Original Research Article
Conservation implications of wildlife translocations; The
state's ability to act as conservation units for wildebeest
populations in South Africa
, Brian K. Reilly
Conservation Beyond Borders, Director's Desk, 3033 Excelsior Blvd, #575, 55416, Minnesota, USA
Department of Nature Conservation, Faculty of Science, Tshwane University of Technology, Staatsartillery Road, Pretoria, 0001, South
Received 30 June 2017
Received in revised form 18 August 2017
Ecological Sustainable Networks (ESN)
Wildlife translocations have historically assisted in re-establishing species in areas of
extinction and are currently employed in over 50 countries. Ironically, they may also be
responsible for the extinction of pure genetic lineages via hybridization, thereby negatively
impacting endangered, indigenous, and rare species. Due to recent evolutionary diver-
gence, black wildebeest (Connochaetes gnou) and blue wildebeest (Connochaetes taurinus)
can mate and produce fertile offspring when sympatric. A total of 6929 translocated black
and blue wildebeest from 273 private ranches and 3 provincial protected areas protected
(PPAs) were documented over 5 years, across 5 South African provinces. We analyzed
dispersal patterns and wildlife ranching economics to identify conservation implications
and to infer if translocations are likely to persist in their current form. Findings indicate (1)
58.45% of sampled private ranches manage for both wildebeest populations, (2) blue
wildebeest males are primarily translocated, (3) wildebeest are introduced across pro-
vincial lines, (4) wildebeest are introduced to within and amongst the private and com-
mercial industry from multiple sources, and (5) wildebeest revenue accounted for 20.8% of
revenue generated from all wildlife translocations. Unwanted conservation implications
concern ecological integrity, genetic swamping, and regulatory efﬁciency. We caution
against risks posed by the game industry upon the PPA's ability to function as nature
conservation units and act as stocking sources and the plausibility that black wildebeest
populations incorporate varying degrees of introgressive hybrids. Moreover, wildebeest
account for 1/5 of revenue generated from all game translocations. This is indicative of its
likelihood to persist in their current form, thereby inducing hybridization and facilitating
outbreeding depression. We caution that concerns are likely to worsen if no intervention is
taken. Lastly, we coin the concept of Ecological Sustainable Network (ESN); we designed a
framework for standardizing procedures to advance effective wildlife translocation prac-
©2017 The Authors. Published by Elsevier B.V. This is an open access article under the CC
BY license (http://creativecommons.org/licenses/by/4.0/).
E-mail addresses: Nicole@Conservationbeyondborders.org (N. Benjamin-Fink), ReillyBK@tut.ac.za (B.K. Reilly).
Contents lists available at ScienceDirect
Global Ecology and Conservation
journal homepage: http://www.elsevier.com/locate/gecco
2351-9894/©2017 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.
Global Ecology and Conservation 12 (2017) 46e58
1.1. The extent of the wildlife ranching industry within South Africa
South Africa encompasses more than 11,600 private wildlife ranches, spreading over 21 million hectares (Dry, 2013; Van
Hoven, 2015). An additional 15,000 ranches manage both domestic livestock and wildlife (Patterson and Khosa, 2005 as cited
in Cousins et al., 2008). During 1991e2000, South Africa experienced an annual increase of 5.6% in land used for wildlife
ranching (Cloete et al., 2015). Currently, 16.8% of private ranches and 6.1% of provincial protected areas (PPAs) utilize the
privatization of wildlife as their primary source of revenue; in fact, wildlife managed on private ranches is almost threefold of
that managed on the provincial land (Bothma, 2002; Cousins et al., 2010).
Wildlife ranching has the higher rate of return per hectare than any agricultural based market (Slabbert, 2013). With an
annual return on investment upwards of 80%, wildlife enterprises generate 4.7 billion Rand/year (Oliver, 2015). They are
subdivided into the following, often overlapping, market segments: hunting (3.1 billion Rand/year), trophy hunting (510
million Rand/year), game translocation (750 million Rand/year), live auctions (1 billion Rand/year), game meat production
(42 million Rand/year), and taxidermy (200 million Rand/year) (Bothma et al., 2009; Du Toit and Van Schalkwyk, 2011;
Grobler et al., 2011).
1.2. Wildlife ranching entails the management of enclosed populations and varying objectives for translocations
South African wildlife ranching practices require fences along the perimeter of all game ranches. As such, populations are
closed and ﬁnite and natural processes do not take place (e.g., dispersal, emigration, and colonization dynamics) (Cousins
et al., 2008). Consequently, ranch owners are faced with the need to intensively manage populations on their land while
weighting economic proﬁtability with genetic concerns stemming from small and closed populations (e.g., inbreeding,
outbreeding depression, and bottlenecks) (Lehmann et al., 2008).
The commercial nature of the wildlife ranching industry has resulted in wildlife translocation practices taking place in over
50 countries worldwide; United States of America and South Africa have the highest utilization (Spear and Chown, 2009a).
