Equus (Hippotigris) quagga (burchelli)
Transcript of 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
Schofthoogte: 120 – 140 cm
Gewicht: 200 – 340 kg
E. grevyi
Schofthoogte: 125 – 160 cm
Gewicht: 350 – 450 kg
“Equus zebra is the largest living zebra” (Churcher 1993, p.1)
E. hemionus ssp.
Schofthoogte: 100 – 142 cm
Gewicht: 200 – 260 kg
E. kiang
Schofthoogte: 142 cm
Gewicht: 250 – 440 kg
“Equus kiang is the largest of the wild asses” (St-Louis & Côté 2009, p.2)
E. zebra
Schofthoogte: 116 – 150 cm
Gewicht: 240 – 372 kg
“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)
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 cap- tive 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.”
(McGreevy 2004, p.134)
“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)
Oorzaak-gevolg risico’s:
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 &
Nihranz 2009; E. burchelli)
“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)
Oorzaak-gevolg risico’s:
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 &
Nihranz 2009; E. burchelli)
“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)
Oorzaak-gevolg risico’s:
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)
Oorzaak-gevolg risico’s:
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.”
(McGreevy 2004, p.42)
“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)
“SUMMARY OF KEY POINTS
• 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)
Oorzaak-gevolg risico’s:
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
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,
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
adapted to desert conditions (Geptner, 1961); that is why
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)
“SUMMARY OF KEY POINTS
• 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)
Oorzaak-gevolg risico’s:
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)
Oorzaak-gevolg risico’s:
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
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tijd
schri
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Vakbla
den
Mondelingen
mededelingen
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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/