equus (hippotigris) quagga (burchelli)

EXCERPT

Diersoort

Wilde soorten:

Equus (Hippotigris) quagga (burchelli) - Steppezebra

Equus (Equus) ferus przewalskii - Przewalskipaard

Equus (Dolichohippus) grevyi - Grévyzebra

Equus (Asinus) hemionus hemionus- Onager

Equus (Asinus) hemionus khur - (= ondersoort Equus hemionus) Indische wilde ezel

Equus (Asinus) kiang – Kiang (3 ondersoorten)

Equus (Hippotigris) zebra – Bergzebra (2 ondersoorten)

Gedomesticeerde soorten (vallen buiten de beoordeling; veel specifieke literatuur

daarom buiten het excerpt gelaten):

Equus (Asinus) asinus asinus - Gedomesticeerde ezel

Equus (Equus) caballus - Paard

Equus (Equus) caballus x Equus (Asinus) asinus asinus - Muildier (hybride, geen (onder)soort)

Equus (Asinus) asinus asinus x Equus (Equus) caballus – Muilezel (hybride, geen (onder)soort)

Genus Equus BRONNEN

Volwassen grootte

en/of gewicht

E. quagga (burchelli)

Schofthoogte: 110 – 145 cm

Gewicht: 175 – 385 kg

“Males are slightly larger than females and usually have thicker necks as

well. This sexual dimorphism is not profound, however.” (Colvin & Nihranz

2009)

E. f. przewalskii



E. grevyi



“Equus zebra is the largest living zebra” (Churcher 1993, p.1)

E. hemionus ssp.



E. kiang

Schofthoogte: 142 cm


“Equus kiang is the largest of the wild asses” (St-Louis & Côté 2009, p.2)

E. zebra



“The subspecies E. z. zebra, of South Africa, is usually smaller then E. z.

hartmannae, of Angola and Namibia.” (Nowak & Walker 1991, p.1314)

Animal Diversity

Web; WIK-EN;

WIK-NL;

McGreevy 2004

p. 15-16;

National

Geographic

Animals

[Online]; Nowak

& Walker 1991;

Afweer- en

aanvalsmiddelen

“All equids are relatively swift, alert runners and generally flee from

danger rather than fight. However, among their own kind or in attempting

defense, they kick with the hind feet, strike with the forefeet and

sometimes bite.” (Ballenger & Myers 2001)

“In fighting among themselves or in attempting a defense, they kick with

the hind feet, occasionally strike with the forefeet, and sometimes bite.

Their teeth, though not adapted for lacerating or tearing (except in male

zebras, which have pointed canines), can give quite hard pinches.” (Nowak

& Walker 1991, p.1306)

Carnivoor/omnivoor/he

rbivoor

Herbivoor, grazers.

“Het zijn herbivoren die vooral gras eten. Wanneer het nodig is, eten ze

ook ander voedsel zoals bladeren, fruit en boomschors.” (Wikipedia

[Nederlands]: Equus (geslacht))

Verspreidingsgebied E. quagga (burchelli) – southeastern Africa, with the highest population

densities in the Serengeti-Mara plains of Kenya and Tanzania. (Colvin &

Nihranz 2009); Burchell’s zebras roam the open savannas of southeastern

Africa. They prefer open grasslands, open woodlands, and open scrub

environments. Occasionally, they may also inhabit taller grasslands,

heavier woodland areas, and even hilly country and mountainous regions

up to 4,400 meters in elevation. However, they avoid dense forests,

deserts, and wetland areas (Colvin & Nihranz 2009). “Plains Zebra live in

all habitats in Africa from sea level to 4,300 m on Mount Kenya, with the

exception of rain forests, deserts, dune forests, and Cape Sclerophyllous

vegetation” (Hack & Lorenzen 2008)

E. f. przewalskii – “Altai Mountains of Mongolia” (Luu 2002); “… grassy

deserts and plains in Western Mongolia, but it has been reported to have

lived at elevations of up to eight thousand feet (Volf 1990).” (Luu 2002).

“Przewalski’s horse occured in both plains and hills up to an elevation of

2.500 meters and apparantly made seasonal migritions in association with

rainfall and vegetation growth.” (Nowak & Walker 1991, p.1311)

E. grevyi – “Grévy zebras occupy arid areas of southern Ethiopia and

northern Kenya” (Churcher 1993, p.1). “Grévy zebras live in semi-deserts

with low vegetation (1 m) or scattered low (3-4 m) Acacia bush with herbs

and grasses. They prefer stony plains or hills, short grassy flats, or broken

country (Churcher, 1982; Willoughby, 1974).” (Churcher 1993, p.4). “(…)

semi-arid grasslands, filling a niche distinct from that of other members of

the genus Equus that live within the same geographical range, such as

wild asses (which prefer arid habitats) and plains zebras (which are more

dependent on water than Grevy's zebras). (Hollingshead 2010)

E. hemionus – “Currently, at least one subspecies has been found in

Russia, China, Iran, and India. However, the major population (over half

the total number) of E. hemionus is found in southern Mongolia. (Feh et

al, 2001)” (Reuter 2002). “It primarily grazes and rests on highland or

lowland desert, semidesert or steppe.” (Reuter 2002). “It currently

occupies four limited areas in southern Mongolia (subspecies E. h.

hemionus), Turkmenia (E. h. kulan), northern Iran (E. h. onager), and

southwestern India (E. h. khur).” (Nowak & Walker 1991, p.1309)

E. h. khur – The Indian wild ass's range once extended from western

India, southern Pakistan, Afghanistan, and south-eastern Iran; Saline

deserts (rann), arid grasslands and shrublands are its preferred

environments. (Wikipedia (Engels))

E. kiang – “Equus kiang is endemic to the Tibetan Plateau of Central Asia

at elevations of 2,700–5,300 m (Schaller 1998; Shah 2002; Fig. 3).” (St-

Louis & Côté 2009, p.3). “Three major vegetation formations cover the

distributional range of Equus kiang: alpine meadow, alpine steppe, and

desert steppe (Miller and Schaller 1996; Schaller 1998).” (St-Louis & Côté

Animal Diversity

Web;

Mammalian

Species; IUCN;

Nowak & Walker

1991;

2009, p.5)

E. zebra – Cape mountain zebras are found only in South Africa.

Hartmann’s mountain zebras range from South West Africa into extreme

southwest Angola. Their distribution is highly discontinuous. (Walker

2005); Mountain zebras inhabit slopes and plateaus in mountainous areas

of South Africa and Namibia (South West Africa). The habitat in South

Africa provides regular precipitation and a fairly constant food-supply year

round. Hartmann’s mountain zebras differ from Cape mountain zebras in

that they occupy an arid region in a mountainous transition zone on the

edge of the Namib Desert. (Walker 2005)

Domesticated species:

E. a. asinus – “Within its natural range, E. asinus inhabits arid, stony

regions dotted with hillocks, which are used as observation posts (Groves

1974, 1986).” (Grinder et al. 2006)

E. f. caballus – “Free-living horses usually are associated with grasslands,

steppes, and semiarid shrub country, though some early reports indicate

the former presence of true wild E. caballus in certain European forests.”

(Nowak & Walker 1991, p.1311)

Hybriden (muilezel en muildier) komen alleen voor in gevangenschap

(bewust gefokt) en worden bij voorkeur ook gehouden in open

landschappen.

Levensduur “The potential lifespan is 25-45 years.” (Nowak & Walker 1991, p.1306;

about Equus in general)

E. quagga (burchelli)

Wild: 20 jaar

Gevangenschap: 35 - 40 jaar

E. f. przewalskii

Wild: 20 – 25 jaar

Gevangenschap: 20 jaar

E. grevyi

Wild: 12 – 18 jaar

Gevangenschap: 22 – 30 jaar

E. hemionus

Wild: 4 – 14 jaar

Gevangenschap: tot 35 jaar.

E. h. khur

Wild: N/A

Gevangenschap: 32 jaar (oudst gerapporteerd)

E. kiang

Wild: 20 jaar

Gevangenschap: 30 jaar

E. zebra

Wild: 20 jaar of ouder

Gevangenschap: tot 30 jaar

Animal Diversity

Web; Wikipedia

(Engels,

Nederlands);

Haltenorth &

Diller 1988;

Nowak & Walker

1991;

Mammalian

Species; Feh et

al. 2001;

National

Geographic

Animals

[Online]; Kumar

et al. 2009;

IUCN-status E. quagga (burchelli) – Least Concern

E. f. przewalskii – Endangered D

IUCN ver 3.1

(Online,

N.B. Toegevoegde

codering na

“Endangered” en

“Vulnerable” hebben

betrekking op redenen

waarom ze op deze lijst

staan; voor uitleg zie:

http://www.iucnredlist.

org/static/categories_cr

iteria_3_1

E. grevyi – Endangered A2ac; C2a(i)

E. hemionus – Endangered A2abc+3bd

E. h. khur – Endangered A2abc+3bd

E. kiang – Least Concern

E. zebra – Vulnerable C1

accessed

December 10,

2014), Wikipedia

(Engels)

CITES-status E. quagga (burchelli) – Not Listed

E. f. przewalskii – App. I

E. grevyi – App. I

E. hemionus – Split-listed. Subspecies E. h. hemionus and E. h. khur are

listed on App. I, other subspecies are listed on App. II.

E. kiang – App. II

E. zebra – Split-listed. Subspecies E. z. zebra is listed on App. I, while

subspecies E. z. hartmannae is listed on App. II

Checklist of

CITES Species

(Online,

accessed

December 10,

2014)

Aantal gebruikte bronnen:

Is voldoende bekend [over de houderij] om oordeel te geven? Ja

Samenvatting van soorteigen gedrag (incl. Klinische indicatoren van gevolgen van houderij voor

dierenwelzijn/diergezondheid)

Hieronder staat per ‘Aspect van soorteigen gedrag’ aangegeven wat de kerneigenschappen (statements)

van de diersoort zijn in de natuur (linker kolom) en in de houderij (rechter kolom). Vervolgens/daaronder

wordt aangegeven welke oorzaak-gevolg relaties gespecificeerd kunnen worden t.b.v. de beoordeling van

het risico op gezondheids- en welzijnsproblemen in de houderij. Met kleurarcering wordt aangegeven welke

statements de oorzaak-gevolgrelaties onderbouwen.

Leefruimte (bewegen en een specifieke leefomgeving)

In de natuur:

Migratory: E. h. kiang (Fryxell & Sinclair 1988)

“Milieu: grasvlakten, savannen met verspreide boomgroei,

heuvellandschap” (Whitfield 1985; Equus burchelli)

“The plains zebra occupies grassland and savanna woodland,

from sea-level to elevations of up to 4,300 metres on Mount

Kenya (1) (3). A highly adaptable species, the plains zebra is

capable of surviving in areas with coarse vegetation with little

nutritional value, but needs to have access to water for daily

drinking.” (ARKIVE)

“Burchell s zebra occurs in a variety of habitats, including

savannah, light woodland, open scrub, and grassland. It

sometimes is found in broken, hilly country and on mountain

slopes up to 4,400 meters.” (Nowak, 1999)

“They travel up to 13 kilometers daily between resting areas

in tall grasslands and grazing areas in short grassland areas.

However, the necessity for fresh water and grazing lead to

movements of 100 to 150 km seasonally.” (Colvin & Nihranz

In de houderij:

“Stereotypic behaviors are not recognized in freeliving

feral horses and are not purely a product of

domestication since they are also reported in captive

examples of wild equids such as the onager mountain

zebra146 and Przewalski horses.147 In the horse, these

behaviors have therefore been linked to a number of

management practices.” (McGreevy 2004, p.23)

“When providing accommodation, one must consider

the natural habitat of the various species. Zebras and

asses require simple shelters in the winter, depending

on the local environmental conditions these will in

individual cases have to provide heating. Flooring must

be nonslippery because these species are easily startled.

To decrease the prevalence and severity of overgrown

hoofs, the areas around feeders may be surfaces with

coarse granite stones and roughened concrete. In the

summer months shade should be available to all

members of a herd.” (Walzer, 2003)

http://www.iucnredlist.org/static/categories_criteria_3_1



2009; Equus burchelli)

“The home range of Burchell’s zebras is dependent on

necessary resources, mostly the availability of fresh water

and grazing areas. Herds are non-territorial and occupy

overlapping home ranges. Typically, groups occupy ranges of

300 to 400 sq km in the wet season and 400 to 600 sq km in

the dry season.” (Colvin & Nihranz 2009; E. burchelli)

“Home ranges vary from a minimum of 30 km2 in a small

rich ecosystem like Ngorongoro Crater, up to 600 km2 in the

migratory Serengeti population (6)” (Estes 1991; Equus

burchelli)

The sedentary population in Ngorongoro, Tanzania, has a

population density of 19.2/km2 (Turner and Watson, 1964)

and group home ranges of 80 to 250 km2 (Klingel, 1969a).

Densities range from 0.7 to 2.2/km2 and home ranges from

49 to 566 km2 (mean 164 km2 ) in the Kruger National Park

where both sedentary and migratory populations occur

(Smuts, 1975b). The numbers of the largest migratory

population, from the Serengeti Plain, Tanzania, are estimated

to be between 57,000 and 280,000 (Grzimek and Grzimek,

1960a, 1960b; Hendrichs, 1970; Kruuk, 1972; Schaller,

1972; Talbot and Stewart, 1964)…

…home range 300 to 400 km2 in the rainy season and 400 to

600 km2 in the dry season (Klingel, 1969b). “(Grub, 1981)

“Zebra populations can be sedentary, with high population

densities and small ranges, or migratory, with relatively low

densities and different wet and dry season home ranges of

great extent.” (Grubb 1981; about Equus burchelli)

“The differing habitats and the need to obtain forage and

water have resulted in both sedentary and migratory

populations.” (Nowak & Walker 1991, p.1316; E. burchelli)

“Grevy’s zebra stallions in Kenya were found to have

territories with an average size of 5.75 sq km and a range of

2.70-10.50 sq km.” (Nowak & Walker 1991, p.1313)

“Live in mesic grasslands and are capable of migrating long

distances. Their ability to utilize coarser vegetation of poorer

quality means that they can thrive and survive in a range of

habitats.” (E. burchelli; Grzimek, 2004)

“Home ranges of dominant males overlapped little,

suggesting that in this population males are territorial.”