Various market sectors within its industry (see above) indirectly act as drivers for translocation efforts; primary examples
include the need to stock animals for hunting (which is employed in over 23 countries and across 1,394,000 km
and national land in sub-Saharan Africa) and ecotourism (driven by wildlife viewing) (Bothma, 2002; Child, 2012; Fischer
et al., 2015, Lewis and Alper, 1997; Lindsey et al., 2007; Loveridge et al., 2007). IUCN (1988) guidelines deﬁne translocation
as the “mediated movement of living organisms from any source (privately managed or wild), with release in another”. This
oversimpliﬁed deﬁnition enables local decision makers to determine if a translocation effort is conservation oriented. The
conservation-based goal of translocating individuals into an existing population is primarily to increase genetic diversity via
outbreeding, whereby genetic diversity is increased through the mating of an unrelated individual and a breeding population
(Balding, 2007). However, such management decisions are complex and there is a need to mitigate the risk of unwanted
consequences (e.g., outbreeding depression). Conservation-based translocations are conducted with the objectives to
counter: genetic bottlenecks, local extinction events, and/or inbreeding processes by re-establishing, recolonizing, replacing,
restoring, relocating, and reinforcing the population, in addition to providing biological control (Rhymer and Simberloff,
Non-conservation related objectives include a substitute for culling, recreation, biological control, aesthetics, religion,
wildlife rehabilitation, color variance, and animal rights activism (Pasquini et al., 2010, Seddon et al., 2012). There is a lack of a
much-needed standardization in the wildlife translocation practice throughout South Africa (Grobler et al., 2011).
1.3. State's role as conservation units
Regardless of whether or not motivations are conservation-based, wildlife translocations have the capacity to shape
ecosystem dynamics. Historically, translocation practices served as an effective conservation tool, bringing various species
back from the brink of extinction through the reintroduction of animals (Hayward et al., 2007). We coin the deﬁnition of the
concept of national parks (i.e., a state) acting as conservation units and deﬁne this as the state's ability to act as a source for
genetically pure individuals for the purpose of serving as population founders on governmental and/or private land. We
emphasize the underlying principle of wildlife management plans concerning the reintroduction of locally extinct pop-
ulations; it is critical that founders are pure breed (i.e., genetic status) in order to reestablish taxonomy-pure populations. We
caution that the risk of translating genetically admixture individuals with the intention to reestablish closed populations is
the facilitation of hybridization, divergence and speciation. Primary examples of reintroductions of locally extinct species in
the South African context include the southern white rhinoceros (Ceratotherium simum), Cape mountain zebra (Equus zebra),
bontebok (Damaliscus pygargus phillipsi), and the black wildebeest (Connochaetes gnou)(Fabricious et al., 1988; Robinson
et al., 1991; Flack, 2003 Hamman et al., 2003).
Ironically, the genetic integrity of the later three is currently jeopardized by the same management practice that originally
enabled them to persist and survive bottlenecks, wildlife translocation. In fact, upwards of 60% of Blesbok (Damaliscus
pygargus phillipsi) are varying degrees of bontebok X blesbuck hybrids (Van Wyk et al., 2013, 2017).
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e58 47
1.4. Introgressive hybridization
Introgressive hybridization refers to the process during which hybridization and backcrossing result in alleles at one locus
from one species introgressing into the gene pool of other loci from another species (http://jhered.oxfordjournals.org/
content/105/S1/795.full,Anderson, 1949). This may occur as a result of artiﬁcial sympatry between two recently diverged
species, which have not had the evolutionary time needed to developed separate reproductive isolation mechanisms (Arnold,
1992). Introgression followed by inbreeding, whereby individuals from a closed population mate and produce offspring, may
lead to genetic admixture (resulting in a new genetic lineage) (Balding, 2007).
Unintentionally overlapping previously ecological allopatric species may facilitate the following deleterious results: (1)
pressure on the local carrying capacity, (2) shifts in ecosystem dynamics, (3) alteration of local vegetation, or (4) ill desired
genetic consequences and therefore decreased ﬁtness (e.g., hybridization, drift, homogenization, admixture, and outbreeding
depression) (Brink, 1993; Coates and Downs, 20 05; Harrison, 1993; Leimu and Fischer, 2010; Watson, 20 06). In fact, many of
these processes are characteristic of hybrid zones; regions where recently diverged lineages are artiﬁcially overlapped, cross-
fertilize and/or backcross occurs, and selection favors hybrid allelic diversity, thereby promoting hybrid vigour (Allendorf
et al., 2001; Arnold and Martin, 2010; Batron and Hewitt, 1985). Such hybrid zones are initiated by species artiﬁcial over-
lap which may result from translocations. This, in turn, may create artiﬁcial bottlenecks in ﬁnite populations, thereby pro-
moting and sustaining hybridization (Arnold, 1992). For these reasons, translocation to closed and ﬁnite populations (typical
of South Africa ranches due fencing requirements) may facilitate the creation of hybrid zones and lead to the extinction of
pure genetic lineages, such as that of the endemic black wildebeest.
1.5. Wildebeest core and extralimital habitat, social structure, and hybridization
The endemic keystone species black wildebeest serves as an example of both the positive and negative conservation
implications embedded in the wildlife management practices of translocation. Its historical distribution included South Africa,
Lesotho and Swaziland (Brink,1993). Hunted to almost extinction in the ﬁrst 2 countries, extensive translocation efforts played
signiﬁcant roles in countering 2 bottleneck effects (Brink, 2005; Corbet and Robinson, 1991). After surviving 2 bottlenecks, the
South African department of environmental affairs formally listed the black wildebeest as a Threatened Or Protected Species
(TOPS). The wildebeests' core habitat was the boundary between the Karoo and the Highveld regions (Brink, 1993; Skinner and
Chimimba, 2005). However, its current extralimital range (i.e., distribution beyond their core habitat) includes the Highveld,
Bushveld, Kalahari, Karoo, and Fynbos ecoregions (Spear and Chown, 2009b). Moreover, its range did not include Namibia
prior to 1972, and yet in 1992, their numbers reached 7177 in Namibia (Barnes and Jager, 1996). The same survey documented
326 blue wildebeest (Connochaetes taurinus) in 1972 and 4935 in 1992 in the same region. Such a dramatic increase over 20
years implies that intensive translocation practices have moved the black wildebeest into an extralimital habitat, overlapping
with the blue wildebeest in their natural habitat (Ackermann et al., 2010; Skinner and Chimimba, 2005). Currently, blue
wildebeest's and black wildebeest's distribution overlaps throughout South Africa and Namibia.
Wildebeest species are social. They interact through three primary social structures: female herd (adult and sub-adult
females, calves and yearlings), bachelor herds (i.e., sub-adults and yearling males), and territorial or solitary bulls (Von
Ricter, 1972; as cited in Helm, 2007). In natural occurrences (i.e., when sympatry is not anthropogenic-driven and
enforced), wildebeest species occupy different ecological niches. The historical distribution of blue wildebeest is in woodland
areas and grasslands, while black wildebeest exclusively inhabits grasslands (Codron and Brink, 2007; Estes, 1991).
The black wildebeest lineage diverged from a shared ancestor merely 1.8 million years ago; this short time on the
evolutionary scale did not allow for the development of reproductive isolation mechanisms that would act as a barrier to gene
ﬂow when the 2 species are artiﬁcially sympatric (Brink, 1993). Changes in biological and ecological barriers may result in an
increased risk for black wildebeest experiencing genetic introgression of regions of the blue wildebeest DNA (i.e., hybridi-
zation) (Ackermann et al., 2010; Grobler et al., 2005). Such hybridization occurs because male blue wildebeest physically
displace male black wildebeest when overlapped. Hybridization between black and blue wildebeest was ﬁrst documented on
Abe Bailey reserve in South Africa, where as a result, the entire population was culled (Grobler et al., 2005). Since then,
additional populations containing varying degrees of introgression and hybrids have been culled (e.g., Spioenkop Dan Nature
Reserve, in KwaZulu-Natal) (Ackermann et al., 2010).
Biological, genetic, ecological, and conservation concerns evolving from the wildebeest hybridization phenomenon have
been the focus of a number of prominent research papers (Ackermann et al., 2010; Brink, 1993; Corbet and Robinson, 1991;
Grobler et al., 2005, 2011; Roed et al., 2011). Alarmingly, a signiﬁcant proportion of game translocations may involve
wildebeest, and yet, there is no comprehensive database indicating the location, or even existence, of pure black wildebeest
populations or revealing the extent of extralimital translocations.
1.6. Intentional and unintentional consequences of the practice of wildebeest translocation; hybridization vs. breeding for color
There are ﬁve subspecies of blue wildebeest throughout the South African continent, the C.t.taurinus is the solely blue
wildebeest inhabiting South Africa and is artiﬁcially overlapped with the endemic black wildebeest (Fabricious et al., 1988).
An undesired by-product of such sympatric conditions is hybridization, which jeopardizes instrumental efforts to conserve
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e5848
the latter's genetic integrity (Ackermann et al., 2010). In contrast, directed wildlife breeding for color variants requires
proactive translocations that induce targeted sympatry as a management strategy towards the production of an offspring of a
certain morphological characteristic (i.e., coat color). With an annual turnover of upwards of 1 billion Rand, breeding forcolor
variance yields an average rate of return on investment (when sold though private auctions) of 50e1000 folds compared to
that generated by the pure breed individuals (Oliver, 2015). Such selective breeding and associated translocations are strictly
economic driven and not considered conservation-based behaviors (Hamman et al., 2003; Nel, 2015; Taylor et al., 2016).
1.7. Aim and objectives
Currently, there is no standardizing framework designed to prioritize decision-making process driven by conservation-
based objectives within the South African game industry (Gallo et al., 2009; Grobler et al., 2011). We hypothesize the
following: (1) that a substantial percentage of private game ranches have a need for a procedural framework to manage for
both wildebeest species, (2) that current translocation practices facilitate hybridization, (3) that both wildebeest species are
translocated across provincial borders, and within and amongst private ranches and PPAs, (4) that the wildlife ranching
industry poses risks to the state's ability to function as nature conservation facilitators, and (5) that wildebeest translocations
make up the lion's share of the market for wildlife translocations (and hence likely to continue).
