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Conservation implications of wildlife translocations; The state's ability to act as conservation units for wildebeest populations in South Africa


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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 divergence , 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 provincial lines, (4) wildebeest are introduced to within and amongst the private and commercial 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 practices worldwide.
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Original Research Article
Conservation implications of wildlife translocations; The
state's ability to act as conservation units for wildebeest
populations in South Africa
Nicole Benjamin-Fink
, 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
article info
Article history:
Received 30 June 2017
Received in revised form 18 August 2017
Applied ecology
Ecological Sustainable Networks (ESN)
Hybrid zones
Wildlife translocation
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 efciency. 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-
tices worldwide.
©2017 The Authors. Published by Elsevier B.V. This is an open access article under the CC
BY license (
*Corresponding author.
E-mail addresses: (N. