ChapterPDF Available

The Red List of Mammals of South Africa, Lesotho and Swaziland Equus quagga | 1

The Red List of Mammals of South Africa, Lesotho and Swaziland Equus quagga | 1
Taxonomy
Equus quagga (Boddaert 1785)
ANIMALIA - CHORDATA - MAMMALIA -
PERISSODACTYLA - EQUIDAE - Equus - quagga
Synonyms: Equus burchelli (Gray 1824) [orth. error]
Common names: Plains Zebra, Burchell’s Zebra,
Common Zebra, Painted Zebra (English), Bontsebra
(Afrikaans), Idube (Ndebele, Zulu), Iqwarhashe (Xhosa)
Taxonomic status: Species
Taxonomic notes: Groves and Bell (2004) described six
subspecies of Equus quagga based on craniometric
taxonomic methods, which showed all subspecies to be
alike, and distinctly different from Mountain Zebras (Equus
zebra). The six subspecies differ from one another in
morphological characteristics, such as coat patterns and
colouration, body size, and the presence (or absence) of a
mane (Groves & Bell 2004). Although there is limited
evidence for genetic diversity between subspecies and the
classification may be a contested topic (Bennett 1980), a
classification of subspecies can be useful as a way to
Equus quagga Plains Zebra
Regional Red List status (2016) Least Concern
National Red List status (2004) Least Concern
Reasons for change No change
Global Red List status (2016) Near Threatened
A2a+3c+4ac
TOPS listing (NEMBA) None
CITES listing None
Endemic No
Recommended citation: Stears K, Shrader A, Castley G. 2016. A conservation assessment of Equus quagga. In Child MF,
Roxburgh L, Do Linh San E, Raimondo D, Davies-Mostert HT, editors. The Red List of Mammals of South Africa, Swaziland
and Lesotho. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa.
E. Do Linh San
emphasise ecological and morphological differences for
management purposes. Following Groves and Bell (2004),
six morphologically defined subspecies are listed as:
Equus q. crawshaii (Crawshay’s Zebra) occurs in
east Zambia, east of Luangwa River, Malawi, south-
eastern Tanzania, and Mozambique as far south as
the Gorongoza district.
Equus q. borensis (Maneless Zebra) ranges over the
northern parts of east Africa from north-west Kenya
to the Karamoja district of Uganda and south-east
Sudan, east of the Nile River (Boma National Park) to
the northern limit of the species’ range (32 °N).
Equus q. boehmi (Grant’s Zebra or Boehm’s Zebra)
is found in Zambia, west of the Luangwa River, west
to Kariba, the Shaba Province of the Democratic
Republic of the Congo north to Kibanzao Plateau,
Tanzania north from Nyangaui abnd Kibwezi into
south-west Uganda, south-west Kenya, and east
Kenya, east of the Rift Valley, into southern Ethiopia
and perhaps to the Juba River in Somalia. This
subspecies constitutes of over 75% of Africa’s Plain
Zebra population, and the Serengeti-Mara
ecosystem supports approximately 200,000
individuals (East 1997; Hack et al. 2002).
Equus q. chapmani (Chapman’s Zebra) ranges from
north-east South Africa, from about 24 °S, 31 °E,
north to Zimbabwe, west into Botswana at about
19 °S, 24 °E, across the Caprivi Strip in Namibia, and
into southern Angola.
Equus q. burchellii (Burchell’s Zebra) formally
occurred north of the Vaal/Orange Rivers, extending
north-west via Botswana to Etosha National Park
and the Kaokoveld, south-east to KwaZulu-Natal and
Swaziland. It is now extinct in the middle of its range.
Equus. q. antiquorum is now included in this
subspecies.
Equus q. quagga (Cape Quagga) occurred in the
former Cape Province, south of the Orange and Vaal
Rivers and west of the Drakensberg. This
subspecies has been extinct since 1883.
Research indicates that the extinct E. q. quagga might
differ morphologically from other Plains Zebra subspecies,
but those differences are not sufficient to justify its
classification as an independent species. Thus the extinct
E. q. quagga should be considered a subspecies of
E. quagga. From north to south of their geographic range,
extant Plains Zebras show a gradual decline in the
contrast between dark stripes and white interspaces, and
an increase in body size is noted from east to south
(Lorenzen et al. 2008). The extinct Cape Quagga, which
was found at the southern limit of the species’ geographic
range, exhibited the extreme case of this trend in pelage
colouration (Groves & Bell 2004; Lorenzen et al. 2008).
Assessment Rationale
Plains Zebras are listed as Least Concern because they
are widespread and abundant throughout the assessment
A recent study in Namibia and Botswana
described a formerly unknown migration of Plains
Zebra from the Chobe River to Nxai Pan National
Park. This migration is now suggested to be the
longest of all documented migrations
by large mammals in Africa
(Naidoo et al. 2016).
Equus quagga | 2 The Red List of Mammals of South Africa, Lesotho and Swaziland
Figure 1. Distribution records for Plains Zebra (Equus quagga) within the assessment region
region and, at present, there are no major threats affecting
the population. Within South African National Parks alone,
there are an estimated 31,94846,237 individuals (counts
conducted from 20102012), with Kruger National Park
(KNP) containing the bulk of the population (29,161
43,450 individuals). The expansion of wildlife ranches
across the country is also increasing the abundance of
this species. The only real threat to this species is the
fragmentation of its habitat, being restricted to fenced
areas, which increases its risk to drought and other
disturbances. Continued protected area expansion
through transfrontier conservation areas, conservancies
and stewardship schemes should be employed to
enhance free movement of Plains Zebra in response to
climatic variability, and changes in the availability of forage
and water.
Regional population effects: Within the assessment
region, migration does take place within transfrontier
Country Presence Origin
Botswana Extant Native
Lesotho Extinct Native
Mozambique Extant Native
Namibia Extant Native
South Africa Extant Native
Swaziland Extant Native
Zimbabwe Extant Native
parks, such as KNP and the Greater Mapungubwe
Transfrontier Conservation Area (TFCA), however the
remaining population is typically restricted to private,
provincial and national protected areas. A degree of
genetic dispersal occurs through the translocation of
individuals between these protected areas via game sales.