This paper explores the delicate ecological interplay and conservation implications of the widely enforced management
practice of translocation. We put forward management and conservation recommendations based on three interlinked as-
pects of the wildebeest hybridization concern: biology, economics, and policy. We aim to improve the regulatory and pro-
cedural aspects of wildlife translocations by questioning whether or not conservation-based agendas are achieved via the
practice of translocation in its current form, and by illuminating the risk posed by the game industry on the state's ability to
act as conservation units as evident by dispersal patters analysis. Our objectives were: (1) to quantify the relative abundance
of private ranches that have both wildebeest species on their land, (2) to quantify the ratio of male blue wildebeest to male
black wildebeest translocated, (3) to analyse dispersal patterns across provincial borders, in addition to in and amongst
private wildlife ranches and provincial protected areas (PPAs), (4) to infer the risk of PPAs acting as stocking sources of
genetically pure wildebeest individuals, (5) to identify the percentage of economic revenue generated from wildebeest
translocations from all wildlife translocations, and (6) to put forward a conservation-based procedural framework that en-
hances management efﬁciency of translocations. We incorporate economic and social incentives as a strategy to increase
participation as well as address conservation implications related tothe black wildebeests' genetic integrityand the existence
of genetically pure black wildebeest populations.
2. Study area
We surveyed three PPAs and 273 privately owned game ranches that inhabit either black wildebeest, blue wildebeest, or
both black wildebeest and blue wildebeest populations. Data sampling occurred across the following ﬁve provinces: Limpopo,
Gauteng, Mpumalanga, Kwazulu-Natal (KZN), and The Free State (Map. 1). These provinces were selected because of the
distribution range of species. Additionally, this large geographical area allowed us to evaluate movement dynamics across
We documented 6929 legally permitted black and blue wildebeest translocations during ﬁve years. Legal translocations
entails that permits be issued as stipulated in provincial regulations, prior to the exchange of animals' ownership. For the
purposes of this study, we deﬁned translocation as an instance when: (1) a game capture company is hired to translocate
game, and (2) the process includes capture, relocation, and release. As such, we excluded sales conducted through public and
private auctions. We documented the following population variables of translocated individuals: their species (i.e., black or
blue wildebeest), sex, and age group (i.e., calves, yearlings, or reproductive age). We utilized SPSS software to analyze
interspecies male ratio. Location (i.e., origin and destination) was documented in order to analyse dispersal patters. Identi-
ﬁcation numbers were generated and given to ranches so that cross refrancing may be done conﬁdentially.
The authors emphasize that data collection focused solely on wildebeest species and population dynamics held on various
farms. Therefore, this study should not be considered as research involving human participants. As such, we did not seek
ethics approval concerning human participants. Research adheres to the highest of ethical codes of conduct provided by the
university. Speciﬁcally, double blinded identiﬁcation numbers were issued so that private ranch ownership remains conﬁ-
dential. This research was approved by the Tshwane University of Technology's ethics committee, by the research ethics
committee, speciﬁcally, by the Faculty Higher Degree Committee (FHDC). The double blinded identiﬁcation numbers were
issued in order to preserve the privacy of ranch ownership and were not strictly advised by the ethics committee.
Next, in order to access the need for a management plan for both species, we created a pie chart illuminating the per-
centage of private ranches that manage for blue wildebeest, black wildebeest, and both blue wildebeest and black wildebeest.
For the purpose of conservation based management recommendations and inferring the efﬁciency of current regulations,
we analyze wildebeest movement patters (i.e., introduction via translocation efforts) across provincial borders in South
Africa, in addition to among private ranches and PPAs. Dispersal patters data was analysed via the statistical program R.
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e58 49
Lastly, in order to asses the economic impact of the translocation of wildebeest translocation within the game ranching
industry, we quantiﬁed the proportional revenue generated from wildebeest translocations relative to revenue generated
from all game translocations. The selling price of animals themselves was used. We provide a detailed table that sums the
revenue generated from all game translocated, and from wildebeest translocated, per year. We then derived the percentage
revenue generated from wildebeest as a whole per year. By excluding private auctions as a data source, highly priced indi-
vidual animals (such as sable antelope (Hippotragus niger) or cape buffalo (Syncerus caffer)) were excluded. And as such, our
data scope incorporates the relevance of understanding the wildebeest market share within the wildlife ranching industry
and translocation practice. Lastly, although revenue generated from game translocation differs amongst species, we reasoned
that a high proportion of revenue generated from wildebeest is indicative of the plausibility that this practice is likely to
persist in its current form long-term.
Lastly, we put forward an applicable framework that involves an electronic procedure; individuals are tagged with an
electronic chip on their ear, the chip is then scanned and updated during each translocation, thereby documenting in-
dividual's location of origin and destination. Such information will enable private game ranchers and PPA managers to make
informed decisions based on the identiﬁed historical place of habitat of speciﬁc wildebeest (e.g., whether or not to accept
wildebeest individuals as founder populations etc'). It will additionally serve as the foundation for a national metapopulation
and monitoring management plan. We outline potential economic and social incentives designed to offset associated costs
and induce widespread implementation.