(Saltz et al. 2000; E. hemionus)

“According to these observations, the social organization of

E. africanus and E. hemionus is of the same type as that of

Grevy’s zebra (Equus grevyi) (Klingel, 1974a.), being

characterized by the very large territories and the conditional

defence by the territorial individual.” (Klingel 1998, p.108)

“Asiatic wild asses (‘kulans’—Equus hemionus spp.) and

Przewalski horses (Equus przewalski) inhabited steppe, semi-

desert and deserts of temperate Eurasia. Unlike kulans,

“Tetanus caused by Clostridium tetani is a significant

risk factor for all equids. Clinical symproms are similar

to those in the domestic horse: muscle rigidity, tetanic

cramping, anxiety, anorexia and prolapsed third eyelid…

Preventive vaccination with a toxoid vaccine with yearly

boosters is suggested. In surgical procedures and wound

management the use of tetanus antitoxin in

nonvaccinated individuals is recommended." (Walzer,

2003)

“Hoof problems have been reported in all species of

nondomestic equids. Several authors have postulated

that hoof trimming is absolutely necessary, especially in

the Przewalski’s horse. However with providing coarse

granite stones the hoofs will crack and eventually break

along a predetermined line, making regular trimming

unnecessary in many instances. The extremely hard and

narrow hoof of the mountain zebras need regular

trimming when they are kept on pastures, thus

mandating a change in enclosure surfacing.” (Walzer,

2003)

Three herds of Plains zebra and eland in multi-species

exhibits in different Danish zoos were observed during

the spring and summer of 1989. The exhibits varied

greatly in size and design: Copenhagen Zoo-0.5 ha,

grassless; Loveparken Zoo Givskud-5 ha, short grass;

Knuthenborg Safaripark-18 ha, long grass. A negative

correlation was seen between exhibit size and amount of

interspecific interactions. When designing multi-species

exhibits, ample resources (food, water, etc.) and space

at feeding sites should be provided. (Andersen, 1992)

“Their natural habitat must be considered when

accommodations are provided for them in captivity. In

the summer shade should be available for all species, as

should shelter in the wintertime. All zebras and most

asses require heated quarters in colder climates.

Exercise is important for their well-being, and inasmuch

as all tend to be easily startled, care must be taken to

provide proper footing. Most wild equids are not adapt

on slippery surface” (Nelson, 1986)

“Animals in all collections have encountered hoof

problems (see Chapter 35). The extent of the problem

and the frequency of trimming can be lessened by

providing the unit with a coarse, decomposed granite

soil or roughened concrete platforms around feeders and

periphery of the enclosure. Przewalski’s horses and

mountain zebras will still require attention.“ (Nelson,

1986)

“Periodic hoof trimming is needed in some individuals

and under certain management conditions. Some

species of equids (e.g. mountain zebras and Przewalski’s

horeses) seem to require more frequent hoof trimming

compared with others. Przewalski’s horses, for example

may require trimming every 6 to 9 months.37” (Janssen

horses preferred more mosaic ecosystems, demonstrated

less local migratory activity and smaller home ranges, being

fixed to a greater extent to watering places (Grum-

Grizimailo, 1892, 1896; Geptner, et al., 1961).” (Bahloul et

al. 2001, p.309-310)

“Territories are very large, measuring on the order of 20 km2

in E. africanus and probably even more in E. hemionus.”

(Klingel 1998)

“For their night rest all three species choose flat, open

spaces, where it is easy to notice an approaching danger

from a distance and where it is possible to flee without

encountering natural obstacles. Extensive plains or even vast

valleys are most attractive for these species. They definitely

avoid places with a greatly dissected relief, but under some

conditions must use them. Thus, in winter they hide from

cold winds or snow storms behind sand dunes or in ravines,

while in the summer they find here cooler places for resting

during the heat of the day.” (Formozov 1966)

“E. kiang may use flat and open areas as escape terrain

where it can flee easily, a common antipredator strategy

among wild equids (Harris and Miller 1995; Schaller 1998).”

(St-Louis & Côté 2009, p.6)

“Generally the present-day equids exploit the semiarid

grasslands, where they play a key role in the functioning of

natural grazing systems... The equidae use a great range of

altitudes within their present day distribution: 60 m (Danakil:

Somali wild ass) and 5000 m (Tibet: Kiang).14” (Walzer,

2003)

Both species of antelopes, and the kulans, are permanently

roaming animals, but regular seasonal migrations are not

typical of all populations, since their nomadic habits vary

considerably depending upon the crop of forage grass, the

conditions of watering places and character of snow cover in

winter (Formozov 1966)

Kulans are typically more migratory-active and better

adapted to desert conditions (Geptner, 1961); that is why

they can regularly do without water, or use water with very

high salinity (Rashek, 1966). (Bahloul et al 2001)

& Allen, 2014)

“Hoof problems commonly occur especially in large

collections of nondomestic equids. Routine hoof

trimming may be necessary in some species and under

some management conditions. The mountain zebras, for

example, seem to require more frequent hoof trims

compared with other species, especially when housed on

soft or sandy soil.” (Janssen & Allen, 2014)

“Enclosure effects:

Skiff (1982) studied the effects of enclosure size on the

behavior of 2 herds of Przewalski's horses. The number

of activities was higher in the 0.05 ha enclosure than in

the 3.5 ha enclosure. When in the small enclosure,

horses did more milling and fence-pacing, had higher

rates of aggression, and spent more time mutually

grooming than when on pasture. Coprophagy and

woodchewing was seen in the small enclosure but never

on pasture. Przewalski's horses isolated in small

enclosures may also begin cribbing, a behavior difficult

to extinguish even when the horse is moved to a

pasture (Bouman-Heinsdijk, 1982; Boyd, 1986).

In the present study, there was no significant effect of

enclosure size on aggressiveness or on the amount of

time spent in mutual grooming or in locomotion

(Kruskal-Wallis ANOVA, P>0.05). The main effects of

enclosure size were that horses in enclosures < 0.4 ha

in size tended to exhibit more behavioral states per hour

and spent significantly less time standing resting than

animals on pasture (Boyd, 1988b ). Horses in small

enclosures were in metropolitan areas, whereas those in

large enclosures were not. It is possible that the noises

in zoos or of the cities in which they are located, disrupt

the diurnal resting behavior of the horses” (Boyd, 1991)

Weaving is one of the most common stereotypical

behaviors of horses. Depending on breed, management

system and use, between 1 % and 9% of all horses

weave. It generally occurs in stalled horses..…In poorly

vegetated corrals, but also in pasture, it can be

observed in isolated horses or those suffering from

sensory deprivation. (Zeitler-Feicht 2004).

Oorzaak-gevolg risico’s:

Onvoldoende bewegingsruimte > Locomotiestereotypieën (o.a. weven), coprofagie and woodchewing

Onvoldoende harde/ruwe ondergrond of gladde ondergrond > Hoefproblemen (overgroeide hoeven)>common en

regular trimming needed (Janssen en Allen 2014) of Trauma a.g.v. uitglijden (mn wanneer dieren schrikken)

Flight animal + external stimuli in captivity (such as noises) > disrupt diurnal resting behaviour [Noot: niet in

argumentenkaart opgenomen vanwege te weinig ruimte (afgesproken max van 16 arg op ArgKaart)]

Sociaal gedrag (sociale of biosociale behoeften) [inclusief agressie]

In de natuur:

“Equids are polygynous herd animals that generally live in

extended family groups occupying large territories in open

country (grasslands, semi-arid areas, deserts, and

mountains).” (Ballenger & Myers 2001)

“In the equids of the arid habitats, some males defend

territories, preferably near water sources, which confers

mating rights over females when they enter the territory,

Asses and Grevy’s zebra form loose groups and only long-

term relationship occurs between mother and offspring for

a period of 2 years. Asses in central Asia may aggregate

to loose groups in excess of several hundred individuals.

The other equid species form permanent family groups,

so-called harem groups. In contrast to the asses, in this

system, lifelong relationships between adult individuals

are formed. Young males form bachelor groups.”

“Two distinct types of social organizations have been

described in wild equids (Klingel 1975, 1977). In the 1st

type, permanent groups contain a few males and several

females and young, and males are not territorial. This

type of social organization occurs in equids from

temperate ecosystems such as E. burchellii (Burchell’s

zebra), E. zebra (mountain zebra), and E. caballus

przewalskii (Przewalski’s horse—Moehlman 1998). In the

2nd type, the only permanent associations occur between

mother and foal. Only temporary groups are formed,

comprising several females and their young or only males

(bachelor groups); adult males are often territorial. This

type of social organization has been documented in equids

adapted to arid environments, such as E. grevyi, E. asinus

africanus (African wild ass—Grinder et al. 2006), and E.

hemionus (Klingel 1977; Moehlman 1988; Rubenstein

1989). (…) Equus kiang seems to display the 2nd type of

social organization (Groves and Willoughby 1981; Schaller

1998).” (St-Louis & Côté 2009, p.7)

“In the wild, Przewalski's horses live in small, permanent

family groups consisting of one adult stallion, one to three

mares, and their common offspring. Offspring stay in the

family group until they are no longer dependent, usually

at two or three years old. Bachelor stallions, and

sometimes old stallions, join bachelor groups. Family

groups can join together to form a herd that moves

together.” (Wikipedia [Engels]: Przewalski’s horse)

In het droge seizoen voegen meerdere kudden zich bij

elkaar tot grotere kudden van soms meer dan duizend

dieren en trekken ze naar vochtigere gebieden. Vaak

sluiten ze aan bij kudden andere hoefdieren, als blauwe

gnoe en Thomsongazelle.” (Wikipedia [Nederlands]:

Steppezebra)

“Zebra social groups are non-territorial and have

In de houderij:

“The various social systems must be considered to manage

these species. Removing individual animals from a harem

group within a program must be regarded especially

critically.” (Walzer, 2003) N.B. Is indicatie voor

gespecialiseerde kennis nodig per soort (opmerking CK)

“Once mature, equids may become dangerous. Intraspecies

and interspecies aggression and trauma is common.

Numerous traumatic injuries and deaths have occurred

when equid males where kept in adjacent enclosures. These

injuries were sustained from the fencing material or from

the fence being broached, thereby causing injuries

subsequent to the contact" (”walzer, 2003)

“Zebras are notorious for killing the young of other species

in mixed exhibits.” (Kleiman et al. 1996, p.42)

“The zebras were the more challenging species as they tend

to be easily agitated; this was particularly evident in the

three individuals being introduced. Zebras may work

themselves into a veritable frenzy once this state is

triggered, and they are often injured in the process.”

(Kleiman et al. 1996, p.207; over opzetten van mixed-

species habitats en introducties van verschillende soorten;

in dit geval zebra’s en neushoorns)

“The establishment of defined social status within any

group of equids promotes stability within the band. A stable

social hierarchy within the band decreases the accumulative

amount of injury by allowing threats of kicks and, perhaps

more importantly, bites to replace the aggressive responses

themselves.6” (McGreevy 2004, p.124-125)

“There is some tendency for males to groom certain

females preferentially. Conversely, males may selectively

attack particular individual females, at least in captivity.”

(Grubb 1981)

“In de dierentuin kunnen halfezels zonder grote

moeilijkheden worden gefokt. De dieren wennen er snel aan

hun nieuwe omgeving; de jongen zijn tamelijk goed

bestand tegen ons wisselvallig klimaat en wennen zonder

moeite aan ander voer. Alleen de hengst veroorzaakt door

zijn aggressiviteit vaak moeilijkheden in een kudde

fokdieren. Hij jaagt de merries op en bijt ze in de poten en

vooral in de keel.” (Grzimek 1973 [deel xii, p. 656, kol. 2])

“Ezels zijn zeer sociaal en dus niet geschikt om alleen

gehouden te worden. Gedomesticeerde ezels kunnen wel in

groepen van allerlei samenstellingen en groottes

samenleven omdat ze evolutionair gezien ook bekend zijn

met een erg losse sociale structuur. Dit in tegenstelling tot

paarden die vaak leven in groepen van een vastere

samenstelling. In een gedomesticeerde omgeving kan het

daarom voorkomen dat er in de groep paarden vaker

overlapping home ranges but females are antagonistic

towards those in other groups.” (Grubb 1981; E.

burchelli)

“Family groups are stable. If the group stallion dies or

disappears, another male replaces him.” (Grubb 1981)

Stable, female groups were the mainstay of group

sociality with male mean residency at 31.6 months being

variable in length or even absent. (Simpson & Nicol,

2012)

Zebra populations can be sedentary, with high population

densities and small ranges, or migratory, with relatively

low densities and different wet and dry season home

ranges of great extent. (Grubb, 1981)

“Bonds of the foal with the mother are lost when she has

a new foal and as opportunities for play-fights with other

males are greater in stallion groups. Family stallions

nevertheless do not show antagonism to their sons.”

(Grubb 1981; E. burchelli)

A prerequisite for the cost-effective viability of female

defense polygyny (stable family/harem groups) is a

spatial and temporal patterning of resource availability

such that it is possible for females to feed in close

proximity. (Grzimek, 2004)

From the female point of view, abundant food allows

closer spacing with other females and gregariousness

enhances predator detection. In addition, the presence of

a dominant male precludes harassment by other males in

the population (Grzimek, 2004)

Several harems come together to form large herds during

their migratory journeys. (Colvin 2009)

Social interactions across all categories of zebras were

free from aggression (Simpson & Nicol, 2012)

“There are two kinds of social groups: family groups

(stallion, mares and their foals) and stallion groups

(males only, including solitary males). There is no social

organization above the level of groups” (Grubb 1981; E.

burchelli)

“With their first estrus, young females are abducted by

males of other family-groups or stallion groups, the latter

thus founding new families. In her first estrus, the female

stands with legs apart and tail lifted in an estrous posture

which attracts up to 18 males. The family stallion is rarely

able to keep her.” (Grubb 1981; E. burchelli)

“Burchell s zebras are social, living in permanent family

groups composed of one male stallion, 1 to 6 females,

and their young. The strong bonds between females are

the central relationships within harems.” (Colvin &

Nihranz 2009; E. burchelli)

sprake is van agressie indien de samenstelling van de groep

aan verandering onderhevig is dan bij een groep ezels.” (de

Graaf-Roelfsema 2009, p.66)

“Wherever possible, horses should be kept in social groups.

The horse responds poorly to isolation and is likely to show

physiological and behavioral distress responses including

stereotypies if deprived of contact with conspecifics.81”

(McGreevy 2004, p.141-142)

“Indien een sterk aan elkaar gehecht paar ezels van elkaar

wordt gescheiden, bestaat er een grote kans dat beide

vervolgens weigeren te eten en apatisch gedrag gaan

vertonen. (…) Om het risico op stress en hyperlipemie te

beperken is het erg belangrijk dat bijvoorbeeld gedurende

een kliniekopname van één van de ezels deze samen met

zijn levenspartner gehuisvest kan worden en als één van

beide sterft, de ander nadien voldoende tijd met het

lichaam krijgt om de situatie te accepteren.” (de Graaf-

Roelfsema 2009, p.67)

“The prominent role of the stallion in regulating group size

could mean that the competition experienced in the safari

park is harder for stallions than it is for mares. Young

females not expelled from their natal harem go through a

"wandering phase", visiting other groups. After that they

exert female choice when they eventually join a group or an

individual.” (Schilder 1992; E. burchelli)

“Trauma, especially bite wounds, are often seen and are

difficult to handle. If the animal must be isolated and

treated, reintroduction and acceptance of the individual can

rarely be achieved. Trauma resulting in fracture in any

adult is usually fatal, as adult wild equids are notoriously

uncooperative patients.” (Nelson, 1986)

“Some interactions and introductions among equids may be

quite violent and aggressive, resulting in injuries from

kicking or from bite wounds on the neck or tail area”

(Janssen & Allen, 2014)

“The dominance hierarchies of Przewalski's horses appear

to be remarkably stable over time. In the 4 years that the

Topeka herd was observed, only one change in rank

occurred.

Przewalski's horses have been reported to direct most of

their aggression toward horses just below them in rank

(Hutson, 1975; Mackler and Dolan, 1980 ). Subordinate

mares are the most aggressive toward new herd members

(Mackler and Dolan, 1980). In this study, 55% of the mares

directed the majority of their aggression toward the horse

just below them in rank (or 63%, if only the mares who

were not at the bottom of their respective hierarchies are

considered). For the 2 cases in which new mares were

introduced to the harem during the course of the study, the

lowest ranking adult female showed the most aggression

toward the newcomers. Foals were always the lowest

ranking animals in the herd but were protected when near

The pattern of stripes is unique for each individual and

serves as a means of identification. Social interactions are

based on this particular identification. Studies show that

individuals with more stripes receive additional social

attention, such as grooming. With these social benefits, it

would have been more advantageous to have more

stripes. (Colvin & Nihranz 2009)

“They form small breeding herds that consist of one adult

stallion and 1 to 5 mares with young. Breeding herds

remain stable over many years and mares usually remain

in a herd for life. (Joubert, 1972a; Nowak, 1999;

Penzhorn, 1984; Penzhorn, 1988)” (Walker 2005; E.

zebra)

“Social organization and reproductive strategies in equids

are a complex interaction of individual feeding

requirements, access to females, and defense against

predation. Individuals will form groups when benefits

exceed costs in terms of feeding, predation, disease, and

reproduction.” (Grzimek et al. 2004)

“A dominance hierarchy is present in harems and employs

a rank order of the dominant male, followed by the

mares, and then the foals. The dominant female

preserves the rank order by leading the group in single

file movements, in which mares line up according to age

correlated rank. Foal rank depends on mother’s rank, they

stand one place directly behind her in the line and the

newest mare of a harem takes the lowest social rank and

is placed at the end. The stallion pulls up the rear of the

line taking a defensive role in case of predator attack.”

(Colvin & Nihranz 2009; E. burchellii)

“Young males in bachelor groups play and engage in mock

fights, preparing for future fights when they begin

searching for available mates and starting their own

harems. When they reach 4 years old, males are finally

prepared to fight for mates and establish a harem.”

(Colvin & Nihranz 2009; E. burchellii)

“Als de hengst 16-18 jaar oud is, wordt zijn plaats op

vreedzame wijze ingenomen door een jongere hengst van

6-8 jaar oud, en leidt hij verder een solitair bestaan”

(Whitfield 1985; E. burchelli)

“Harems are organized into a dominance hierarchy.

Females of higher rank have been found to produce more

offspring and to have shorter interbirth intervals. Stallions

show mating preference towards these high ranking

females. Immediately after birth, foals take a position in

the dominance hierarchy at a position below their

mothers.” (Colvin & Nihranz 2009; E. burchellii)

“Males fight fiercely to obtain or retain mares, circling one

another and attempting to bite or kick the opponent.

Dominant males also are highly protective of their units

and take up a defensive position to the rear while the

other animals flee. Intergroup relations vary, with the

their mother and so initiated and received little aggression.

Age was positively correlated with dominance order for the

adults in the 10 herds studied and significantly correlated

with rank in 4 of the 6 herds large enough to permit

statistical testing. Following age in importance was body

size. Aggressiveness was positively correlated with

dominance in all but 1 herd. Horses ranking high in the

hierarchy initiated more aggressive acts per hour. Factors

of lesser importance were length of residence in the herd

and maternal rank.

Mares in a harem form close, stable associations that

persist even in the absence of stallions (Kaseda, 1981;

Keiper, 1985; L.E. Boyd, personal observations of 3 herds

of Przewalski's mares without a stallion present, 1986,

1988 ). It is not unusual for mares in the same harem to be

relatives (mother and daughters, full or half sisters), which

probably contributes to the close ties between harem

females. Closely related mares preferred one another as

mutual grooming partners. As mares form these long-term

bonds, it seems logical that the present practice of rotating

stallions from harem to harem to reduce inbreeding more

closely approximates the natural social processes than

would a program that traded particular mares

As of 1980, 75 zoos worldwide maintained Przewalski's

horses in a total of 97 enclosures (Bouman, 1982 ). Thirty-

two enclosures housed males only. The males in 26 of these

enclosures were solitary, and therefore, likely to pace.

Horses are gregarious; in the wild, stallions which do not

possess harems are rarely solitary but form bachelor

groups instead. In captivity, this process can be mimicked.

Males not currently being used for breeding can be grouped

to form bachelor herds. This reduces the number of

enclosures needed and makes the stallions less likely to

pace. Serious aggression is not a problem with juvenile

males; forming herds containing older experienced males

requires more care, however. By setting up herds in areas

large enough (0.4-0.8 ha) to allow subordinates to flee and

take cover, Tilson et al. (1988) were able to form bachelor

groups containing males ranging in age from 1 to 9 years.

Wild stallions seldom have harems of more than 5 or 6

females. When the harem of the Przewalski's stallion "Basil"

grew to 18 members, he became apathetic about breeding

(Knowles, 1980). Isolating him and reducing the harem size

prior to his return restored his libido. The stallion "Bars"

became aggressive toward his mares when the harem size

reached 13 animals (Keverling Buisman and van Weeren,

1982 ).” (Boyd, 1991)

“It has been suggested that the restrictions associated with

human management may precipitate higher rates of

aggression than may be seen in the free-ranging state.4,7

This should be borne in mind when horse-holding facilities

are designed: e.g. paddocks should have rounded corners

to prevent subordinate animals becoming trapped and

adult mares being antagonistic, the dominant stallions

greeting one another in a ritualized ceremony, and the

young sometimes playing together (Klingel 1969).”

(Nowak & Walker 1991, p.1316; E. burchellii)

“Equus zebra is a social species with populations

consisting of breeding herds and bachelor groups.”

(Walker 2005; E. zebra)

“Other methods of plains zebra communication include

facial expressions, body movements, and sounds, such as

a braying bark.” (ARKIVE)

E. burchelli communicates by a variety of gestures, facial

expressions, and sounds. An explosive braying bark

seems to identify stallions and maintain group cohesion.

(Nowak & Walker, 1991)

Stallions of different groups greet each other with their

ears up. When they sense threat, especially in the form of

combat, they will put their ears down. Greetings are also

achieved through nose sniffing, rubbing, and genital

smelling. (Colvin 2009)

Six calls and two facial expressions are used in

communication between individuals. Three of the calls are

used as predator alert or threat calls, one is used to

communicate injury, another is used in distress, and the

last one is used in contact between individuals. (Colvin

2009)

Relationships between harems are relatively cordial and

males have a ritual greeting. When they meet, males

keep their

ears standing up and sniff each other’s bodies, especially

their necks, nostrils, flanks, and tails. Females from other

harems tend to be antagonistic towards each other.

(Colvin 2009)

“The unusal social behaviour of E. grevyi has been

reviewed by Kingdon (1979) and Klingel (1974). Although

thousands of animals formely congregated in response to

temporary environmental conditions, there are no lasting

bonds between adults such as those of E. burchelli and E.

zebra. The only strong association is that of a female and

her offspring of one or more years.” (Nowak & Walker

1991, p.1313)

“The observed three-level society in this khulan

population reveals a complexity of social organisation

unheard of before in any ungulate.” (Feh et al. 2001,

p.59)

“Herd size may vary. In E. hemionus, average herds

contain between 10 and 20 animals, with one male and

many females. However, in some ecological conditions,

pressure from predators causes small single male groups

to come together. (Nowak, 1999)” (Grogan 2005; E. h.

kicked by dominant ‘bullies’.68” (McGreevy 2004, p.140)

“Even in captivity, male Przewalski horses devoted more

time to behaviors that would be necessary for harem

acquisition and defense in the wild. Males were more active

than females; they locomoted more and exhibited greater

numbers of behaviors per hour. Females devoted more time

to foraging and interacting with other herd members.

Females mutual groomed more than males: perhaps

because they had kin in the herd, whereas males did not.

Solitary males and bachelors exhibited more transitions per

hour than harem males. Solitary males paced more than

bachelors or harem males. Pregnant and/or lactating

females spent more time feeding than barren females.

Horses in small enclosures spent less time stand-resting

and changed behavior states frequently.” (Boyd, 1988)

Self-mutilation and autoaggression represent management-

and handling-induced aberrant behaviors, which, according

to a Canadian study (Luescher et al. 1996), occur in 1.9%

of stallions and 0.7 % of geldings…..Self-mutilation is

almost exclusively shown by stallions, with domestic

stallions as well as Przewalski stalions in zoos being equally

affected…..Although the optimal solution would be free

contact with other horses, there is a risk of injury to other

horses with this type of aberrant behavior…. In stallions in

particular, sufficient social contact should be provided.

(Zeitler-Feicht 2004)

The prevalence of the various types of self-mutilation in

horses is not known. In one survey, results suggested that

self-mutilation of one form or another has been observed in

as many as 2% of domestic stallions. Among and between

stallions, self-mutilation varies in frequency and intensity,

and can reach levels that are fertility and/or life

threatening.(Mc Donnell 2008: domestic horses)

A chronic dilemma of captive breeding is that institutions

are often unable to provide sufficient housing for those

species they wish to propagate. This is particularly evident

to managers of Przewalski horses (Equus przewalskii).

Orthodox management of the species in harem bands (1

stallion with several mares and foals) and a roughly even

sex ratio at birth (Buisman and van Weeren, 1982 ) have

combined to produce an abundance of so-called "surplus"

(i.e. non-harem band) males. Traditionally maintained

singly (or rarely in pairs or trios) because of their

aggressive nature (Reindl and Tilson, 1985a), these males

are accumulating to the point where they present one of

the largest remaining obstacles to the successful long-term

maintenance of the species in captivity (Bouman, 1982;

Bouman-Heinsdijk, 1982; Ryder and Wedemeyer, 1982).

….For many polygynous species, maintenance of surplus

males in bachelor (i.e. all-male ) groups is the strategy of

choice because of the unique management and exhibition

advantages conferred (Berry, 1985). However,

acknowledging the aggressive nature of Przewalski horses,

is this a realistic alternative, or simply a euphemistic means

of eliminating problem animals through combat-related

onager)

In onagers, greater turnover of individuals within groups

and lower selectivity in association choices appear to

result in nonconcurrent interactions: individuals tend to

be in groups with their various preferred associates at

divergent times, rather than occurring with all these

associates in the same group. (Sundaresan et al 2007)

Khulans of this subpopulation [in Gobi], unlike other Asian

and African wild asses, form year-round stable, non-

territorial families. These families and all-male groups join

together into ‘‘bands’’ in winter, and herds of several

hundred animals, where reproductive rate is highest, form

throughout the year. The existence of such herds may

thus be critical for the breeding success of the population.

[p.51] (Feh et al 2001)

“We conclude that male Asiatic wild asses in our study

population are territorial.” (Saltz et al. 2000)

“During the breeding season, males spent more time in

close association with female groups, adopting what may

temporarily appear to be a harem breeding strategy.”

(Saltz et al. 2000)

“E. kiang usually form family groups of 5-400 animals.

The group is led by an old female, and is tightly

cohesive.” (Wang 2002)

“Equus kiang is often found alone or in small groups but

also may form herds of several hundred individuals

(Foggin 2000; Paklina and van Orden 2003; Schäfer

1937; Schaller 1998). (…) Group size appears to vary

seasonally. Individuals of E. kiang are dispersed in

summer and tend to congregate in fall and winter after

rut.” (St-Louis & Côté 2009, p.7)

“Khur have territorial harem type social organization

(Shah and Qureshi 2007).” (Moehlman et al. 2008)

“In the wild they travel in herds of several individuals to

up to one hundred individuals, with an average groups

size of 4.7 animals.” (Huggins 2002; E. asinus)

“Equids form two types of social structure, Type I, female

defence polygyny is seen in wild and feral horses that live

in bachelor bands or family bands/harems consisting of a

stallion (occasionally multiple stallions) and a number of

breeding mares and their offspring. (…) In contrast, Type

II, territorial defence polygyny is often seen in wild asses

that are more territorial, with a dominant male tending to

guard an area and breeding with females in the territory

(Klingel, 1975; Linklater, 2000).” (Proops et al. 2012,

p.337)

“Similarly, a study of the dispersion of kulans, Asiatic Wild

asses (Equus hemionus kulan) and takhi, Asian Wild

horses (Equus przewalski) showed that the two species

actively avoid one another, visiting areas at different

mortality? ….Combat-related permanent disability and

death have been observed in captive Przewalski horses.

Boyd (1986) reported that an 8-year-old stallion per-

manently lamed a 3-year-old colt when the 2 horses were

placed together. ….. In equids other than Przewalski horses,

this phenomenon has been observed in Moun- tain zebras

(Penzhorn, 1984) and feral horses (Berger, 1986). Berger

(1986) reports that 96% of the adult male feral horses he

examined showed signs of bite-related wounds -- 1 stallion

displayed over 50 distinct scars!.(Tilson 1988)

times of day. The kulans completely stopped using areas

that became horse strongholds, suggesting that the

horses are the dominant species (Bahloul et al., 2001).”

(Proops et al. 2012, p.341; over dat paarden vaak

dominant lijken te zijn over ezels in gemixte groepen)

“Horses familiarize themselves with foreign objects by

smelling them. Social exchange by sniffing one another’s

breath with or without an open mouth, represents an

important part of greeting rituals between horses.”

(McGreevy 2004, p.46)

“Each horse’s position within the band is held through a

blend of aggression and appeasement behavior.

Aggressive behavior can be biting, kicking, circling and

displacement, but the most common response to a

competitor is a threat to kick or bite.” (McGreevy 2004,

p.125)

“Under free-range conditions, even where the territory is

extensive, group bonding is important to the extent that

horses maintain continual visual and, to an extent

olfactory, contact with each other.36 A central mechanism

of band cohesion is the establishment of pair bonds and

mare-foal bonds.55 Pair bonds in bachelor groups are

generally weaker than those observed in natal bands and

become more tenuous as bachelors mature and are driven

to establish reproductive relationships.16” (McGreevy

2004, p.129)

“The most common form of aggression among all horses

is the head threat, which involves the extension of the

aggressor’s head and neck towards another individual

while flattening the ears against the head.” (McGreevy

2004, p.133)

“The hindlegs can be especially effective in aggression,

and threats to use them involve simply moving the

hindquarters near another animal, or lifting, occasionally

hopping and ultimately kicking with the hindlegs.”


“Horses are remarkably accurate when they choose to

strike with their hindlegs,68 and this is why kicks that do

not engage on the protagonist are described as threats

rather than misses.” (McGreevy 2004, p.134)

Features of the ethogram employed in aggression: bite,

bite threat, chase, ears laid back/pinned, grasp, head

threat, interference, kick, kick threat, lunge, nip, pawing,

push, rear, rump presentation, stomp, strike, strike threat

(McGreevy 2004, Table 5.5, p.123-137)

“The social structure adopted by donkeys in any particular

area is dependent on the availability of resources such as

food and water. In bountiful environments, donkeys use a

natal band system, similar to those of horses and ponies,

with complex hierarchies within the groups, in which rank

is not a simple function of age, sex, aggression or weight

(Jane French, personal communication). In arid and semi-

arid regions, a loose social structure (also typical of

African asses and Grevy’s zebras) exists with temporary

groups of males, or females, or males and females

predominating while some jacks become solitary. Small

aggregations in this system rarely last more than a few

days – membership is very fluid with mixing and splitting

of groups occurring when animals congregate, for

example at watering sources. There is no aggression

between groups. Dominant jacks do not maintain a harem

but dominate breeding activity within a large area, called

a lek. The only permanent association is between a

female and her foal, who travel together unassociated

with others. This behavioral characteristic has a profound

effect on the frequency of interactive behaviors, e.g.

social play is rare in donkeys when compared with harem

equids.74” (McGreevy 2004, p.139)

Equus hemionus are monogamous. Stallions tend to stay

with the mare and foal year-round. (Feh et al, 2001)

(Reuter, 2002)

According to these observations, the social organization of

E. hemionus is of the same type as that of Grevy’s zebra

Equus greÍyi. Klingel, 1974a., being characterized by the

very large territories and the conditional defence by the

territorial individual. (Klingel 1998)

During lengthy steppe migrations, which were

characteristic of kulans in Central Kazakhstan, they (E.

hemonius) assembled in herds of 1,000 and more

(Rychkov, 1762). (Formozov 1966)

In the networks of onager nonzero and preferred

associations, most individuals are in one component. Thus

we would expect diseases and information to spread

through most of the onager population. (Sundaresan et al

2007)

All authors having studied Mongolian khulans report the

existence of big herds, such as Dulamtseren (1989),

Bannikov (1958) or Andrews (1932) for the first half of

this century. More recently, Owen and Munkhtuya

(personal communication) saw two herds of ca. 900

individuals in the Galpin Gobi (South-Gobi district). In

1994, Schaller, (1994; personal communication) met two

herds of 234 and ca. 500 khulans. Finally, Mix et al.

(1995) identified four herds consisting of 500–650

khulans each during an aerial count, and on our own

expedition in September 1996 we saw one herd of 1241

individuals. [p.59] (Feh et al 2001)

E. kiang: uitgesproken kuddedier [volgens de Duitse

zooloog en Tibetreiziger Ernst Schaefer] ... Ik ken geen

ander in het wild levend dier waarbij de orde in de kudde

sterker en imposanter tot uiting komt dan bij de Kiang.

Als de kudde van een paar honderd kiangs in wilde vaart

door de zonovergoten steppe raast, lopen de dieren

precies achter elkaar’ ... ondanks deze orde in de kudde

verdragen de dieren elkaar slecht; [vol XII, p660, kol. 1]

(Grzimek 1973)

“Combat over young mares in estrus is not ritualized, and

involves running, circling, neck-wrestling, biting from

standing and sitting positions, and kicking.” (Grubb 1981)


Sociale stress (incl. [orale] stereotypieën) en verwondingen a.g.v. veranderingen van groepssamenstelling bij sociale

soorten (m.n. E. burchellii (Burchell’s zebra), E. zebra (mountain zebra), and E. caballus przewalskii (Przewalski’s horse)

die in harem groups leven).

Agressie: Gevechten tussen mannen ivm territorialiteit en competitie merries– (bijt)trauma, verwondingen mn aan

nek en staart (incl tetanus) tot zelfs sterfte (ook bij elkaar zien!) Wonden zijn vaak lastig te behandelen bij deze dieren

door onbenaderbaarheid en moeite bij herintroductie in de groep! En Verhoogde agressie in gevangenschap (schoppen

en bijten) intergroup (merries) en within group (merries onderling en hengst naar merries) > verwondingen, trauma

(Bijten naar merries is normaal gedrag in kader van reproductie, personal communication FV)

Sociale stress automutilatie en pacing) a.g.v. sociale deprivatie (solitair houden komt voor in captivity bij hengsten)

Personal communication Chantal: bij hengsten die solitair gehouden worden komt automutilatie voor; obv vergelijkbare

stereotypieën bij andere paardensoorten ook hier te verwachten

(Aversive) stimulation (e.g. introduction of other animals) ->Zebra’s are easily agitated -> veritable frenzy ->often

injured (Kleiman et al. 1996, p.207) (zie Schuilen en benaderbaarheid)

Trauma resulting in fracture in any adult is usually fatal, as adult wild equids are notoriously uncooperative patients.”

(Nelson 1986) (Zie schuilen en benaderbaarheid); Volgens ref Nelson 1986 van de 39 dieren die voor pathologie warden

aangeboden 14 dieren die trauma hadden

Schuilen en benaderbaarheid (veiligheid en schuilgelegenheid)[Incl. thermoregulatie en klimaat]

In de natuur:

“Within their natural range, many equids are subjected to

extreme weather conditions. For example, temperatures

range from -45°C in the winter to approximately 40°C in

the summer, with diurnal temperature ranges in excess of

35°C at the reintroduction sire for Przewalski horses in

the Dzungarian Gobi in Mongolia” (Walzer, 2003)

“The stripes may function in thermoregulation and provide

as a natural suncreen. Differences in cooling of the black

and white stripes create a rotary breeze.” (Colvin &


“During cold weather, they often look for shelter in

wooded ravines and shallow caves, and visit east-facing

slopes on cold mornings to sun themselves.” (Walker

2005; E. zebra)

“During cold weather, mountain zebras seek shelter in

wooded ravines and shallow caves (Penzhorn, 1984b).

They often graze in the lee of a hill or ridge, protected

from the wind.” (Penzhorn 1988, p.5)

Przewalski horses were less adapted to semi-desert

In de houderij:

“Physical restraint is unsatisfactory for any wild equid

except when it is very young. In fact, tranquilization

without use of an immobilizing agent is dangerous to

personnel involved, because of the unpredictable behavior

of these animals.

Anesthetic shock has occurred in a number of species and

care must be used during immoblizations. Handling zebras

in ambient air temperatures above 26.6 C is dangerous.

Certain individuals in all species immobilized have

experienced a catatonic state, which includes profuse

perspiration even in moderate ambient conditions. The

rigidity and tremors make intravenous injection difficult, but

intramuscular injection of the antagonist to etoprhine does

not produce satisfactory results in these situations.

Hyperthermia resulting in death in frequently seen if

recovery is prolonged….

Wild equine species are also affected by capture myopathy”

(Nelson, 1986) (N.B. Aantal dieren met anesthestic shock

betrof 6/39 dieren voor pathologie – informatie echter

achterhaalt in deze tijd; aantal dieren uit groep en over

conditions (both water and vegetation needs) than kulan.

[p309] (Bahloul et al 2001)

When escaping from a pursuit, dzerens develop speeds up

to 60-65 km an hour, kulans-up to 60-70 km an hour.

These and saigas have great endurance. For their night

rest all three species choose flat, open spaces, where it is

easy to notice an approaching danger from a distance and

where it is possible to flee without encountering natural

obstacles. Extensive plains or even vast valleys are most

attractive for these species. They definitely avoid places

with a greatly dissected relief, but under some conditions

must use them. Thus, in winter they hide from cold winds

or snow storms behind sand dunes or in ravines, while in

the summer they find here cooler places for resting during

the heat of the day. (Formozov 1966)

All equids are relatively swift, alert runners and generally

flee from danger rather than fight. (Ballanger & Myers

2001)

E. hemionus has a well-developed strategy for anti-

predator defense. Stallions from more than one family

group cooperate to chase off predators. The frequent

occurrence of large groups aids this ability. Wolves are

the only known predator of the Asian wild ass other than

humans. (Feh et al 2001)

Khulans have developed an elaborate anti-predator

defence strategy through group formation. Khulan

stallions of the Gobi B population stay with the mares and

foals year round and actively take part in the defence of

offspring (Feh et al., 1994, this study). When several

family groups join up, as is usually the case in winter,

khulan stallions from different families chase wolves

cooperatively. [p.59] (Feh et al 2001)

the usual flight distance of the animals from cars was ca.

2 km),.p.59] (Feh et al 2001)

welke periode is overigens onbekend)

“Physical restraint cannot be recommended for any wild

equid except possibly for newborn foals.” (Walzer, 2003)

“As in other ungulates, capture myopathy is a potential

problem in nondomestic equids in relation to capture events

and anaesthesia.“

“While domestic horses tend to rest in sheltered areas,

Przewalski horses when released into extensive enclosures

have been noted to rest in the highest parts of the available

terrain.42” (McGreevy 2004, p.232)

We have found that great care must be taken with newly-

caught young wild asses if they are to adjust to captivity. If

trapping operations are not carried out properly the wildass

will often die either during or immediately after capture. In

the past, newly-captured foals often refused to eat and

died. However, we have evolved a method that works well.

We place the newly-captured young animal with another

young conspecific. This companionship seems to give them

a sense of security and confidence and they soon start

eating. We give them sweetened milk (sweetened with

jaggery) with proteins and carbohydrates added. To this,

moist bran and gram is gradually added. The young wild

asses are also fed carrots, lucerne and hay. When they are

six months old the milk diet is discontinued. We provide

them with plenty of rock salt, placed in different corners of

the enclosure, and from the start they love to lick it. Even

when young, the foals are difficult to approach, for like the

adults they kick and bite simultaneously. Gradually they

become friendly but they never become really tame and

suitable for riding. [p. ] (David 2007)

“A condition described but still poorly understood is capture

myopathy. This is usually seen in animals subjected to a

great amount of exertion, either before or after darting and

before immobilization is achieved. Most consistent

postmortem findings include pulmonary edema, myocardial

hemorrhages, and muscle degeneration” (Nelson, 1986)

“Physical restraint is not often practical in adult equids

because of their size and strength…. Animal care staff must

work cautiously around any nondomestic equid because

stay may startle and bolt unexpectedly into solid obstacles,

which may result in fatal neck or head injuries”(Janssen &

Allen, 2014)

“Hoof abscesses from wounds, cracks, bruises, which is

more common in Somali wild asses resulting in sudden

severe lameness and can be prevented by avoiding extreme

wet or dry conditions” (Janssen & Allen, 2014)

“Most equids are managed in a similar way in zoos. Equids

are generally hardy and withstand normal to severe

temperature variations as long as shelter and protection

from wind and sun are available. Equids acclimated to the

southern Californian climate do well in temperatures

ranging from 0 to over 38 C. Grevy zebras are reported to

be less cold tolerant in zoos compared with other zebras.13

Shelter should be provided to keep food dry and to provide

shade.” (Janssen & Allen, 2014)


Lastigheid van hanteren > vaak sedatie of speciale box nodig

Anesthesie en vangen: Shock en sterfte a.g.v. anesthesie (risico lijkt wat groter dan bij andere soorten), Risico op

hyperthermie en sterfte indien hanteren tijdens warmere dagen (>26.6 C), Vangen/anesthesie en bijbehorende stress >

Capture myopathy (nauwelijks meer gezien bij zebra’s e.d – personal communication FV)

Schrikreacties door plotselinge prikkels of benadering > Trauma als gevolg van rennen of opspringen tegen obstakels

(personal communication FV – met name bij veulens)

Extreme natte of koude condities kunnen leiden tot ontwikkeling van brokkelige hoeven, hoefabcessen vanuit wonden,

scheuren of kneuzingen (mn in Somali wilde paard en kou in Grevy Zebra) (niet op drassige gronden plaatsen helpt dit

voorkomen en beperkt de problemen; personal communication FV)

Wilde jonge ezels eten niet in gevangenschap > sterfte

Zebras and asses require simple shelters in the winter, depending on the local environmental conditions these will in

individual cases have to provide heating. (Walzer, 2003)

Toiletteren (schoonmaakgedrag)

In de natuur:

“In addition to maintaining the health of the integument,

grooming behaviors can contribute to the affirmation of

social bonds not least by reinforcing affiliations and

sharing odors.” (McGreevy 2004, p.223)

“Allogrooming is frequent in the species. Grooming zebras

usually stand side by side, head to tail.” (Grubb 1981)

“Lastly, these distinctive stripes may protect against

tsetse flies. One study demonstrated that tsetse flies

prefer solid verses striped objects. “(Colvin, 2009)

“Allogrooming occurs between individuals, especially

mares and their foals and stallions and their preferred

mares. This is achieved when the grooming zebras stand

side-by-side, head to tail and is effective in removing

parasites and strengthening social bonds.” (Colvin &


“Grooming patterns are essentially similar in all equids

(Klingel, 1972; Trumler 1959).” (Penzhorn 1988, p.5)

“Grévy zebras do not mutually groom (…). All zebras can

twitch their skins to dislodge flies, using a subcutaneous

musculature that is absent where the hind-quarters are

flicked by the tail. Toilet is conducted by tail swishing,

rubbing the head against the legs or against parts of a

conspecific, scratching with a hindhoof, and nipping with

incisors. Rolling in dust, water or mud helps protect

against insects; when dried muds falls away it dislodges

ticks or other ectoparasites. Zebras roll or scratch against

trees, rocks, or termite mounds to relieve itchy skin, hair

In de houderij:

“De vacht van de ezel bevat veel stugger haar en is meer

waterdoorlatend dan het paard doordat zij minder vet in

hun haarkleed hebben. Hierdoor zijn ze vatbaarder voor

kou en nattigheid en dient de poetsfrequentie laag

gehouden te worden om het weinige vet in de vacht te

behouden ter bescherming.” (de Graaf-Roelfsema 2009,

p.25)

Rolling is an elementary need. Well cared-for horses have

the same urge to roll on a daily basis as free roaming

hoofed animals….Rolling has probably additional

communicative function. This is suggested by the intense

olfactory investigation of the ground prior to lying

down….(Zeitler-Feicht 2004)

or parasites (Haltenorth and Diller, 1980; Lydekker,

1926).” (Churcher 1993, p.6)

“In both species the stallions defecate preferentially onto

their own dung or on dung of conspecifics, thereby

producing conspicuous dung piles.” (Klingel 1998; E.

hemionus & E. africanus)

There is some tendency for males to groom certain

females preferentially. Conversely, males may selectively

attack particular individual females, at least in captivity.

(Grubb 1981)


Ezels en –achtigen stugger haar dat meer waterdoorlatend is + gebrek aan schuilmogelijkheden > Vatbaar voor kou

en nattigheid met secundair ziekteproblemen (zie schuilen en benaderbaarheid)

Gebrek aan mogelijkheden tot grooming: allogrooming > evt. parasieten; sociale stress en stereotypieën: zie ook

sociale gedrag),

Gebrek aan modder, zandbad of materialen om zichzelf te krabben > Slechte vachtconditie en ectoparasieten (bij

paarden voldoet aan criteria ethologial need).

Time-budget (periodes van activiteit of inactiviteit gedurende een dag of een seizoen)

In de natuur:

“Equids are active both day and night but are mainly

crepuscular.” (Ballenger & Myers 2001)

“Roberts (1977) indivated that E. h. khur, of India and

Pakistan, grazes at night.” (Nowak & Walker 1991,

p.1308)

“Other observations indicate that feral horse activity may

be either diurnal or nocturnal, that most of the day is

spent seeking and consuming food, that feeding peaks

may occur in the early morning and late afternoon, and

that there are several daily periods of rest.” (Nowak &

Walker 1991, p.1311, about E. c. przewalski and feral

horses)

“Although the only true wild horses (E. przewalskii) are in

or from captive populations, we can use their behavior

and that of feral E. caballus as guidelines for what could

be regarded as normal equine behavioral organization. Of

the major groups of behaviors, those that occupy most of

a free-ranging horse’s day are the search for choice

grazing spots and the ingestion of forage. Equine feeding

control mechanisms appear to have evolved to maintain a

high gut-fill.77” (McGreevy 2004, p.194)

“DAILY RHYTHM: generally active throughout the day. At

sunrise leave sleeping place for the grazing areas (in

single file) where eating, drinking, rolling, resting

alternate. In late afternoon return to sleeping area. At

night 3 rest periods with short grazing spells in between.

In cold or rain stay near the sleeping area.” (Haltenorth &

In de houderij:

“Equids spend more than half of the day feeding. Lactation

is a very demanding form of maternal investment. In an

environment such as a zoo, where no grass but several

feeding sites are present, conflict between suckling

behaviour of the foal and feeding behaviour of the lactating

mother should occur. The results of our study show that

feeding while suckling could reduce parent-offspring conflict

and improve welfare of captive foals and mares.” (Pluhácek

et al. 2010; Equus burchellii)

“Overall, the horses spent 46.4 _+ 5.9% of their time

feeding, 1.3 _+ 0.1% nursing, 0.5 +_ 0.1% drinking, 20.6

+_ 5.4% standing, 15.7 ± 3.2% stand-resting, 1.7 _+

0.2% self-grooming, 2.2 _+ 0.7% mutual grooming, 7.4 _+

1.0% locomoting, 1.2 ± 0.3% playing, 1.2 _+ 0.5% lying

laterally and 4.1 ± 3.0% lying sternally. The horses

averaged 45.2_+ 5.8 behavioral states per hour, and 0.2 +

0.0 defecations, 0.3 ± 0.0 urinations, 1.5 ± 0.3 aggressions

and 0.7 _+ 0.1 vocalizations per hour. The horses spent the

greatest amount of time foraging between 20.00 and 04.00

h, when the temperatures were lower. They spent 68.2 +

2.2% of their time between 20.00 and 24.00 h feeding, but

only 31.2 _+ 2.1 ?~ of their time feeding between 08.00

and 12.00 h. Recumbent rest was most common between

00.00 and 04.00 h. As temperatures rose during the

daylight hours, the horses spent more time drinking and

standing, rather than grazing. Stand-resting was the most

common form of rest during the day. The horses exhibited

the greatest number of activities per hour from 08.00 to

20.00 h. While standing in close proximity to one another

during these hours, the horses exhibited the highest

Diller 1988)

“Zebra showed activity phases after midnight, in the

morning, afternoon, and in the evening, but never around

noon.” (Scheibe et al. 2009)

“Grazing and resting occupy most of the daylight hours.

There are generally three main grazing periods during the

day: a few hours after dawn, late morning, and from mid-

afternoon to dusk.” (Penzhorn 1988, p.5; about E. zebra)

“They are active nocturnally and they feed primarily at

night” (Wang 2002; E. kiang)

“There is a daily cycle of movement covering as much as

13 km and generally from the sleeping sites in higher,

more open areas to the lower, lusher grazing grounds.”

(Novak, 1999)

Despite an absence of non-human predators, the

proportion of dusk time budget allocated to vigilance was

high, at 41% for males during periods when they

accompanied stable female groups and 12% for females

during these same periods (Simpson & Nicol, 2012)

number of aggressions per hour ( 1.9-2.4 ).” (Boyd, 1988b)

“The horses then spent the evening grazing and recumbent

away from the barn. Grazing peaked between 20:00 and

24:00 h when the horses spent 68.2% of their time in this

activity. Recumbency peaked at 21.4% of the time budget

between the hours of 00:00 and 04:00 (Boyd et al., 1988).

Thus, the horses were making use of their entire enclosure

and were taking advantage of the forage present but were

waiting until evening to do so. Grazing during summer

evenings may have minimized thermal stress, and at night

there was no reason to linger near the barn as no visits by

staff were expected.” (Boyd, 1991)

“Social integration within the Takhi herds was very high

from the beginning, as described by the spatial relation and

synchronisation data. Between 50 and 89% of the

observation time, the behaviour of all herd members was

synchronised. The amount of time spent grazing by the

Takhi (30-68% of the daylight period) was similar to that of

feral horses and Takhi in captivity and semi-reserves. The

Takhi tended to rest in the morning and have a bimodal

period of grazing at dawn and in the afternoon. The Takhi

displayed clear habitat preferences for certain activities.

They had a strong preference to rest at the highest point in

their enclosure. They fed preferably on two or three

different vegetation types (with five types available in each

enclosure). The amount of time spent grazing during the

non-growing seasons (49 + 15%) indicates that the feeding

value and availability of food were sufficient.” (Van

Dierendonck, 1996)

“Because the wild equid counterparts spend up to 18 hours

grazing, it is important to establish a feeding regimen in

captivity that emulates this behavioral necessity. This is

ideally carried out by providing pasture but can be

simulated with scatter feeding and multiple feeding

installments. Though most species are pure grazers, the

provision of browse has shown to be beneficial from a

feeding and environmental enrichment standpoint in many

species…” (Walzer, 2003)


Onvoldoende foerageer mogelijkheden > Orale stereotypieën (zie foerageren en voeding)

Zintuigen (prikkels en afleiding)

In de natuur:

“Communication of moods and other information takes

place with changes in ear, mouth, and tail positions. Also,

some vocal communication through nickering takes place

in horses and zebras.” (Ballenger & Myers 2001)

In de houderij:

“One of the most important elements provided by the

environment is mental stimulation. Without this stimulation,

problem behaviours develop; these behaviours are often

designed to relieve the boredom of domestication.” (The

Donkey Sanctuary: ‘Understanding donkey behaviour’

“Horses have been described as being among the most

perceptive of animals.1” (McGreevy 2004, p.37)

“As a herbivorous flight animal, the horse has good

distance vision, allowing it to scan widely for danger and,

despite being relatively poor at accommodation, with a

vertical field of 178°,4 is able to visualize the ground

immediately ahead while grazing.” (McGreevy 2004, p.37)

“The high proportion of rods to cones (generally 20:1)13

gives the horse excellent night vision but insufficient to

make horses innately fearless of areas that are poorly lit.”


“SUMMARY OF KEY POINTS

The horse has:

• almost 350° vision

• a caudal blind spot that accounts for a proportion of

startle responses

• dichromatic color vision (i.e. like a color-blind person)

• a sense of taste that discriminates between safe and

toxic plants with variable accuracy

• highly developed accessory olfaction

• the ability to hear within and beyond the range of

human hearing

• predictable zones of very sensitive cutaneous

sensation.” (McGreevy, p.51; summary of chapter 2:

Perception)


• Because horses have excellent vision, the nuances of

equine body language can be very subtle.

• Ear position and head posture are the most important

variables in non-vocal communication.

• Tail positions can help to coordinate the movements of a

group of horses.

• The olfactory cues in urine and the visual stimuli offered

by characteristic estrous urination postures are used by

mares to communicate their readiness to mate.