No rescue effects are necessary.
Distribution
This species ranges from the northern region of South
Africa northwards to southern Sudan and Ethiopia, and
westwards into northern Namibia and southern Angola
(Hack et al. 2002). Historically, Plains Zebra exhibited a
much broader range across the African continent, and
fossil remains have been discovered from North Africa
(Churcher & Richardson 1978). More recently, Plains
Zebra ranged throughout eastern and southern Africa,
with only the exception of the forest belts of Uganda
(Duncan 1992), and extended southwards across the
Orange and Vaal Rivers in South Africa to the Cape, with
the subspecies E. q. quagga, which is now extinct (Groves
& Bell 2004). The species is also now extinct in Burundi
and Lesotho, and there remains a question over their
presence in Angola.
Within the assessment region, the natural range of the
Plains Zebra is found in the eastern Lowveld and the
KwaZulu-Natal lowlands, and into the Northern Cape
(Figure 1). Subpopulations have been widely reintroduced
across their natural range, for example, in North West
Province where they once occurred throughout the
province (du Plessis 1969; Rautenbach 1982; Power
2014). A migratory subpopulation exists in the KNP;
Table 1. Countries of occurrence within southern Africa
The Red List of Mammals of South Africa, Lesotho and Swaziland Equus quagga | 3
however, most other populations do not have sufficient
area in which to migrate seasonally. Subpopulations of the
species have been introduced into areas outside of its
natural range (Figure 1). The Greater Mapungubwe
Transfrontier Conservation Area (between South Africa,
Botswana and Zimbabwe) also supports a subpopulation,
and approximately 300 individuals are within the Botswana
section.
Population
The Plains Zebra is common, widespread and amongst
the most abundant of all grazing mammals in Africa, with a
total population size of 663,212 individuals estimated in
2002 (Hack et al. 2002). Recent information regarding
global overall population numbers is limited. However,
during the previous decades, this species has been
extirpated from portions of its range (Hack et al. 2002).
Within the assessment region, the population is
increasing, both on formally protected areas and the
expansion of private wildlife ranches. For example, the
subpopulation on Golden Gates Highlands National Park,
Free State Province, has increased from 131 to 1,592
individuals between 1994 and 2016 (Bissett et al. 2016).
Similarly, on Chelmsford Nature Reserve, KwaZulu-Natal
Province, the Plains Zebra population has increased from
97 to 876 individuals between 1986 and 2014 (KwaZulu-
Natal Ezemvelo Wildlife unpubl. data). The generation
length of the Plains Zebra has been calculated as 10 years
(Pacifici et al. 2013), which makes the three generation
window 30 years (19862016). While few long-term
datasets area available to quantify the overall national
population increase, nearly all protected areas show an
increasing or stable trend.
Within South African National Parks, there are an
estimated 31,94846,237 (seven protected areas; 2010
2012 counts) individuals (Ferreira et al. 2013), with KNP
containing the bulk of the population (29,16143,450
individuals, estimated using distance sampling in 2012).
Overall, there is a minimum estimate of 59,204 individuals
on 803 protected areas and wildlife ranches across the
country (20102015 counts; Endangered Wildlife Trust
unpubl. data), which equates to 35,52241,443 mature
individuals assuming a 6070% mature population
structure. Thus, the wild population is well established and
appears to be increasing. There are three main groups:
1. Kruger National Park and the Associated Private
Nature Reserves: The KNP is estimated to contain
around 5478% of the total the free-roaming
population.
2. Northern KwaZulu-Natal: The Plains Zebra
population was estimated to comprise 12,166
individuals. However, these data do not represent the
entire population within the province as they are
limited to 22 Ezemvelo-KZN Wildlife protected areas
and 31 private reserves. As a result, this value better
represents a minimum population estimate.
3. Limpopo, North West, and Northern Cape
provinces: Large numbers in private populations
through the region. The North West alone, for
example, contains 4,450 individuals on provincial
reserves and 8,920 individuals on private properties
(Power 2014).
Population density ranges from estimates of 22 animals /
km² in Ngorongoro (Klingel 1969), 16.6 animals / km² in
Nechisar National Park, Ethiopia (Doku et al. 2007), to
0.9 animals / km² in KNP (Smuts 1976). Densities are also
expected to vary seasonally in migratory regions, as not
all individuals migrate (Hack et al. 2012).
Current population trend: Increasing
Continuing decline in mature individuals: No
Number of mature individuals in population: 35,522
41,443
Number of mature individuals in largest subpopulation:
17,49730,415 animals in KNP.
Number of subpopulations: At least 97 in formally
protected areas.
Severely fragmented: Yes, with the exception of Greater
Limpopo Transfrontier Park and Mapungubwe
Transfrontier Conservation Area, all reserves are fenced,
and do not allow for natural movements and migrations.
Translocations of individuals do contribute towards gene
flow.
Habitats and Ecology
With the exception of a few extreme environments, such
as deserts, dune forests and rain forests, Plains Zebra
thrive across a broad range of habitats in Africa within
both tropical and temperate climates (Duncan 1992; Hack
et al. 2002; Stuart & Stuart 2007). From sea level to
elevations of 4,300 m on Mount Kenya, Plains Zebra utilise
open grasslands, as well as savanna woodland (Duncan
1992). In southern Ethiopia, Plains Zebra revealed a
distinct preference for open grassland habitats, and
habitat preference showed seasonal variability (Regassa &
Yirga 2013). The dietary flexibility and its tolerance for
highly fibrous grass material are the factors considered to
contribute to the Plains Zebra’s broad geographic range
(Schulz & Kaiser 2013).
Sward height largely affects Plains Zebra feeding patterns
and, in northern Botswana, once grass height had
declined to 20 cm, they moved off in pursuit of taller
grasses (Joos-Vandewalle 2000). In the Hluhluwe-iMfolozi
Park Zebra foraged on a range of grasses, but again, in
grazing lawns of Digitaria argyograpta, Urochloa
mosambicensis and Sporobolus nitens, they limited their
grazing to grass swards with a height of at least 20 cm
(Arsenault & Owen-Smith 2008). Within medium-tall bunch-
grasslands comprised of Themeda triandra, Cymbopogon
plurinodis and Heteropogon contortus, they selected for
grass swards at least 10 cm tall (Arsenault & Owen-Smith
2008). The availability of water directly influences their
movements and habitat selection (Estes 1991; Duncan
1992; Skinner & Chimimba 2005). A migratory
subpopulation in Tanzania congregates in the Serengeti
Plains during the rainy season (November to May), but
migrates north-westwards during the dry season, and into
Kenya’s Maasai-Mara National Reserve (Hack et al. 2002).
A recent study in Namibia and Botswana described a
formerly unknown migration of Equus quagga from the
Chobe River (border of Botswana and Namibia) to Nxai
Pan National Park (Botswana) (Naidoo et al. 2016). The
round-trip migration covers a straight-line distance of
500 km, which is longer than the seasonal migratory
movement of Blue Wildebeest (Connochaetes taurinus)
from the Serengeti to the Maasai-Mara in East Africa
(Naidoo et al. 2016). In fact, Naidoo et al. (2016) suggest
that this is the longest of all documented migrations by
large mammals in Africa. Within the assessment region,
fences surrounding protected areas predominantly inhibit
Equus quagga | 4 The Red List of Mammals of South Africa, Lesotho and Swaziland
the migration of the Plains Zebra, with the exception of
Greater Limpopo Transfrontier Park and Mapungubwe
Transfrontier Conservation Area between Botswana, South
Africa and Zimbabwe.
Plains Zebra are gregarious and highly social, forming
herds comprised of one dominant stallion, several females
and their offspring (Klingel 1969). Males without harems
form unstable bachelor herds of up to 50 individuals or
remain solitary (Klingel 1969; Hack et al. 2002). Breeding
in the Plains Zebra is not considered seasonal, as young
are born throughout the year (Klingel 1969), however, a
peak in births occurs between December and January,
and the vast majority of foals are born during the summer
months (Smuts 1976).
Ecosystem and cultural services: As hindgut fermenters,
Plains Zebra process their food at a faster rate than
ruminants. To compensate, they eat more bulk, including
vegetation that is too fibrous and low in protein for most
ruminants to digest effectively. Resultantly, they have the
ability to survive on poor quality vegetation providing it is
abundant. Additionally, Zebra have a low relative muzzle
width index, better suited for foraging in taller grass
(Arsenault & Owen-Smith 2008). By feeding on taller grass
they can facilitate the grazing of other herbivores that
require shorter grazing grass.
Plains Zebra have aided livestock farmers in reducing the
predation pressure of dogs and jackals on livestock
calves, because Zebra chase and deter these predators.
Thus, farmers keep small herds of Zebras on their
properties for this reason (K. Stears, unpublished data).
Finally, as a highly charismatic and extremely visible
species, the Plains Zebra is a powerful attraction for
ecotourism and a valuable species for international
support of wildlife and ecosystem conservation.
Use and Trade
Plains Zebra is popular amongst both subsistence and
commercial hunters for skins, meat and trophies. They
were one of the top three most exported animal trophies in
Namibia in 2012 (Di Minin et al. 2016). Live animals are
frequently traded at game auctions. Trade and hunting of
this species is not expected to have a severely negative
impact on the population. However, the wildlife ranching
industry may be artificially selecting individuals to produce
desired colour morphs and/or hybridising native with
exotic subspecies. Although this is not expected to impact
the wild population, regulations should be established to
prevent such individuals from entering formally protected
areas. A worrying trend is the increasing intensity of
bushmeat hunting in southern Africa (Lindsey et al. 2013),
including some areas within the assessment region
(Hayward 2009; Nel 2015). Presently, however, these
reductions are localised and are unlikely to have an effect
on the overall population.
Threats
Across much of eastern Africa, habitat encroachment by
cattle threatens the forage quality and quantity of Plains
Zebra while illegal hunting can cause local declines (Hofer
et al. 1996; Grange et al. 2004, 2015; Regassa & Yirga
2013). As a result, Regassa and Yira (2013) suggested the
implementation of effective management policies to
combat illegal settlements and encroachment by humans
in the Yabello Wildlife Sanctuary of southern Ethiopia.
Within the assessment region, this species is mostly
restricted to protected areas; therefore, these threats are
unlikely to become a major problem, although localised
problems with poaching do occur in protected areas, such
as in Borakalalo Nature Reserve, North West Province (Nel
2015). Constrained movements do, however, increase the
species’ vulnerability to drought (Walker et al. 1987;
Georgiadis et al. 2003). This is particularly true within
biomes most at risk to the effects of climate change, water
scarcity and enhanced aridity. Subpopulations have
Category Applicable? Rationale Proportion of
total harvest Trend
Subsistence use Yes Bushmeat trade. Unknown Stable
Commercial use Yes Trade in skins and trophies. Non-consumptive ecotourism
on provincial and private reserve.
Unknown Stable
Harvest from wild
population
Yes Live animal sales. Unknown Stable
Harvest from
ranched
population
Yes Meat, skins and trophies. 1015% per
annum (Hack et
al. 2002)
Increasing with
wildlife ranching
expansion.
Harvest from
captive population
Yes Captive breeding for skins. This is rare for this species
because it is mostly focused on Mountain Zebra species,
E. z. zebra and E. z. hartmannae. However, there is an
increasing trend to intensively breed for colour morphs in
captivity.
Unknown Increasing with
wildlife ranching
expansion.
Table 2. Use and trade summary for the Plains Zebra (Equus quagga)
Elsa Bussière
The Red List of Mammals of South Africa, Lesotho and Swaziland Equus quagga | 5
Equus q. antiquarum, previously considered a distinct
subspecies, is currently included under the subspecies E.
q. burchelli (Groves & Bell 2004). Occurring in a few
isolated subpopulations in protected areas of KwaZulu-
Natal, these subpopulations have been considered under
threat from population isolation, and possibly the
detrimental results of inbreeding (Bowland et al. 2001).
Bowland et al. (2001) recognises the importance of
improving management practices and the active
translocation of small, isolated populations between
shown increases and declines associated with high and
low rainfall years, respectively (Gandiwa et al. 2016).
Nonetheless, following population declines associated
with drought years, Plains Zebra subpopulations have
demonstrated fairly rapid recoveries in population
abundance to levels above those prior to the drought
(Hack et al. 2002). Thus, when provided with adequate
habitat and protection from overhunting, the Plains Zebra
is recognised as a relatively resilient species.
Net effect Unknown
Data quality Suspected
Rationale The overall population size and geographic range of Plains Zebra has been increased significantly through the
practices of wildlife ranching. However, the effects of genetic manipulation, sexually-skewed hunting pressure and the
spread of diseases threaten to impact on overall populations.
Management
recommendation
Research into the effects of management practices on Plains Zebra kept on private properties is required.
Table 3. Possible net effects of wildlife ranching on the Plains Zebra (Equus quagga) and subsequent management
recommendations
Rank Threat description Evidence in the
scientific literature Data quality Scale of
study Current trend
1 2.1.3 Agro-industry Farming: loss of habitat from
agricultural expansions. Current stresses 1.2
Ecosystem Degradation, 1.3 Indirect Ecosystem
Effects and 2.3.5 Inbreeding: habitat degradation
and increased vulnerability to drought through
fragmentation of habitat; inbreeding in small
fenced reserves.
Walker et al. 1987
Bowland et al. 2001
Georgiadis et al. 2003
Gandiwa et al. 2016
Empirical
Empirical
Indirect
Empirical
Local
Regional
Local
Local
Increasing due to
climate change and
ongoing agricultural
expansion.
2 2.3.3 Agro-industry Grazing, Ranching or
Farming: loss of habitat from ranching
expansions. Current stresses 1.2 Ecosystem
Degradation, 1.3 Indirect Ecosystem Effects and
2.3.5 Inbreeding: habitat degradation from
overgrazing and increased vulnerability to
drought through fragmentation of habitat;
inbreeding in small fenced reserves.
Walker et al. 1987
Bowland et al. 2001
Georgiadis et al. 2003
Gandiwa et al. 2016
Empirical
Empirical
Indirect
Empirical
Local
Regional
Local
Local
Increasing due to
climate change and
ongoing agricultural
expansion.
3 1.1 Housing & Urban Areas: loss of habitat from
human settlement expansion. Current stresses
1.3 Indirect Ecosystem Effects, 2.1 Species
Mortality and 2.3.5 Inbreeding: increased
vulnerability to drought through fragmentation of
habitat and increased levels of poaching;
inbreeding in small fenced reserves.
Walker et al. 1987
Bowland et al. 2001
Georgiadis et al. 2003
Nel 2015
Gandiwa et al. 2016
Empirical
Empirical
Indirect
Empirical
Empirical
Local
Regional
Local
Local
Local
Increasing due to
climate change and
continuing expansion
of human
settlements.
4 2.3.2 Small-holder Grazing, Ranching or Farming:
loss of habitat from small-holder ranching.
Current stresses 1.2 Ecosystem Degradation, 1.3
Indirect Ecosystem Effects, 2.3.1 Hybridisation
and 2.3.5 Inbreeding: habitat degradation from
overgrazing and increased vulnerability to
drought through fragmentation of habitat;
inbreeding in small fenced reserves;
hybridisation and decreased genetic diversity in
small subpopulations (< 100 zebra) on wildlife
ranches.
Walker et al. 1987
Bowland et al. 2001
Georgiadis et al. 2003
Gandiwa et al. 2016
Empirical
Empirical
Indirect
Empirical
Local
Regional
Local
Local
Increasing due to
climate change and
expansion of wildlife
ranches.
5 8.1.1 Invasive Non-Native/Alien Species/
Diseases: spread of pathogens and parasites
through translocations into new areas and/or
mixing subspecies.
- Anecdotal - Possibly increasing
through expansion of
wildlife ranching and
translocations.
Table 4. Threats to the Plains Zebra (Equus quagga) ranked in order of severity with corresponding evidence (based on IUCN
threat categories, with regional context)
Equus quagga | 6 The Red List of Mammals of South Africa, Lesotho and Swaziland
protected areas in order to sustain genetic diversity and
population viability. On the other hand, an important
consideration is that the expansion of the species’ range
through translocation could introduce new pathogens and
parasites to populations without co-evolved resistance to
them (Hack et al. 2002). Plains Zebra occur across a
range of habitats, and it is probable that they have distinct
adaptations to local conditions (Hack et al. 2002).
Expanding its range through translocation from
ecologically diverse regions may result in a decline in
these particular adaptations. Consequently, the spatial
scale, ecological conditions, genetics and resistance to
diseases need to be considered before translocations
occur. Similarly, while wildlife ranching is considered
largely beneficial for Plains Zebra conservation in South
Africa, and has effectively increased the extent and
abundance of the species, some concern exists over the
genetic implications of selective breeding for particular
colour morphs and outbreeding depression caused by
mixing subspecies. Further research is required to
quantify this potential threat.
Current habitat trend: Stable. Savannah habitat is not
threatened within the assessment region (Driver et al.
2012). Habitat available to Plains Zebras has increased
through an increase in transfrontier conservation areas,
privately owned reserves, and the expansion of wildlife
ranching. However, there is some loss in habitat quality
through agriculture and livestock farming. Grassland
biomes form key habitat for Zebra, and is one of South
Africa’s most threatened ecosystems with only ~2.4%
being formally conserved (Carbutt & Martindale 2014).