4.1. Wildebeest specie's abundance
Amongst the 273 private ranches surveyed, 9.2% managed for black wildebeest, 32.35% managed for blue wildebeest, and
58.45% managed for both black and blue wildebeest (Fig. 1).
4.2. Translocation efforts alter interspecies and intraspecies dynamics
Interspecies dynamics were analyzed based on the ratio of male wildebeest species translocated. The ratio of male blue
wildebeest to male black wildebeest translocated is skewed towards male blue wildebeest (Fig. 2). Additionally, complete
herds were not captured and translocated in full.
4.3. Wildebeest translocations occur across provincial borders
Black wildebeest were translocated to their extralimital habitat, and blue wildebeest were translocated into the black
wildebeest's core habitat (Fig. 3). After applying grouping around each province, a lack of geographic separation of the
network is illuminated; this indicates that translocations are occurring across provincial borders (Fig. 3).
Fig. 1. The relative abundance of black wildebeest and blue wildebeest species on private ranches in South Africa during 2006e2010. Solid black area indicates
private ranches that have black wildebeest (25 ranches, 9.2% of surveyed ranches), faded blue area indicates private ranches that have blue wildebeest (88
ranches, 32.35% of surveyed ranches), and shaded line area indicates the percentage of private ranches that have black wildebeest and blue wildebeest pop-
ulations (160 ranches, 58.45% of all ranches).
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e5850
4.4. Wildebeest translocations occur from multiple sources, in addition to within and amongst private ranches and PPAs
Almost all locations received both black wildebeest and blue wildebeest from more than one origin, with one destination
in particular accepting blue wildebeest from 34 different ranches over the course of ﬁve years (as indicated by thickness of
arrow linked to PPA in Fig. 4). In another example, a private ranch received 35 adult males, 98 adult females, and 55 juveniles
(22 males and 33 females) from 12 separate locations (Farm ID number 51, highlighted by a red triangle, Fig. 4).
Moreover, wildebeest were translocated within and amongst private ranches and PPAs (Fig. 4). All three PPAs, from various
provinces, accepted blue wildebeest and two also accepted black wildebeest from multiple sources (Fig. 4). In fact, the direct
location of origin of black wildebeest in one PPA was from three separate privately owned ranches across provincial borders,
which previously accepted black wildebeest fromfour private ranches (Farm ID number 300, highlighted by a purple triangle,
Fig. 4). Additionally, blue wildebeest originating in one protected area were dispersed into ﬁve different provinces (See ID
number 127, highlighted by a blue triangle, Fig. 4).
4.5. Black and blue wildebeest translocation account for a ﬁfth of the generated revenue
Wildebeest were found to be a substantial proportion of game translocations, accounting for between 15% and 34% of all
translocations, and generating an excess of 21 million Rand/year on average(Table 1). The highest costs of a single transaction
amounted to Rand335,901. This extensive market for wildebeest translocations, on average, accounted for 20.8% of revenue
generated from all game translocated (Table 1).
5.1. More than half of private game ranches encounter a need to manage for both blue wildebeest and black wildebeest populations
on their land
Currently, each province provides ranches with species ownership permits contingent on certain disclosures (e.g., a
species list, the abundance of suitable habitat, and adequate fencing). The purpose of such documentation is to ensure that
permits are not granted to owners that keep blue wildebeest and black wildebeests populations in sympatric conditions.
However, 58.45% of surveyed private ranches have blue wildebeest and black wildebeest populations on their land (Fig. 1).
Additionally, two of the three surveyed PPAs received both black wildebeest and blue wildebeest individuals (Fig. 4, See
below). Our ﬁndings indicate that sympatric conditions is widely practiced within South African private ranches, and thereby
suggest the need to reevaluate the efﬁciency of regulatory enforcement. Equally important, such a high percentage em-
phasizes the importance of better understanding of conservation implications that may result from wildebeest translocations,
Fig. 2. The ratio of male black wildebeest and male blue wildebeest translocated within and amongst private ranches and provincial protected areas (PAAs) in
South Africa. The sex and species of translocated wildebeest was documented during 20 06e2010. Y axis indicates the density as a percentage. All males are of
reproductive age. Zero on X axis means that number of blue wildebeest translocated is equal to number of black wildebeest translocated. Density is inferred by
blue males minus black male wildebeest. We factored in ratio rather than actual values as an accommodation to landowners who wished to understand this
relationship, as it is relevant to probability of hybridization occurrences. This is because hybridization is attributed to the displacement of male black wildebeest
by male blue wildebeest, followed by the mating of the later with cow black wildebeest. This ratio is skewed towards male blue wildebeest.
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e58 51
and thereby the importance of similar research. It additionally provides evidence of the need to equip managers with
standardizing conservation-based metapopulation management guidelines. The focus of such guidelines should be to
manage sympatric populations while accounting for risk posed by various hybridization facilitators.