• Dung-piles and the rituals attached to them are effective

means of avoidance of aggressive interactions between

stallions.” (McGreevy 2004, p.162, summary of chapter 6:

Communication)

“Six calls and two facial expressions are used in

communication between individuals. Three of the calls are

used as predator alert or threat calls, one is used to

communicate injury, another is used in distress, and the

last one is used in contact between individuals.” (Colvin &

Nihranz 2009; about E. burchellii)

“Horses maintain visual contact with their family and herd

at all times and have a host of ways to communicate with

one another, including vocalizations, scent marking, and a

wide range of visual and tactile signals.” (Wikipedia

[Engels]: Przewalski’s horse)

The male sniffs the urine or feces of estrous females and

then may show flehmen, followed by urinating or

[Online])

defecating on the deposits of the female. Precopulatory

grooming of young mares occurs, with copulation only on

the third or fourth day of estrus. (Grub 1981)

hearing good, sight not bad, scent moderate. (Haltenorth

& Diller, 1988)


Onvoldoende stimulatie > Probleemgedrag en stereotypieën (obv bevindingen bij vergelijkbare diersoorten); zie ook

foerageren en voeding (deze arg komt echter van ezel en niet van overige equiden)

Foerageren en voeding (foerageren en eten, waaronder de rantsoensamenstelling) [incl. drinkwater]

In de natuur:

“They are entirely herbivorous, feeding largely on grass

and some browse. Most drink water daily, although they

can go without water for long periods of time.” (Ballenger

& Myers 2001)

“Grazing is the preferred means of ingestion in adult

horses, but browsing is also adopted when grass becomes

particularly scarce.37–39 Although, like horses, donkeys

graze, they tend to select coarser grasses and

demonstrate their greater agility as they exercise their

preference for browsing. Because of their motivation to

select a variety of forage, donkeys are in greater danger

of ingesting poisonous plants than horses are.” (McGreevy

2004, p.192)

“Live in mesic grasslands and are capable of migrating

long distances. Their ability to utilize coarser vegetation of

poorer quality means that they can thrive and survive in a

range of habitats.” (Grzimek et al. 2004; E. burchellii)

“Het zijn echte grazers, die zich aan alle grassoorten en -

hoogten kunnen aanpassen. Ook eten ze twijgen,

scheuten en bladeren van bomen en struiken. Ze zijn

zelden ver van water te vinden, maar mijden modderige

grond.” (Wikipedia [Nederlands]: Steppezebra)

“The diet is 90 percent grass but also includes some

browse.” (Nowak & Walker 1991)

“Burchell’s zebras gather grass by clipping it with their

upper lip and lower incisors. They are also well-equipped

with large grinding molars which are able to process the

tough plant material. Their diet is low in protein, but they

process large amounts” (Colvin & Nihranz 2009; E.

burchelli)

“Kulans are typically more migratory-active and better


they can regularly do without water, or use water with

very high salinity (Rashek, 1966).” (Bahloul et al. 2001,

p.320)

“Przewalski horses were less adapted to semi-desert

conditions (both water and vegetation needs) than kulan.”

In de houderij:

“In general the equids are bulk-feeding grazers that spend

up to 18 hours a day feeding. Equids have high-crowned

teeth that resist the fibrous defenses of the various grasses

(Table 57-3)… The equids can use forage with high levels of

fiber because bacteria in the cecum and colon break down

hemicellulose and cellulose. However, unlike in the

ruminants, the site of fermentation is distal of the small

intestine – the primary site of absorption of bacterial

protein and water-soluble proteins – therefore amino acids

and vitamins are necessary in the diet. Whereas mountain

zebras do not use any browse, the Grevy’s zebra diet is

reported to comprise up to 30% browse.

In a captive environment wild equids do not have unique

feeding requirements. They are maintained on good quality

mixed grass in the summer months and on mixed hay in

the winter months. Supplementation with 12% protein

pelleted horse feed may be carried out in winter and in

lactating females, but because obesity is a problem in many

captive equids, it is normally unnecessary. Depending on

the size of the herd and the social system, an adequate

number of feeders need to be provided because the

dominant female and male may not allow other animals to

feed while they are at the feeder. Mineralized and plain salt

blocks should be provided.” (Walzer, 2003)

“Obesity is a familiar occurrence in most wild equids kept in

small areas with free access to feed. Where available,

pelleted horse feed, usually 12 to 12.5 percent protein, is

an efficient method of supplying adequate nutrients. Most

animals in confinement get bored with little to do, so a

limited amount of pellets, from 1.0 to 1.5 kg/100 kg body

weight, in addition to a low-quality high-fiber hay of equal

amount makes a good ration.

In cold weather and for lactating females and young foals, a

supplement that provides more energy should be added.

The practice of giving a higher-protein ration to mares

during late gestation or to foals during the first few months

of growth does not appear necessary in wild equids

At least two feeders should be provided in any unit housing

these species. If a sizable group is housed together

(Bahloul et al. 2001)

“While the horses eat a variety of different plant species,

(…) the Przewalski horses have seasonal food

preferences.” (Wikipedia [Engels]: Przewalski’s horse)

Foraging strategy affected the extent of directed

movement: zebras with a confined dispersion of grazing

patches around the central place directed their

movements over a longer distance. Zebras may extend

the distance at which they can direct their movement

after improving their knowledge of the local environment.

(Brooks, 2008)

The Asian wild ass is strictly herbivorous. They tend to eat

perennial grasses (noncotyledons) that are of species of

Stipa or Agropyron. They also eat herbs and bark. (Glenn,

1999) (Reuter 2002)

Kulans are typically more migratory-active and better


they can regularly do without water, or use water with

very high salinity (Rashek, 1966). [p320] (Bahloul et al

2001)

Dental abnormalities in animals are often artifacts of their

domestication or captivity, e.g., crib biting in horses

resulting in uneven wearing of the incisors (Jubb and

Kennedy, 1970, Pathology of Domestic Animals, Vol. 2,

Academic Press, New York, 697 pp.). Little has been

documented on dental abnormalities in free-ranging wild

animals, especially of the African species, although some

descriptions were included in a general survey based on

mu seum specimens (Colyer, 1936, Variations and

Diseases of the Teeth of Animals, John Bale, Sons and

Danielsson, London, 750 pp.)… During a 4#{189}-yr

study (1971-1976) of the ecology and behavior of Cape

mountam zebras in the Mountain Zebra National Park

(MZNP), Republic of South Africa, all skulls found in the

veld were collected for age determination (Penzhorn,

1982, Koedoe 25: 89-102). A few of the skulls exhibited

dental al)normalities and these are described here. The

zebra population of the NIZNP is about 200. (Penzhorn

1981)

“All equids are bulk-feeding grazers, feeding primarily on

grass and roughage. In the wild, grass constitutes over

90% of the common zebra’s diet. They may resort to

some browsing and digging of plant materials in the dry

periods. Grevy zebras reportedly eat some browse in the

wild.19 As with other perissodactylids, equids are hind gut

fermenters and have a large cecum and colon.” (Janssen

& Allen, 2014)

“Strangles (Streptococcus equi) acquired from domestic

horses at shared water points has proven to be a

significant morbidity and mortality factor in reintroduced

Przewalski’s horses in the Gobi desert in Mongolia.”

(Walzer, 2003)

observation will dictate the number of feeders and feeding

space required. As basic guideline would be 1.0 m of feeder

length for every two animals. Often the males or dominant

female will not permit others to feed while they are at the

feeder.

Sand colic is not uncommon in wild equids but can be

greatly reduced by covering the area around the feeder

with another type of surface

Blocks of both mineralized and plain salt should be

available; these should also be placed on surfaced area and

protected so moisture runoff does not result in a natural

salt lick in the surrounding soil” (Nelson, 1986)

“… Fresh water must be provided at all times. Wild equids

may be trained to use automatic drinkers. ” (Walzer, 2003)

“Enteritis – either infectious or resulting from changes in

feeding – is a typical precursos to colic. Sand colic is not

uncommon and is best managed by providing concrete

feeding pads. Furthermore parasite controle is of utmost

importance in this respect Diagnostic and therapeutic

management is difficult because of the necessity of

chemical restraint.” (Walzer, 2003)

“Equine degenerative myeloencephalopathy (EDM) has

been described on numerous occasions in Przewalski’s hors,

zebras, and kulans. This disease is an idiopathic, diffuse,

degenerative disease of the spinal cord and selected parts

of the brain… Vitamin E deficiencies in the first year of life

have been implicated in the development of this disease. In

most cases the onset and progression of the clinical signs –

gait deficits, ataxia and hypometria that affect all four

extremities – is insidious and progressive. In zoo settings,

the prevalence of the disease appears to be correlated to

the absence of pasturage. In the early stages of the

disease, significant dosages of vitamin E... may result in

complete recovery.” (Walzer, 2003)

“Traditionally, horses are fed long fiber from a haynet or

hayrack. This is intended to reduce wastage from

contamination with urine or feces and possibly to decrease

the likelihood of endoparasite transmission. However, there

is growing evidence that this unnatural foraging position

can have a deleterious effect on the efficacy of the

mucociliatory escalator in clearing the upper airways of

inhaled particles (Fig. 1.8), especially those inhaled from

dried foodstuffs (which are to some extent unnatural in

themselves).72 Similarly, haynets reduce the space

available in the stable, may increase the risk of the horse

becoming snared (e.g. by trapping its hoof), and elevate

the forage so increasing its potential as a source of ocular

foreign bodies. It is also argued that feeding from a net or

rack may adversely affect muscles and nerves in the

neck.73 Perhaps this is why, when financial considerations

are almost insignificant relative to the value and

maintenance of performance – for example, in the majority

of racing yards74 – horses are fed roughage, whether it be

"Free-ranging ungulates frequently eat soil or visit mineral

licks. This has also been reported in equids (Vesclovsky &

Volf 1965: Smuts 1972: Feist & McCullogh 1976) During

an ecological and behavioural study of Cape mountain

zebras Eq1111s zebra zebra in the Mountain Zebra

National Park (MZNP). Republic of South Africa. stallions.

mares and foals were observed eating soil (Fig. I). A

number of licks was located throughout the Rooiplaat area

of the Park….Calcium was the only mineral with higher

concentrations at the licks than in all surrounding

samples. At two licks magnesium had a greater

concentration than in the surrounding samples. but the

third lick had a low magnesium concentration. The sodium

concentrauon of all three licks was less than that of most

of the surrounding samples. Potassium and phosphorus

had relatively high concentrations at one lick only…It is

likely that the mineral sought by wildlife at a lick is lacking

or insufficient in their diet. Rats with a specific nutritional

deficiency choose a corrective diet if available (Biedler

1961). No definite conclusions can be drawn from the

MZNP licks. but it appears that calcium may have been

the essential element. This may infer a shortage of

calcium in the diet of Cape mountain zebras in the Park.”

(Penzhorn 1982)

hay, haylage or silage, from the stable floor.” (McGreevy

2004, p.14)

“Koliek is een verschijnsel bij een aantal bij het paard

voorkomende ziektebeelden die allemaal worden

gekenmerkt door pijn in de buik. Koliek kan in ernst

variëren van mild tot levensbedreigend, het kan paarden

van iedere leeftijd of ras treffen.” (Wikipedia [Nederlands]:

Koliek (paarden))

“Restrictions on feeding behavior, and especially the

provision of discrete meals, lead to digestive anomalies and

behavioral frustration.” (McGreevy 2004, p.196)

Hyperlipaemia: “Donkeys (as well as some native pony

breeds) are particularly susceptible to this devastating

condition, which has been recognised since the late 1960’s.

These types of equines are designed to live in harsh

environments where vegetation may be sparse and of poor

quality, necessitating walking for up to 16 hours a day in

search of food. As a result they tend to put on weight and

gain excess fat reserves when living on the relatively lush

pasture and with limited exercise. Unfortunately when these

animals stop eating for any reason hyperlipaemia may

develop.” (The Donkey Sanctuary: ‘Hyperlipaemia’

[Online])

“… over-feeding frequently leads to health issues including

a propensity to obesity which increases the risk of related

diseases such as laminitis.” (The Donkey Sanctuary:

‘Feeding donkeys’, p.1 [Online])

“De meest voorkomende problemen van het

digestiesysteem zijn bij de ezel grofweg onder te verdelen

in gebitsproblemen, in het bijzonder bij geriatrische ezels,

en koliek.” (de Graaf-Roelfsema 2009, p.32)

“Obesitas is een algemeen en veel voorkomend probleem

bij niet-werkende ezels in West Europa. Dit heeft een aantal

nadelige gevolgen voor de ezel, namelijk verhoogde risico’s

op: hyperlipemie, hoefbevangenheid en witte lijn defecten.”

(de Graaf-Roelfsema 2009, p.38)

“We provide them with plenty of rock salt, placed in

different comers of the enclosure, and from the start they

love to lick it. Even when young, the foals are difficult to

approach, for like the adults they kick and bite

simultaneously.” (David 2007; E. h. khur)

“Free-ranging horses regularly consume soil,76 but it is not

clear what elements they are seeking when they do so. It

has been suggested that sodium, iron and copper

supplementation are among the benefits of this

activity.50,111” (McGreevy 2004, p.196)


• Food selection allows horses to adjust their intake of

nutrients to suit their current situation, while avoiding

poisonous alternatives.

• A horse is born with innate dietary preferences which

generally include:

– sweet-tasting foods

– grasses high in fiber and carbohydrate

– short, young growth.

• There is individual variation in preferences and aversions.

• Horses may learn from conspecifics and personal

experience which foods to select.

• Concentrated feeds are associated with reduced saliva

production and increased gastric acidity.

• Periods without food are associated with increased gastric

acidity and risk of gastric ulceration.

• Lack of forage is the most important management factor

linked with the development of stereotypic behaviors in

cross-sectional epidemiological studies.

• Lack of forage and the provision of concentrate feed are

important causal factors that precede the development of

oral stereotypies in young horses in prospective

epidemiological studies.” (McGreevy 2004, p.208, summary

of chapter 8: Ingestive behavior)

Finally, the exotic equids (eg, zebra, Przewalski's wild

horse) possess the same dental formulae as their domestic

counterparts, and, in many instances, some of the same

dental problems. (Knightly & Emily, 2003)

Reported manifestations of vitamin E deficiency amongst

exotic animals include… myodegenerative disorders in …

zebras Equus burcheli ssp (Brush & Anderson, 1986)

“Carotenoids may be important because of their role as

vitamin A precursors and as free radical trapping agents. It

has been suggested that dietary carotenoids influence

longevity in humans as well as animals (Cutler 1984a and

1984b) and may be essential for proper immune function in

humans and other species (Chew 1987, Chew et al. 1993).

Perissodactyla (Grant’s zebra, Grevy’s zebra, South

American tapir, black rhinoceros). Eighty percent of the

zebra diet was timothy hay; 20% was low fiber herbivore

pellets. Tapir diets were 25% timothy hay, 25% low fiber

herbivore pellet and 50% fruits and vegetables on an as-fed

basis. Black rhinos consumed primarily timothy hay (81%).

Low fiber herbivore pellets and fresh produce made up 10

and 9% of the diet, respectively, as fed. These diets were

moderate in carotenoid content.