Conservation
Within the assessment region, Plains Zebra occur in
numerous well-managed protected areas, where KNP and
its surrounding reserves protect the core free-roaming
population. The majority of the reserves surrounding KNP
are not fenced and allow for unrestricted movement of
Plains Zebra and consequently, enhanced genetic
diversity. KNP also forms part of the Greater Limpopo
Transfrontier Park which allows movement over a much
larger scale. The Greater Mapungubwe TFCA also allows
unrestricted Plains Zebra migration between Botswana,
South Africa and Zimbabwe. Continued expansion of
transfrontier conservation areas will enable this species to
adapt to changing climates and environments and thus
enhance the resilience of the population (for example,
Bartlam-Brooks et al. 2011).
While no specific interventions are required at present,
Hack et al. (2002) recommended the following
conservation strategies for Plains Zebra that are relevant
for the assessment region:
1. Improve coverage and frequency of monitoring.
Annual or biennial sampling of population abundance,
location and conservation status is required in order
to detect problems in their primary stages and
respond effectively. This is particularly relevant for
private protected areas and wildlife ranches to gather
baseline information needed to develop a Biodiversity
Management Plan and translocation policy.
2. Improve risk assessment. A detailed and local
investigation into site- and subpopulation-specific
hazards, particularly with regards to mitigating
potential inbreeding, hybridisation and disease
transmission on small fenced properties, is necessary.
3. Quantify and manage genetic diversity, both globally
and locally. Management plans may need to be
specifically reviewed and implemented for genetically
distinct populations.
4. Investigate the economics of alternative utilisation
strategies. Quantifying and comparing the economic
uses of this species may guide future global
management strategies, and incentivise landowners
to further reintroduce subpopulations within the
natural range of the species.
Recommendations for land managers and
practitioners:
Improve management of small subpopulations to
prevent inbreeding and a loss of genetic diversity.
This can be achieved through the development of a
Biodiversity Management Plan and associated
translocation policy. For the latter, a harem should
be translocated every 5 years for a subpopulation
size of c. 9 individuals, while for a population size of
c.110 individuals, translocations should take place
every 15 years if heterozygosity is to be maintained
at more than 90% within each subpopulation over
100 years (Bowland et al. 2001).
Remove fences to create conservancies, which allow
for an increase in free-roaming subpopulations.
Rank Intervention description
Evidence in
the scientific
literature
Data
quality
Scale of
evidence
Demonstrated
impact
Current
conservation
projects
1 1.1 Site/Area Protection: expand transfrontier
areas to create corridors for migration and
adaptation.
Bartlam-Brooks
et al. 2011
Empirical Regional Migration route
restored after
creation of
corridor.
Peace Parks
Foundation
2 3.3.1 Species Reintroduction: translocations of
harems in small intensively managed
populations to increase genetic diversity,
possibly through metapopulation strategy.
- Anecdotal - - -
3 4.3 Awareness & Communications: inform game
farmers and the hunting industry on dangers of
selective breeding and inbreeding.
- Anecdotal - - -
Table 5. Conservation interventions for the Plains Zebra (Equus quagga) ranked in order of effectiveness with corresponding
evidence (based on IUCN action categories, with regional context)
The Red List of Mammals of South Africa, Lesotho and Swaziland Equus quagga | 7
Maintain accurate abundance, distribution and
population structure records.
The management of artificial waterholes plays a vital
role in the population dynamics of this species. The
spatial distribution of active waterpoints should vary
temporally. This will create variation in habitat use
and movements across the landscape, thereby
sustaining landscape heterogeneity.
Research priorities:
Investigating the effects of wildlife ranching,
including factors associated with sexually-skewed
hunting pressure and the corresponding influence
on female reproduction success. It is suggested that
high turnover rates of dominant, harem males can
negatively influence female body condition and
hormone fluctuations (Berger 1983). Research is
necessary to determine whether this is the case for
Plains Zebras.
Studies into the factors regulating natural population
fluctuations of Plains Zebra would aid in predicting
their responses to human-induced changes to their
habitats.
Examining the spread of pathogens and parasites
via harem translocations into new areas.
Encouraged citizen actions:
Citizens can refuse to hunt artificial colour morphs
and hybrids.
Landowners could remove fences and create
conservancies and corridors for this species,
encouraging natural migrations and movements.
Subpopulations outside of the species’ natural
distribution (for example, Western Cape Province)
could be removed.
Private landowners, provincial and national reserves
can contribute data on subpopulation numbers and
structures to virtual museum platforms (for example,
iSpot and MammalMAP) to ensure accurate
assessments of species through the Red Listing
process.
References
Arsenault R, Owen-Smith N. 2008. Resource partitioning by grass
height among grazing ungulates does not follow body size
relation. Oikos 117:17111717.
Bartlam-Brooks HLA, Bonyongo MC, Harris S. 2011. Will
reconnecting ecosystems allow long-distance mammal
migrations to resume? A case study of a zebra Equus burchelli
migration in Botswana. Oryx 45:210216.
Bennett DK. 1980. Stripes do not a zebra make, Part I: A cladistic
analysis of Equus. Systematic Biology 29:272287.
Berger J. 1983. Induced abortion and social factors in wild
horses. Nature 303:5961.
Bissett C, Ferreira S, Bezuidenhout H, Smit I, Daemane E,
Mokoena V, Sikhosana T. 2016. Golden Gate Highlands National
Park herbivore off-take recommendations 2016: An integrated
approach combining local knowledge with data derived from
animal census, herbivore models, vegetation field monitoring and
satellite imagery. Internal Report 09 / 2016, Scientific Services,
South African National Parks, South Africa.
Bowland AE, Bishop KS, Taylor PJ, Lamb J, van der Bank FH,
van Wyk E, York D. 2001. Estimation and management of genetic
diversity in small populations of plains zebra (Equus quagga) in
KwaZulu-Natal, South Africa. Biochemical Systematics and
Ecology 29:563583.