The practice of translocation may alter mating encounters.
Blue and black wildebeest sympatric coexistence with minimal hybridization may be attributed to differential niche
occupation by family groups or herds (Codron and Brink, 2007). However, translocation practices do not preserve these
dynamics. We show that the ratio of male wildebeest translocated is skewed towards that of male blue wildebeest (Fig. 2). We
attribute this to the nature of the game translocation industry: (1) owners buy individuals (and not entireherds) from several
ranches with the management goal of outbreeding in order to increase genetic diversity, and (2) game captures are task with
capturing a certain number of individuals (a process that does not require the preservation of herd structures).
This additionally serves as evidence that sex composition was not preserved relative to locations of destination or origin.
Moreover, hybridization is documented to occur as a result of male blue wildebeest displacing male blackwildebeest, thereby
promoting mating encounters between bull blue wildebeest and cow black wildebeest. Grobler et al. (2011) suggest that this
displacement is possible due to the size of the bulls; bull blue wildebeest are bigger and will outcompete bull black males. As
such, we put forward that a skewed male ratio may induce interspecies mating encounters and hybridization. In addition,
maintaining such skewed male ratios is not favorable for the replacement of hybrid populations. Lastly, all surveyed locations
received and/or exported wildebeest individuals (Fig. 4). We found no indication that population dynamics (i.e., intraspecies
or interspecies sex composition) was preserved within populations that received or exported wildebeest. In fact, we provide
evidence of the contrary (Fig. 2).
As such, conservation implications are rooted in the risk of displacement in mating encounters and the low probability of
hybrid replacement in populations from which wildebeest individuals are either introduced or extracted. Thereby, providing
evidence for the creation of hybrid zones and the facilitation of hybridization. Such risks draw into question the efﬁciency of
Fig. 3. Black wildebeest and blue wildebeest movement across provincial borders in South Africa. Locations of origin and destination of translocated wildebeest
were documented during 2006e2010 and analyzed via a network coded in R. Overlapped networks were constructed when using gray circles to group locations
by province. Arrows indicate direction of movement (i.e., introduction of translocated wildebeest). Squares indicate PPAs and dots indicate private ranches. Colors
represent the provinces as follows: red (Limpopo), blue (Gauteng), green (Mpumalanga), purple (KwaZulu-Natal), and orange (Free State).
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e5852
the practice of translocation to serve as a conservation tool aimed at increasing genetic diversity while preserving the genetic
integrity of pure lineages. There is a need for additional research to explicitly quantify the link between sex and species
composition of receiving and exporting destinations, and hybridization rates.
5.2. Translocations from multiple origins, and across provincial borders, may introduce hybrid individuals of varying degrees into
Wildebeest were translocated within and across provincial borders, and from multiple sources (Figs. 3 and 4). We caution
against risks associated with artiﬁcially grouping wildebeest individuals from such various regions, subjected to different
provincial regulations and degrees of enforcement. The primary concern is the plausibility of introducing varying degrees of
hybrid and backcrossed individuals into parental pure breed populations across provincial borders, thereby inducing back-
crossing and genetic admixture.
Fig. 4. The extent of black wildebeest and blue wildebeest translocations within and among private game ranches and provincial protected areas in South Africa
during 2006e2010. Locations of origin and destination were classiﬁed as private ranches or provincial protected areas for translocated wildebeest during
2006e2010 and analyzed via a network coded in R. Arrows document directions of movement (i.e., indicative of the direction of introduction). Blue arrows
indicate blue wildebeest movement, whereas black arrows indicate black wildebeest movement. Arrow thickness is proportional to the amount of individuals
translocated. Ranches introduced blue and black wildebeest from more than one source of origin. Dotted colors and location ID ranges represent the provinces as
follows: red and >99 (Limpopo), blue and 100e199 (Gauteng), green and 200e299 (Mpumalanga), purple and 300e399 (KwaZulu-Natal), and orange and
400e499 (Free State). Circles represent privately owned ranches and squares represent PPAs. Lastly, farm IDs 51, 300, and 127 are additionally discussed in the
text and therefore highlighted by colored triangles corresponding to their provincial location.