Ungulates (Perissodactyla and Artiodactyla). With some

exceptions, the wide range of taxa which were grazers and

browsers and thus consuming moderate amounts of lutein

and b-carotene even in captivity, had nondetectable serum

concentrations of any carotenoid. Grant’s zebra and the

South American tapir had low concentrations of lutein, a-

carotene (tapir only) and b-carotene. Seven samples from

three black rhinoceros had no detectable serum carotenoid

concentrations despite moderate intakes of lutein and b-

carotene.

The horse-related species (Perissodactyla) had carotenoids

ranging from not detectable in the black rhinoceros to 233

nmol/L in the Grant’s zebra. Lutein was found in both the

Grant’s zebra and tapir serum, but only the tapir had a-

carotene. The diets offered were classified as moderate in

carotenoids, which makes these animals moderate or, in

the case of the black rhino, low accumulators. The serum

concentration of b-carotene in the Grant’s zebra was similar

to the 261 nmol/L reported in horses by Baker et al.

(1986), but considerably lower than the 939 nmol/L

carotene reported for horses by Vander Noot et al. (1964).

Horse-related species that accumulate may be selective

carotene accumulators like the bovids. (Slifka et al 1999)

“Although wild equids may go without water for long

periods in the wild, they drink water when it is accessible so

they should have a constant supply of fresh water when

managed in zoos.” (Janssen & Allen, 2014)

“In general, non-domestic equids have no specialized

feeding requirements and may be fed like domestic

equines. Specifically, feeding a diet combining high-fiber

pellets and grass hay to nonruminant grazers in zoos is

recommended. The pellet serves as a source of nutrients

and may be designed to compensate for specific regional

dietary deficiencies or for deficiencies in the hay. In regions

where enteroliths are a problem reducing or eliminating

alfalfa in the pellet or hay source is advisable. Pellet and

hay may be fed at a ratio of approximately 50% pellet and

50% hay. Intake should be about 1.5% to 3.0% body

weight.25 In group-housed animals, adequate feeders are

necessary to avoid competition from dominant animals. Salt

and trace mineral blocks may be used If the pelleted diet

cannot be specially formulated. Feeding of produce is not

recommended since readily fermentable substances may

lead to digestive upset. Often these items are desired for

behavioral husbandry or enrichment. It is recommended

that produce be offered at no more than 2% to 5% of the

diet on a dry matter (DM) basis. 25 The document ‘Nutrient

Requirement for Horses’ from the National Research Council

(NRC) may be consulted for specific nutrient and energy

requirements for maintenance, growth, gestation, and

lactation.14

Obesity may be a problem in nondomestic equids

maintained in zoo environments. Encouraging exercise and

restricting the amount of pelleted feed may help.” (Janssen

& Allen, 2014)

“One of the most significant noninfectious diseases in the

author’s experience is enterolithiasis.20 This disease most

often manifest as acutre onset of colic… Reducing dietary

alfalfa because of its excessive magnesium content may be

useful to prevent enteroltihs.14” (Janssen & Allen, 2014)

“In one group each of Plains zebra (six mares, one foal, one

subadult) and Asiatic wild asses (seven mares, two foals) at

Nuremberg Zoo, food distribution was experimentally

changed from clumped (all food in one standard hay rack)

to dispersed (one heap per animal). Plains zebras had an

individually structured system of social relationships in a

dominance order. During the dispersed feeding situation

frequencies of agonistic interactions in both species

decreased (however nonsignificantly), individual distances

increased but mares also frequently ''visited'' each others:

heaps.” (Ganslosser & Dellert 1997)

The presence of forage ad libitum eliminated coprophagy

and greatly reduced the amount of pacing observed.”

(Boyd, 1988)

“Laminitis resulting from consumption of carbohydrate rich

feed; occurs rarely in Przewalski’s horse. Causes lameness

and can be prevented by avoiding rich pasture and carbo-

hydrate rich feeds” (Janssen & Allen, 2014)

“Colic arising from intestinal accidents, sand impactions or

enterocolitis appears to be occasional problem resulting in

abdominal pain, distress, recumbency and shock and can

be prevented by managemental adaptations (sand

impactions” (Janssen & Allen, 2014)

“Enteroliths caused by diet of alfalfa hay (struvite stones)

which is predominantly reported in kiangs, somali wild ass

and appears to be a regional problem. Causes acute severe

abdominal discomfort and requires rapid surgical

intervention to prevent fatal outcome. Prevention by dietary

changes” (Janssen & Allen, 2014)

“Red maple leaf toxicity (gallic acid in dry or wilted leaves

has been reported in Grevy zebras resulting in

methemoglobinemia, hemolytic anemia. Can be prevented

by not planting Acer spp. Tree in or around housing areas.”

(Janssen & Allen, 2014)

“Equine degenerative myeloencephalopathy caused by

vitamin E deficiency reported in Przewalski’s horses, zebras

and kulans resulting in hind limb incoordination, particularly

in foals and can be prevented by oral vitamin E

supplementation” (Janssen & Allen, 2014)


Te hoog energiegehalte van het voer (overmatige krachtvoer) > Obesitas en hyperlipemie, hoefbevangenheid, witte

lijn defecten

Onvoldoende foerageren en te weinig ruwvoer > maagzweren, orale stereotypieën, coprofagie, pacing, tandproblemen

(diersoorten zijn daarvoor lastig te behandelen)

Imbalans: Vitamine E deficiënties (mn bij veulens en bij voeren van uitsluitend ruwvoer; personal communication FV)

> Myopathie; goed te voorkomen wanneer vitamine E of multivitamine/mineralen supplement bij ruwvoer (of goede brok

speciaal voor deze soorten

Andere problemen met imbalans van voeding: vitamine E-deficientie of gebrek aan weidemogelijkheden- Equine

degenerative myeloencephalopathie (neurologische verschijnselen), Afwezigheid zout/mineralenblock > Deficiënties (mn

calcium wordt genoemd), Opname van alfalfa (magnesiumrijke voeding) > Enterolithen en koliek (allen zeldzaam,

personal communication FV)

Voederplaats: onvoldoende voederplaatsen > (territoriale) agressie, (zeer zelden onvoldoende voeropname en

afvallen; personal communication FV)

Plotselinge voedingsveranderingen > Enteritis en koliek (geldt voor alle hindgut fermenters; zeldzaam; personal

communication FV)

Groter risico op eten van giftige planten vanwege weinig selectief eten (ezels) > intoxicaties, evt met dood tot gevolg

(speelt bij de ezel, daarom niet op argkrt)

Endoparasieten a.g.v. voeren op de grond, icm evt. verhoogde druk in populatie in gevangenschap en

moeilijkheidsgraad om deze dieren te behandelen (komen echter zelden voor; personal communication FV)

Zandkoliek a.g.v. fourageren vanaf de grond, en/of mineralen tekort ( likken aan grond), en/of abnormale gedragingen

(komt relatief zelden voor; als koliek dan wel vaak hierdoor of door endoparasieten maar bij dergelijke paardachtigen in

verhouding tot het paard zelden koliek; personal communication FV – mn indien voeren op een volledig kale weide waar

geen gras meer staat maar alleen zand ligt)

Voeren in een ruif > Luchtwegproblemen (inhalatie stofdeeltjes), nek/rugproblemen, entrapment/trauma (geldt voor

paardachtigen algemeen) (probleem wordt nauwelijks gezien bij overige equiden anders dan gedomesticeerde paard;

personal communication FV)

Reproductie (voortplanting en grootbrengen van jongen)

In de natuur:

“Equids are polyestrus species whose reproductive season

is not narrowly defined. In the wild most foals are born in

early to late spring.” (Walzer, 2003)

“Females come into heat several times a year or until they

become pregnant. Most species give birth every 2 years

to a single offspring after a gestation period of 11-13

months. Weaning occurs after about 6-8 months and

offspring become sexually mature at about 2 years.

Potential lifespan is 25-35 years.” (Ballenger & Myers

2001)

“The Przewalski's horse has 66 chromosomes, compared

to 64 in all other horse species.” (Wikipedia [Engels]:

Przewalski’s horse)

“Weaning is complete after 7 to 11 months but females

may lactate up to 16 months. Young reach independence

after 1 to 3 years, when they leave their natal groups.”

(Colvin & Nihranz 2009; E. burchelli)

“Both male and female offspring disperse from the natal

group” (Gale, 2004)

“Plains zebra: The young disperse voluntarily from the

group when aged between one and three years, with the

males joining bachelor groups until able to compete at

around four years old

Females that have come into first oestrus advertise

readiness to breed by adopting a distinctive stance, with

head lowered, tail raised and straddled legs. All males in

the area compete fiercely for such females, and may

abduct them from harems prior to voluntary dispersal.”

(ARKIVE)

In de houderij:

“Although reproductive problems (e.g. cystic ovaries,

endometritis) have been reported, breeding generally is not

a problem in these species.” (Walzer, 2003)

“Immunocontraception with porcine zona pellucida (PZP)

has been used extensively in feral horses in the United

States and in Przewalski’s horses in Europe… Whereas no

negative effects of PZP have been noted in short-term use

(1-3 years) these may not be generally excluded at the

present.” (Walker, 2003)

“Infanticide occurs in captive E. kiang (Berger 1986). (…)

Infanticide has often been observed in other equid species

(Cameron et al. 2003; Duncan 1982; Linklater et al. 1999).

Induced abortion has been observed in wild horses (Berger

1983), but whether it occurs in E. kiang is unknown.” (St-

Louis & Côté 2009, p.5)

“On 3 March 1965 a female foal was born at Ahmedabad

Zoo. .... When the foal was born the stallion was in the

same enclosure and to our consternation he vigorously

rushed at the new-born foal and attacked it, picking it up

by the neck and trying to kill it.. [p. ] (David 2007 Indian

wildass)

“Het speenproces bij ezels zal gewoonlijk vanzelf

plaatsvinden tussen de 7 en 12 maanden. Bij wilde paarden

is de gemiddelde speenleeftijd 8-9 maanden en hoewel

deze leeftijden elkaar overlappen, lijkt een ezelveulen

veelal wat langer bij de merrie te blijven dan een

paardenveulen.” (de Graaf-Roelfsema 2009, p.47)

In de dierentuin kunnen halfezels zonder grote

moeilijkheden worden gefokt. De dieren wennen er snel aan

hun nieuwe omgeving; de jongen zijn tamelijk goed

bestand tegen ons wisselvallig klimaat en wennen zonder

A rival male may attempt to drive off the family stallion. If

the stallion leaves the family, his sons may follow him.

Young males normally leave the family at 1 to 3 years of

age, and exceptionally up to 4.5 years. (Grubb, 1981)

Bonds of the foal with the mother are lost when she has a

new foal and as opportunities for play-fights with other

males are greater in stallion groups. Family stallions

nevertheless do not show antagonism to their sons.

(Grubb, 1981)

Mature adult females only exceptionally leave families.

(Grubb, 1981)

As females mature, the estrous stance becomes less

conspicuous and confined to the period immediately

before mounting, so without the optically conspicuous

stance, there is no competition for mature mares. (Grubb,

1981)

“A rival male may attempt to drive off the family stallion.

If the stallion leaves the family, his sons may follow him.

Young males normally leave the family at 1 to 3 years of

age, and exceptionally up to 4.5 years.” (Grubb 1981)

“Stallions offer parental care by defending the group from

predators.” (Colvin & Nihranz 2009; E. burchelli)

Harems are organized into a dominance hierarchy.

Females of higher rank have been found to produce more

offspring and to have shorter interbirth intervals. Stallions

show mating preference towards these high ranking

females. Immediately after birth, foals take a position in

the dominance hierarchy at a position below their

mothers. (Colvin & Nihranz 2009; E. burchelli)

“When preparing to give birth, mares separate from the

rest of the herd to hide from predators.” (Colvin & Nihranz

2009; E. burchelli)

“Male infanticide and feticide has been noted in captive

individuals of Equus burchellii and occur when a new male

takes over a harem. In order to gain reproductive

advantage the new stallion will kill young foals

(infanticide) or force them to abort (feticide) via forced

copulation. By gaining reproductive rights to a harem, the

stallion is able to ensure that he is only investing parental

care in his own offspring.” (Colvin & Nihranz 2009; E.

burchelli)

Harassment involving chases and copulations can

negatively affect a female’s feeding rate and may even

result in abortion or involve infanticide. (Grzimek 2004)

“The male sniffs the urine or feces of estrous females and

then may show flehmen, followed by urinating or

defecating on the deposits of the female. Precopulatory

grooming of young mares occurs, with copulation only on

the third or fourth day of estrus.” (Grubb 1981)

moeite aan ander voer. Alleen de hengst veroorzaakt door

zijn aggressiviteit vaak moeilijkheden in een kudde

fokdieren. Hij jaagt de merries op en bijt ze in de poten en

vooral in de keel. [deel xii, p. 656, kol. 2] ... In de

dierentuin verliezen halfezels nooit hun aangeboren

schuwheid en ze kunnen nooit zo afgericht worden, dat ze

als last-, rij- of trekdier bruikbaar zijn. [p.657] (Grizimek

1973)

“Sexual maturity occurs somewhat later in most wild equids

than it does in domestic horses, especially the male; some

do not reach maturity until the age of five years. This is

most often seen in the Przewalski’s wild horse. This species

also has a high percentage of infertile males.

Estrus in wild mares is similar to that in domestic mares. It

usually starts in early spring and, if not settled, may go to

late summer. Births are recorded from April through

September. Gestation periods are not well documented, but

the accepted range is 330 to 350 days for most species.

Grevy’s zebra has a longer period of gestation.

Whether or not to leave the male in the herd when the foals

are dropped depends mostly on size and type of enclosure

and disposition of the individuals, both stallion and mare.

Biting and kicking can be extremely viscious and

occasionally foals are injured or killed in the melee,

especially when mares are in foal estrus

Most wild equine mares are very protective mothers.