Carbutt C, Martindale G. 2014. Temperate indigenous grassland
gains in South Africa: Lessons being learned in a developing
country. PARKS 20:101121.
Churcher C, Richardson M. 1978. Equidae. Pages 379422 in
Maglio V, Cooke H, editors. Evolution of African Mammals.
Harvard University Press, Cambridge, USA.
Di Minin E, Leader-Williams N, Bradshaw CJ. 2016. Banning
trophy hunting will exacerbate biodiversity loss. Trends in
Ecology & Evolution 31:99102.
Doku Y, Bekele A, Balakrishnan M. 2007. Population status of
plains zebra (Equus quagga) in Nechisar plains, Nechisar
National Park, Ethiopia. Tropical Ecology 48:7986.
Driver A, Sink KJ, Nel JN, Holness S, Van Niekerk L, Daniels F,
Jonas Z, Majiedt PA, Harris L, Maze K. 2012. National Biodiversity
Assessment 2011: An Assessment of South Africa’s Biodiversity
and Ecosystems. Synthesis Report. South African National
Biodiversity Institute and Department of Environmental Affairs,
Pretoria, South Africa.
du Plessis SF. 1969. The past and present distribution of the
Perissodactyla and Artiodactyla in southern Africa. M.Sc. Thesis.
University of Pretoria, Pretoria, South Africa.
Duncan P. 1992. Zebras, Asses and Horses: an Action Plan for
the Conservation of Wild Equids. IUCN, Gland, Switzerland &
Cambridge, UK.
East R. 1997. Current status of Burchell’s Zebra in Africa, with
additional information on Grevy’s Zebra and Cape Mountain
Zebra. IUCN SSC Equid Specialist Group.
Estes R. 1991. The Behaviour Guide to African Mammals.
University of California Press, Berkeley, USA.
Ferreira S, Gaylard A, Greaver C, Hayes J, Cowell C, Ellis G.
2013. Animal abundances in Parks 2012/2013. Scientific Services,
SANParks, Skukuza, South Africa.
Gandiwa E, Heitkönig IM, Eilers PH, Prins HH. 2016. Rainfall
variability and its impact on large mammal populations in a
complex of semi-arid African savanna protected areas. Tropical
Ecology 57:163180.
Georgiadis N, Hack M, Turpin K. 2003. The influence of rainfall on
zebra population dynamics: implications for management.
Journal of Applied Ecology 40:125136.
Grange S, Barnier F, Duncan P, Gaillard J-M, Valeix M, Ncube H,
Périquet S, Fritz H. 2015. Demography of plains zebras (Equus
quagga) under heavy predation. Population Ecology 57:201214.
Grange S, Duncan P, Gaillard J-M, Sinclair AR, Gogan PJ, Packer
C, Hofer H, East M. 2004. What limits the Serengeti zebra
population? Oecologia 140:523532.
Groves CP, Bell CH. 2004. New investigations on the taxonomy of
the zebras genus Equus, subgenus Hippotigris. Mammalian
Biology 69:182196.
Data sources Field study (unpublished)
Data quality (max) Estimated
Data quality (min) Estimated
Uncertainty resolution Best estimate
Risk tolerance Evidentiary
Table 6. Information and interpretation qualifiers for the
Plains Zebra (Equus quagga) assessment
Data Sources and Quality
Equus quagga | 8 The Red List of Mammals of South Africa, Lesotho and Swaziland
Hack AM, East R, Rubenstein DI. 2002. Status and Action Plan for
the Plains Zebra. Pages 4360 in Moehlman PD, editor. Equids:
Zebras, Asses and Horses. Status Survey and Conservation
Action Plan. IUCN, Gland, Switzerland.
Hayward MW. 2009. Bushmeat hunting in Dwesa and Cwebe
Nature Reserves, Eastern Cape, South Africa. South African
Journal of Wildlife Research 39:7084.
Hofer H, Campbell KLI, East ML, Huish SA. 1996. The impact of
game meat hunting on target and non-target species in the
Serengeti. Pages 117146 in Taylor VJ, Dunstone N, editors. The
Exploitation of Mammal Populations. Chapman and Hall, London,
UK.
Joos-Vandewalle ME. 2000. Movement of migratory zebra in
northern Botswana. Ph.D. Thesis. University of the Witwatersrand,
Johannesburg, South Africa.
Klingel H. 1969. The social organisation and population ecology
of the plains zebra (Equus quagga). Zoologica Africana 4:249
263.
Lindsey PA et al. 2013. The bushmeat trade in African savannas:
Impacts, drivers, and possible solutions. Biological Conservation
160:8096.
Lorenzen ED, Arctander P, Siegismund HR. 2008. High variation
and very low differentiation in wide ranging plains zebra (Equus
quagga): insights from mtDNA and microsatellites. Molecular
ecology 17:28122824.
Naidoo R, Chase MJ, Beytell P, du Preez P, Landen K, Stuart-Hill
G, Taylor R. 2016. A newly discovered wildlife migration in
Namibia and Botswana is the longest in Africa. Oryx 50:138146.
Nel P. 2015. Population estimates for large herbivores and
predators in protected areas in the North West Parks Board
November 2015. North West Parks Board. Mahikeng, South
Africa.
Pacifici M, Santini L, Di Marco M, Baisero D, Francucci L, Marasini
GG, Visconti P, Rondinini C. 2013. Generation length for
mammals. Nature Conservation 5:8994.
Power RJ. 2014. The Distribution and Status of Mammals in the
North West Province. Department of Economic Development,
Environment, Conservation & Tourism, North West Provincial
Government, Mahikeng, South Africa.
Rautenbach IL. 1982. Mammals of the Transvaal. No. 1, Ecoplan
Monograph. Pretoria, South Africa.