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e58 53
5.3. Current translocation practices may facilitate hybridization
Genetic markers that differentiate between black, blue, and hybrid wildebeest have not yet been identiﬁed (Ackermann
et al., 2010; Corbet and Robinson, 1991; Corbet et al., 1994; De Klerk, 2008; Grobler et al., 2005, 2011). However, a number
of prominent papers have indicated the occurrence of hybridization amongst black wildebeest and blue wildebeest, and the
production of fertile offspring based on morphological characteristics abnormal of parental taxa such as horn shape ab-
normalities in horn shape has previously served to identify hybrids populations prior to two culling expeditions on national
land (Ackermann et al., 2010; Grobler et al., 2005). Our ﬁndings show that there is no current standardized procedure that
documents the genetic purity of translocated wildebeest, and that herd composition is not preserved in introduction loca-
tions. We additionally provide noble evidence that illuminates translocation practices across provincial borders and between
and among private ranches and national parks. Such data provides reasoning for two underlying assumptions: (1) wildebeest
hybridization has occurred on private ranches, and (2) the lack of genetic testing of translocated individuals creates the
possibility that a proportion of these are hybrids. We therefore extrapolate that translocation efforts may foster the intro-
duction of hybrid individuals and that management guidelines should be created in order to identify and counteract such
5.4. The impact of the private game ranching industry on the state's ability to act as nature conservancies
The black wildebeest has previously experienced two population bottlenecks (Grobler et al., 2005). Conservation concerns
about bottlenecks steam from a decreased ability to adapt to stochastic events. This is because genetic drift occurs at an
accelerated rate in small populations and results in decreased genetic variance. A common management solution that ad-
dresses genetic drift concerns aims to counter bottlenecks by relocating individuals into existing populations in order to
provide population reinforcement, thereby increasing genetic diversity (Rhymer and Simberloff, 1996). Similar conservation-
driven translocations re-establish, recolonize and replace populations by founder effects (Rhymer and Simberloff, 1996). A
founder effect occurs when a new population is started by a few members. Often times, these founders will be provided by a
source likely to not have hybrids in their population.
Map 1. Map of South African provinces. Surveyed locations are illustrated by circles and within the following provinces: Limpopo, Gauteng, Mpumalanga,
Kwazulu-Natal, and The Free State.
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e5854
Alarmingly, despite provincial regulations prohibiting the sympatric stocking of both wildebeest species on national land,
2/3 surveyed PPAs stock both black and blue wildebeest population on their land (Fig. 4). Our ﬁndings we show that black
wildebeest and blue wildebeest are translocated across provincial borders (Fig. 3), in addition to within and between the
private ranching industry and PPAs (Fig. 4). In fact, PPAs accepted individuals from multiple private ranches which previously
accepted individuals from other multiple sources (Fig. 4). This raises concerns about whether or not national parks within
South Africa are able to provide genetically pure black wildebeest to act as population founders on private lands or on other
national parks. We therefore draw into question the PPA's ability to act as conservation units and serve as stocking sources.
Risks posed by the game industry are underlined by the inability to identify all historical habitat conditions of each in-
dividual's historical locations. Consequently, we draw into question the functionality of PPAs as conservation units, and their
ability to act as stocking sources or founder populations. Such concerns are future illuminated when considering the genetic
status of the black wildebeests within its contextual history; hybrid populations have been previously culled on at least two
PPAs (Ackermann et al., 2010; Grobler et al., 2005).
Such evidence suggests the need to consider potential outbreeding depression, hybridization, backcrossing, and genetic
swamping, and their implications, within existing populations whose founders originated in such PPAs. We caution that at a
minimum, PPAs may suffer economic lose due to a reputational risk from stocking both species on their land. In a worse case
scenario, all black wildebeest populations in South Africa incorporate varying degrees of hybrid individuals.
5.5. The economic viability of wildebeest translocations
Black wildebeest and blue wildebeest translocations were responsible for generating on average 20.8% (and up to 34%) of
the economic revenue resulting from all translocated game (Table 1). Attributing such a substantial revenue to wildebeest
translocations suggest that its role as a management practice is likely to persist long term in its current form, and that
wildebeest populations are likely to continue to be impacted by it. This is concerning in light of our evidence warning against
deleterious conservation implications which include hybrid facilitation, outbreeding depression, and genetic swamping. As
such, translocation practices are possibly positioning the black wildebeest in dangerof genetic admixture and swamping, and
are likely to worsen if no intervention is taken.
5.6. Strategies to decrease possible hybridization
Our data indicates a fundamental need to consider whether or not conservation-based management goals for wildebeest
populations are achieved via the current practice of translocations. Primarily conservation-based objective are twofold: (1) to
avoid inbreeding (and inbreeding depression by which ﬁtness is reduced due to limited allelic diversity), and (2) to promote
outbreeding (i.e., increased allelic diversity). However, the process of translocation itself induces overlap (Fig. 1), alters and
shifts interspecies and intraspecies sex composition (Fig. 2), and introduces individuals within and across provincial borders,
from multiple sources, and within and amongst private ranches and PPAs (Figs. 3 and 4). Finally, our ﬁndings suggest that this
process is likely to be enforced as a management practice in the near future (Table 1).
The lack of accountable documentation of variables in places of historical origins (i.e., sympatric/parapatric/allopatric
populations, known hybrids, etc') implies the risk of introducing individuals that are hybrids and/or backcrossed individuals
of varying degrees to PPAs and private ranches. This may facilitate unwanted result such as outbreeding depression, which
may manifest in one of two negative effects: (1) decreased ﬁtness compared to parental populations, and/or (2) accelerate the
evolution of locally adapted gene complexes (Allendorf et al., 2001; Arnoldand Martin, 2010; Burke and Arnold, 2001). In both
cases, offspring are likely to face signiﬁcant disadvantages when translocated back into parental environments. A primary
example of such decreased ﬁtness in wildebeest hybrids is horn shape and skull shape (De Klerk 2008).