Most foals hold a strong attachment for their mothers for a

long time. Nursing will continue for four to six months or

even longer if the mare will permit. Often the mare must

forcibly drive the youngster away when her next foal is

born” (Nelson, 1986)

“Several cases of infanticide have been documented in wild

equids.16 Care should be exercised when introducing a

male to a new herd or a pregnant female to a new stallion.

Stallions may be aggressive toward new foals, so mares

that are close to foaling should be separated from

stallions.” (Janssen & Allen, 2014)

“As all equids have a similar reproductive biology, the

domestic horse is a good model. Equids are seasonally

polyestrous, with estrous behavior recurring until

conception or the end of the breeding season. In temperate

regions, seasonality is determined primarily by photoperiod.

In tropical species, seasonal birth peaks relate to other

environmental factors such as the rainy season.” (Janssen

& Allen, 2014)

“The gestation length of Przewalski's horses is 11 months

and foals are usually born in the late spring or early

summer (Dobroruka, 1961; Mohr, 1971 ). Mares come into

estrus within 1 week of foaling. The mares in this study

who conceived promptly weaned their foals just prior to the

birth of their next offspring. Mares failing to conceive during

that breeding season, permitted their current foals to nurse

“Equus hemionus are monogamous. Stallions tend to stay

with the mare and foal year-round. (Feh et al, 2001)”

(Reuter 2002)

“The newborn frees itself from the amniotic membranes

and crawls towards its mother’s head. The mother licks it

clean, and ingests the membranes and some amniotic

fluid, which may be important in initiating lactation or the

maternal bond (King, 1965; King et al. 1966).” (Churcher

1993, p.4)

“The young are semi-independent at ca. 7 months, often

being left by mother or leaving her for periods of hours.

(…) It is weaned by 9 months (32-42 weeks – King, 1965)

but stays with hits mother. (…) A young Grévy female

becomes independent from her mother at 13-18 months

(…). A young Grévy male stays with his mother until at

least 3 years old, when he joins a bachelor group or a

mixed herd.” (Churcher 1993, p.4)

The single young weighs about 32 kg, can stand almost

immediately, runs within an hour and starts to eat grass

within a week. However, weaning usually is completed

after 7-11 months, and lactation sometimes continues for

up to 16 months. The young normally leave their family

after 1 - 3 years (Nowak 1991)

There is little or no precopulatory activity with adult

mares, which are mounted repeatedly during the whole of

estrus, but with intromission every one to three hours

only at the height of estrus. (Grub 1981)

SEXUAL MATURITY: female at 1.25 years. first fully

sexually capable at 2.5 years, Male at I.25 years, fully

capable of breeding at 2.5-3 years and first capable of

being a family stallion at 5-7 years. Female fertile up to

20 years, producing on average 1 young per year

(Haltenorth & Diller, 1988)

Female in oestrous allows close following by male, who

smells, licks and grooms her. Female at height of ostrous

is first to be ready. Mating lasts only a few seconds, Male

lays head on female withers. Mating takes place

repeatedly at intervals of 1-2 hours throughout 1-2 days.

(Haltenorth & Diller, 1988)

Foals are especially vulnerable with 50% of juveniles

annually dieing due to predation. This high rate of juvenile

mortality is also partly due to disease, death of mothers,

low nutrition, and drought. (Colvin 2009)

Throughout the species range, mortality rate can vary in

the first year, from 19–47%. Adult mortality is lower and

varies from 3–17%. (Grizimek 2004)

much longer. Three mares whose 2-year-old daughters

remained in their natal band failed to conceive during the 2

years following the birth of their daughters. Each of these

mares nursed their daughters through 26 months of age.

Mares continue investment in lactation for their older

offspring until they have another offspring in utero which

begins to make significant demands on their resources. The

timing of weaning of older foals can, thus, be used as a

non-invasive diagnosis for pregnancy, with the exception of

mares on a high plane of nutrition. Mares at two zoos

received 17% more digestible energy than horses at the

zoo feeding the next highest energy level, and nursed both

their yearlings and their latest foals. Klimov (1988) has also

reported that mares may allow older offspring to continue

to nurse along with their newborn foals. The high level of

nutrition apparently permitted the mares to invest in more

than 1 offspring simultaneously.” (Boyd, 1991)

“In the wild, sires probably played an important role in

protecting their offspring against predators and in retrieving

foals when they strayed from the herd, as has been

observed in feral horses (Feist and McCullough, 1975;

Boyd, 1980; Berger and Rudman, 1985; Berger, 1986 ). In

captivity, sires rarely have occasion to provide this form of

parental investment. When the number of feeding stations

are limited, sires do appear to aid their offspring by sharing

grain with them. Mackler and Dolan (1980) also observed a

Przewalski's stallion sharing food with his son but chasing

adult mares away. Stallions also tolerate foals grazing in

their vicinity (Dobroruka, 1961 ). (Boyd, 1991)

“Several cases of infanticide by Przewalski's stallions have

been reported…. Infanticide may not be a pathological

behavior resulting from captivity, as it is also seen in free-

ranging domestic horses and appears to be adaptive as it is

directed toward unrelated foals rather than the sire's own

offspring. By killing unrelated foals, a stallion eliminates

future competitors who do not share his genes. In domestic

horses, unrelated male foals are more commonly victimized

than unrelated female foals, perhaps because male foals

grow up to be competitors whereas female foals represent

future mates (Duncan, 1982 ). By killing young unrelated

foals, a stallion might also benefit by freeing the mares

from the physiological stress of lactation, which improves

their chances of producing a foal by him the following

season (Boyd, 1986)… Infanticide does not always occur

when 1 stallion replaces another. Several stallions did not

attack unrelated foals born into their harem after they

replaced the former stallion. As zoos cannot predict

whether a stallion will be infanticidal, precautionary

measures may be warranted. Introducing stallions only to

harems containing non-pregnant mares and older foals,

who are less likely to be attacked, should help.

Alternatively, mares pregnant by other stallions could be

isolated prior to foaling, although this disrupts the normal

social structure of the herd. Zoos must strike a balance

between the need to rotate stallions frequently in order to

promote gene flow in inbred populations and the increased

risk of infanticide that this practice brings“ (Boyd, 1991)

“The sexual selection hypothesis explains infanticide by

males in many mammals. In our 11-year study, we

investigated this hypothesis in a herd of Przewalski's horses

where we had witnessed infanticidal attacks. Infanticide

was highly conditional and not simply linked to takeovers.

Attacks occurred in only five of 39 cases following a

takeover, and DNA paternity revealed that, although

infanticidal stallions were not the genetic fathers in four

cases out of five, stallions present at birth did not

significantly attempt to kill unrelated foals. Infanticide did

not reduce birth intervals; only in one case out of five was

the infanticidal stallion, the father of the next foal; mothers

whose foals were attacked subsequently avoided

associating with infanticidal stallions. Therefore, evidence

for the sexual selection hypothesis was weak. The “human

disturbance” hypothesis received some support, as only zoo

bred stallions which grew up in unnatural social groups

attacked foals of mares which were pregnant during

takeovers” (Feh & Munkhtuya, 2008)

“On the assumption that infanticide exists in plains zebra,

as reported for horses, Equus caballus, we tested the

following hypothesis. Introducing a new zebra male into a

herd of breeding females should increase foal mortality in

comparison with herds in which the sire of the foals is still

present. The younger the foal, the more likely infanticide

should be. We collected data from five herds in two

zoological gardens in the Czech Republic. We found nine

records of infanticide in plains zebra and three cases of

abortions that were probably induced by forced copulation.

We analysed additional indirect data to investigate the

possibility of introduced males causing other abortions.

Abortions were three times more likely in herds with

introduced males than with only fathers present. Postnatal

mortality of the foals was four times greater with

introduced males than with fathers. No indication of a sex

preference was observed for infanticide by a new male for

either abortions or postpartum deaths. When we combined

all records involving introduced males, the probability of

foal death was greatest when the new male joined the herd

just after conception and decreased with increasing time

between conception and date of the new male introduction

(the chance of a foal surviving was less than 5% just after

conception and more than 50% at the time of delivery).

Mortality of foals did not depend on whether the new male

was introduced before or after the foal was born. Survival

increased to more than 60% after the foal reached 1 month

of age. Our results suggest that captive plains zebra show

the highest occurrence of infanticide reported among

ungulates.” (Pluhacek & Bartos, 2000)


Introductie van een nieuwe hengst in een groep met veulens of drachtige merries (geeft ook verstoring van de

groep/stress) > Infanticide, abortus(Mannetje hiervoor zoveel mogelijk uit de groep plaatsen, rust behouden in groep).

Wisseling van hengst is nodig om inteelt te voorkomen. Verhoogd risico op agressie naar merrie en veulen ook vaak

a.g.v. opgroeien in onnatuurlijke condities in gevangenschap (menselijke verstoringsfactor)

Gevaar voor de mens (Inzetbaarheid van afweer- en aanvalsmiddelen)

“… among their own kind or in attempting defense, they

kick with the hind feet, strike with the forefeet and

sometimes bite.” (Ballenger & Myers 2001)

“Aggressive behavior can be biting, kicking, circling and

displacement, but the most common response to a

competitor is a threat to kick or bite.” (McGreevy 2004,

p.125)

“Both subspecies of E. zebra are good climbers and have

exceptionally hard and pointed hooves compared to other

equines.” (Walker 2005; pointed hooves…)

“All equids are relatively swift, alert runners and generally

flee from danger rather than fight. However, among their

own kind or in attempting defense, they kick with the hind

feet, strike with the forefeet and sometimes bite.”

(Ballenger & Myers 2001)

“In fighting among themselves or in attempting a

defense, they kick with the hind feet, occasionally strike

with the forefeet, and sometimes bite. Their teeth, though

not adapted for lacerating or tearing (except in male

zebras, which have pointed canines), can give quite hard

pinches.” (Nowak & Walker 1991, p.1306)

Ook dieren die van jongs af in gevangenschap leven

verliezen hun wildheid niet. De kainghengst in de dierentuin

van Praag, die in gevangenschap is geboren, valt

blindelings iedereen aan die in debuurt van zijn hek komt;

[vol XII, p660, kol. 1] (Grzimek 1973)

“By the time that they reach maturity, most individuals are

unpredictable, at best, and many are actually dangerous.

They are best kept in unites by themselves. The males and,

even at times, the females will traumatize other animals.

Biting, especially tail biting, is common, and they have

frequently used their front feet to stomp a newborn

antelope to death. Zebra mares have been known to

attempt killing another mare’s newborn foal. Hartmann’s

mountain zebras (Equus zebra hartmani) are more tolerant

than other species but are usually aggressive enough to

drive any other species away from feeders.” (Nelson, 1986)

“The zebras were the more challenging species as they

tend to be easily agitated; this was particularly evident in

the three individuals being introduced. Zebras may work

themselves into a veritable frenzy once this state is

triggered, and they are often injured in the process.”

(Kleiman et al. 1996, p.207; over opzetten van mixed-

species habitats en introducties van verschillende soorten;

in dit geval zebra’s en neushoorns)

“… tranquilization without use of an immobilizing agent is

dangerous to personnel involved, because of the

unpredictable behavior of these animals”

“Danger potential: Members of this group (horse, ass,

zebra) are efficient and effective kickers, strikers and

biters. They should not be considered as just slightly wild

horeses, because they lash out much faster than the

domestic horse and are far more likely to kick or strike.

Physical restraint: There is no effective way to physically

restrain adult animals of this category without risk of injury

to the animal or handlers. The twitch, lip chain or other

similar devices customarily employed when restraining

domestic equids are not suitable for use on wild equids.”

(Fowler, 1995)

“Stallions may also be aggressive toward keepers, and

extreme caution should be used when working around

them. 37” (Janssen & Allen, 2014)

Skiff (1982) studied the effects of enclosure size on the

behavior of 2 herds of Przewalski's horses. The number of

activities was higher in the 0.05 ha enclosure than in the

3.5 ha enclosure. When in the small enclosure, horses did

more milling and fence-pacing, had higher rates of

aggression, and spent more time mutually grooming than

when on pasture.=> N.B. Betreft hier wel reactie naar

andere dieren (Bouman-Heinsdijk, 1982; Boyd, 1986).

The “human disturbance” hypothesis received some

support, as only zoo bred stallions which grew up in

unnatural social groups attacked foals of mares which were

pregnant during takeovers” (Feh & Munkhtuya, 2008)

This review elucidates the strong need for specific research

and training of humans working with horses in order to

improve the human–horse relationship that, as shown by

the high incidence of accidents and increasing number of

horses with a decreased welfare is far from optimal…..They

have to be aware that deficits in the management

conditions (housing, food, social context, and training) may

lead to disturbances in the horse’ behaviour and ultimately

to relational problems. (Hausberger 2008; equus callabus)

Ernstig letsel inclusief inwendig trauma (vergelijkbaar met het paard) maar risicio hoger dan bij paard doordat ze

sneller opgewonden zijn

Extra risico (offensieve agressie) bij moeder met jong en/of hengsten

Extra risico doordat ze minder snel gewend raken aan prikkels en hun schuwheid moeilijker verliezen indien niet

gehabitueerd op jonge leeftijd

Extra risico op agressie in gevangenschap bij beperkte ruimte en vluchtmogelijkheden, onnatuurlijk opgroeien,

suboptimaal management

Literatuurlijst Equus

Bronnencategorie

Geciteerde literatuur Diersoort

Wete

nschappelijk

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tijd

schri

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Vakbla

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Mondelingen

mededelingen

Overi

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1 Wikipedia (Engels) X

2 Wikipedia (Nederlands) X

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Niet gebruikt in excerpt (wel quotes in Statements file): AL AnimalsWorld

Accessed December 10, at http://animalia-life.com/horse.html

AnAge

AnAge: The Animal Ageing and Longevity Database. Accessed December 10, 2014 at http://genomics.senescence.info/species/

equus (hippotigris) quagga (burchelli)

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