Assessors and Reviewers
Keenan Stears1, Adrian Shrader1, Guy Castley2
1University of KwaZulu-Natal, 2Griffith University
Contributors
Claire Relton1, Matthew F. Child1, Patricia Moehlman2,†,
Jeanetta Selier3, Sarah King2
1Endangered Wildlife Trust, 2EcoHealth Alliance, 3South African
National Biodiversity Institute
IUCN SSC Equid Specialist Group
Details of the methods used to make this assessment can
be found in Mammal Red List 2016: Introduction and
Methodology.
Regassa R, Yirga S. 2013. Distribution, abundance and
population status of Burchells zebra (Equus quagga) in Yabello
Wildlife Sanctuary, Southern Ethiopia. Journal of Ecology and the
Natural Environment 5:4049.
Schulz E, Kaiser TM. 2013. Historical distribution, habitat
requirements and feeding ecology of the genus Equus
(Perissodactyla). Mammal Review 43:111123.
Skinner JD, Chimimba CT. 2005. The Mammals of the Southern
African Subregion. Third edition. Cambridge University Press,
Cambridge, UK.
Smuts GL. 1976. Population characteristics of Burchell’s zebra
(Equus burchelli antiquorum, H. Smith, 1841) in the Kruger
National Park. South African Journal of Wildlife Research 6:99
112.
Stuart CT, Stuart MD. 2007. Field Guide to Mammals of Southern
Africa. Random House Struik (Pty) Ltd, Cape Town, South Africa.
Walker BH, Emslie RH, Owen-Smith RN, Scholes RJ. 1987. To
cull or not to cull: lessons from a southern African drought.
Journal of Applied Ecology 24:381401.
ResearchGate has not been able to resolve any citations for this publication.
Article
Full-text available
International pressure to ban trophy hunting is increasing. However, we argue that trophy hunting can be an important conservation tool, provided it can be done in controlled manner to benefit biodiversity conservation and local people. Where political, and governance structures are adequate, trophy hunting can help address the ongoing loss of species.
Article
Full-text available
. The population was female biased. The sex ratio of adult male to adult female was 1.0:1.27 and adult to young ratio was 2.9:0.6. Age composition of Burchell’s zebra comprised 78.2% adult, 13.0% subadult, 5.3% juvenile and 3.5% foal. Group size changed seasonally and the mean group size was 12.5. The average herd sizes of one male harem and bachelor stallion herds were 7.8 and 5.3, respectively. Distribution and vegetation utilization of the animal showed a marked preference for open grassland habitat. However, there was a seasonal change in the preference of habitat. Increase in human and livestock population was observed in the study area. Overgrazing by cattle and encroachment are the primary factors that affect the population status of Burchell’s zebra by reducing the grass quality in the Sanctuary. The study recommends an effective and realistic management policy to control illegal human settlement and encroachment in the sanctuary.
Article
Full-text available
Migrations of most animal taxa are declining as a result of anthropogenic pressures and land-use transformation. Here, we document and characterize a previously unknown multi-country migration of Burchell's zebra Equus quagga that is the longest of all recorded large mammal migrations in Africa. Our data from eight adult female zebras collared on the border of Namibia and Botswana show that in December 2012 all individuals crossed the Chobe River and moved due south to Nxai Pan National Park in Botswana, where they spent a mean duration of 10 weeks before returning, less directly, to their dry season floodplain habitat. The same southward movements were also observed in December 2013. Nxai Pan appeared to have similar environmental conditions to several possible alternative wet season destinations that were closer to the dry season habitat on the Chobe River, and water availability, but not habitat or vegetation biomass, was associated with higher-use areas along the migratory pathway. These results suggest a genetic and/or cultural basis for the choice of migration destination, rather than an environmental one. Regardless of the cause, the round-trip, straight-line migration distance of 500 km is greater than that covered by wildebeest Connochaetes taurinus during their well-known seasonal journey in the Serengeti ecosystem. It merits conservation attention, given the decline of large-scale ecological processes such as animal migrations.
Article
Full-text available
In natural ecosystems, ungulate densities show strong temporal variations. The ecological processes driving these fluctuations are complex: food limitation and predation are both important and can interact. Survival rates are central to this debate, but data are sparse for tropical ecosystems. Here, we estimate age- and sex-specific survival rates for plains zebra in Hwange National Park, a nutrient-poor savanna with a high predator–prey ratio. We estimated survival from a detailed Capture-Mark- Recapture (CMR) monitoring based on 248 individual life histories, for the first time in an African grazer. We controlled for variations in detection probabilities among adult females, which resulted from their social structure. As expected, annual survival was low during the first year (0.441); increased in yearlings (0.560) and peaked at 0.795 and 0.847 in adult males and females respectively. The survival of adult females was lower during the dry season, which probably resulted from higher predation due to predictable movements of zebras to waterholes. Survival at all ages was low compared to ungulates without predators. The demographic model we constructed showed a declining trend (k = 0.94), which was consistent with the data from road counts (^k = 0.92). Life Table Response Experiment (LTRE) analyses using the Serengeti and Kruger populations as references showed that the main cause of this declining trend in the Hwange population was low survival in yearling and adult females; low foal survival also contributed. In this ecosystem, predation is likely to be the main ecological process causing low survival, and therefore a decline in the zebra population.
Article
Full-text available
The fragile state of temperate indigenous grasslands globally has galvanised action in the form of the Temperate Grasslands Conservation Initiative of the International Union for Conservation of Nature’s World Commission on Protected Areas. However, despite this initiative raising the profile of temperate grassland conservation on the global conservation agenda, one still requires country-based interventions at the hands of local conservation authorities, in collaboration with non-governmental organisations (NGOs), to improve protection levels on the ground. To this end we report on progress made with temperate indigenous grassland conservation in South Africa since 2006, a landmark heralding the birth of biodiversity stewardship in our temperate grassland biome. Since then an additional 124,983 ha of temperate grassland have been brought under formal protection with a further 96,641 ha in the declaration process, most of which should be secured by the end of 2014. We also discuss the driving forces underpinning these gains - namely the National Protected Area Expansion Strategy, the Grasslands Programme of the South African National Biodiversity Institute, provincial biodiversity stewardship units and funding channelled through the Critical Ecosystems Partnership Fund into civil society to augment the state’s contribution. Given the clear benefits derived from each intervention, we encourage other relevant countries with temperate indigenous grasslands to develop similar structures in the quest to safeguard representative, viable samples of one of the world’s great terrestrial biomes.