6. Management implications
6.1. The entire black wildebeest species in South Africa may be composed of varying degrees of hybrids
We show that there is a non-zero probability that the genetic integrity of the black wildebeest has been jeopardized and
that black wildebeest populations are hybrids of varying degrees. Especially because farmers are largely unaware of the
Revenue generated in South Africa from all game translocations and wildebeest translocations from 2004 to 2008. Revenue is documented per million South
Revenue generated from WB
% of revenue generated from WB
Revenue generated from all game
2010 16.8 17.99 93.7
2009 18.9 15.65 120.8
2008 18.5 17.27 107.4
2007 19.9 18.48 107.8
2006 32.8 34.83 94.3
N. Benjamin-Fink, B.K. Reilly / Global Ecology and Conservation 12 (2017) 46e58 55
degree to which introduced wildebeest are pure breed. Moreover, the black wildebeest recently survived two genetic bot-
tlenecks (Brink, 1993), supporting the strong possibility that all populations are genetically interlinked (particularly without
knowledge concerning founder origin of these populations). Careful consideration of our ﬁndings within the context of
genetic laws applicable to ﬁnite and isolated populations (typical of South African ranches) suggest plausible hybridization,
backcrossing, introgression, outbreeding depression and genetic swamping. And that in fact, translocation efforts may play a
signiﬁcant role in accelerating unwanted evolutionary processes (Grobler et al., 2005, 2011). We caution against deleterious
eventuality; pure black wildebeest will eventually become genetically extinct.
Our ﬁndings provide evidence that the wildebeest genetic heritage will beneﬁt from an applicable trans-provincial
conservation-based framework that will serve as a metapopulation management plan. Due to the lack of genetic markers
distinguishing parental species, we suggest the need for a standardizing procedure that documents wildebeest individuals'
historical locations of origin. Such information will serve as inference for its genetic status and enable the cross reference of
origin and destination of individual wildebeest prior their acceptance in private ranches or PPAs.
6.2. The Ecological Sustainable Network (ESN); an accountable and transparent recommended wildebeest procedural-framework
We coin the concept of an Ecological Sustainable Network (ESN), a procedural framework that documents the place of
origin and destination of each individual wildebeest, thereby increasing regulatory efﬁciency by standardizing accountability
and transparency. Individuals will be tagged with an electronic chip. Chips will be scanned and updated thereby documenting
the following information: (1) historical ownership (i.e., locations of origin and destinations), (2) habitat conditions (i.e.,
sympatric, parapatric, or allopatric conditions), and (3) the occurrence of known hybrids. This information will serve as a
determining factor for informed decisions by private ranches and PPA managers as to whether or not to introduce speciﬁc
wildebeest onto their property.
Implementing the ESN is easy because it follows the three steps currently employed during translocation practices: (1)
during capture, man power and needed resources will be assessed, (2) during the handling phase, wildebeest individuals will
be tagged with an electronic chip on their ear (or scanned and updated, in cases where wildebeest already have electronic
chips), and (3) during the release phase habitat conditions will be documented. The ESN additionally establishes a platform
for collaborative efforts by creating a network amongst managers. Moreover, we recommend that the initial associated costs
(i.e., mass tagging) will be the responsibility of provincial ofﬁces, while the costs associated with updating (i.e., updating
chips, and tagging new born individuals) will be the responsibility of private ranch owners. The strategy of such diminished
economic responsibility has previously shown to increase implementation. Furthermore, ESN supports the private owners'
willingness to pay and willingness to conserve, once the economic incentives are reduced and removed (as suggested by
Ramsdell, 2014; Sonnekus and Bretyenbach, 2001; Sorice et al., 2013). An example of such implications is that private
ranchers from the Lowveld area may have equal proﬁt margins to those from the Highveld.
A similar framework exists aimed at preserving the genetic integrity of the bontebok. Hybridization resulting from
breeding between bontebok and blesbok has been the focus of a number of studies. Currently, there are genetic markers that
identify bontebok, blesbok, and their hybrids (Van Der Walt et al., 2013). This enables the translocation of the bontebok to be
optimally managed; a DNA-based genetically purity certiﬁcate is required to accompany bontebok translocations (Van Wyk
et al., 2013, 2017).
Lastly, enhancing management transparency and accountability will position South Africa at the frontier of wildlife
conservation. Similarly, our suggested protocol may be modiﬁed to address hybridization in other ungulate species whose
genetic markers have not been identiﬁed, and serve as an effective conservation tool for wildlife managers in countries where
current translocation has similar implications.
We seek to acknowledge T. Klagsbrun, S, Vrahimis, S. Klippies, and NZGB for ﬁeld assistance, coordination and insights, and
E.M. Lind for assistance in data analysis as well as various colleagues for insightful comments and Conservation Beyond
Borders for support. We also extend special thanks to the many South African conservation scientists, conservationists, and
game stakeholders that willingly donated their time and information. Thanks to anonymous reviewers whose comments
contributed to the ﬁnal version of this paper.
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