb|right|alt=Mountain zebra suckling a foal |Mountain zebra suckling a foal]]
Usually, a single foal is born, which is capable of running within an hour of birth.<ref name="MacDonald" /> A newborn zebra will follow anything that moves, so new mothers prevent other mares from approaching their foals as they become more familiar with the mother's striping pattern, smell and voice.<ref name="Churcher 1993" /> At a few weeks old, foals begin to graze, but may continue to nurse for eight to thirteen months.<ref name="MacDonald" /> Living in an arid environment, Grévy's zebras have longer nursing intervals and young only begin to drink water three months after birth.<ref name="Becker 1990">{{cite journal |last1=Becker|first1= C. D.|last2= Ginsberg|first2= J. R.  |year=1990 |title=Mother-infant behaviour of wild Grevy's zebra |journal=Animal Behaviour |volume=40 |issue=6 |pages=1111–1118 |doi=10.1016/S0003-3472(05)80177-0|s2cid= 54252836}}</ref>

In plains and mountain zebras, foals are cared for mostly by their mothers, but if threatened by pack-hunting hyenas and dogs, the entire group works together to protect all the young. The group forms a protective front with the foals in the centre, and the stallion will rush at predators that come too close.<ref name="Estes 1991" /> In Grévy's zebras, young stay in "[[Crèche (zoology)|kindergartens]]" when their mothers leave for water. These groups are tended to by the territorial male.<ref name="Becker 1990" /> A stallion may look after a foal in his territory to ensure that the mother stays, though it may not be his.<ref name="Rubenstein 1986"/> By contrast, plains zebra stallions are generally intolerant of foals that are not theirs and may practice [[Infanticide (zoology)|infanticide]] and [[feticide]] via violence to the pregnant mare.<ref>{{cite journal|last1=Pluháček|first1=J|last2=Bartos|first2=L|year=2005|title=Further evidence for male infanticide and feticide in captive plains zebra, ''Equus burchelli''|journal=Folia Zoologica-Praha|volume=54|issue=3|pages=258–262|url=https://www.ivb.cz/wp-content/uploads/54_258-262.pdf}}</ref>