Article
Full-text available
We investigated the rainfall patterns and associated fluctuations of wild large herbivore species in the Great Limpopo Transfrontier Conservation Area (GLTFCA), southern Africa. The study objectives were to: (i) establish the synchrony in rainfall and drought occurrence patterns in Gonarezhou National Park, Zimbabwe, and four adjacent areas, and (ii) determine the responses of different large herbivore species’ populations to droughts. We used annual rainfall data collected from the five sites within the GLTFCA and large herbivore population data collected from multispecies aerial surveys in Gonarezhou and Kruger National Park, South Africa. Our results showed that between 1970 and 2009, Gonarezhou recorded three wet years (1977, 1978 and 2000) and six drought years (1973, 1983, 1989, 1992, 1994 and 2005). However, there were some variations in the drought occurrences between Gonarezhou and the four adjacent areas indicating a weak synchrony in rainfall patterns. Furthermore, seven large herbivore species showed dips in their populations associated with the 1992 severe drought, with most of the species’ populations recovering thereafter. Our study suggests that rainfall does have a strong influence on large herbivore population dynamics especially in really dry years in African savanna ecosystems. Our findings underscore the need for further detailed studies on bottom-up processes influencing large herbivore population trends in savanna ecosystems with high rainfall variability.
Article
Species and subspecies of zebras are examined using traditional (pelage and craniometric) taxonomic methods, including multivariate analysis. Mountain Zebras are split into two species, Equus zebra and Equus hartmannae, because they differ absolutely (100%); Burchell's or plains zebras, however, are placed in Equus quagga because there are no absolute differences. The subspecies of Equus quagga are revised; six subspecies are recognisable. E. zebra, E. hartmannae and E. grevyi are monotypic.
Article
(1) The ecological consequences of a severe drought on four wildlife conservation areas in southern Africa (Tuli, Klaserie, Kruger, Umfolozi) are compared. (2) Herbivore mortality occurred during the second year of the drought, when failure of the spring rains coincided with depleted food reserves. It was severe in Klaserie, with up to 90% mortality. In Tuli, mortality was high but restricted mostly to wildebeest and zebra; impala (a mixed grazer-browser) and browsers were little affected. Mortality was up to 35% in Kruger; it included browsers, but wildebeest and zebra were little affected. Mortality was low in Umfolozi, even in an unculled block, although pre-emptive culling exceeded the average mortalities at Kruger. Ungulate populations recovered to predrought levels within 2 years in Kruger but, apart from impala, will take longer in the more severely affected areas. (3) Mortality of grass tufts was extensive in Klaserie, Umfolozi and Tuli, and was limited to areas near water in Kruger. Initial recovery was rapid in three areas after one season of above-average rainfall, but was retarded in Tuli, and later in Klaserie because of continuing drought and (possibly) continued impala grazing. Little tree mortality was recorded. (4) Ungulate mortality was high in the most arid areas (Tuli and Klaserie) where grass mortality was particularly high and, most importantly, where extensive surface water permitted a high pre-drought biomass and even distribution of ungulates. (5) It is concluded that culling is ecologically unnecessary where sufficient spatial heterogeneity exists to provide reserve forage. Some drought-related mortality is natural and probably beneficial to both animal and plant populations. Wild ungulate populations should be allowed to fluctuate within limits set by management objectives and culling is likely to be necessary only where the provision of water points has eliminated reserve forage areas.
Chapter
In the Serengeti National Park (SNP), illegal game meat hunting is largely carried out using snares in the south-western, western and north-western areas. Game meat hunting provides cash income and protein to communities outside the SNP. The economic benefits of game meat hunting have drawn people to villages close to the park boundary, causing a rise in human population density well above the regional average. Game meat hunting has already drastically reduced populations of Cape buffalo and must in the long term be considered unsustainable for a number of other herbivore species. In this chapter an estimate of the current wildlife offtake from the National Park is made and the impact of unselective hunting methods on carnivore species, the most common non-target species, is considered. The analysis demonstrates that game meat hunting poses a threat to both target and non-target species of the Serengeti wildlife community. Optimality models, commonly used in behavioural ecology and economics, are introduced to assess a hunter’s profit in relation to hunting effort (costs) and to ask whether unchecked illegal hunting is likely to be sustainable in the long term. A review of studies on African systems demonstrates that whenever costs are reduced, the impact on wildlife due to illegal hunting is dramatically increased and reaches unsustainable levels. Proposals to limit wildlife offtake to sustainable levels, including limited legalization of game meat hunting in areas adjacent to SNP and the development of alternative sources of income and protein for local communities, are considered. The evaluation of these proposals suggests that the situation in the Serengeti does not meet the pre-conditions and assumptions of programmes developed elsewhere for maximizing economic returns from wildlife utilization as an incentive to preserve wildlife; hence such programmes are unlikely to be successful here. This is because the Serengeti is a wildlife system dominated by migratory herbivores, exacerbating the problem of assigning unambiguous ownership of wildlife outside the protected area to a given local community — a pre-condition for any successful privatization or commercialization scheme. Also, if future community conservation services are focused only on those communities that currently benefit most from illegal exploitation, i.e. communities adjacent to the protected areas, then such programmes are likely to reinforce a vicious cycle. They are likely to attract more people to villages close to the protected area and ultimately put greater demands on the proted areas, just as currently people are attached to these villages because of enhanced oppurtunities for illegal hunting. The analysis suggests that in ecosystems dominated by migration herbivores and wih low levels of law enforcement a large investment is required in both law enforcement and rural development of local communities, that the success of the latter may be linked to investment in the former, and that without both of these the long-term conservation of Serengeti wildlife populations is unlikely to be ensured.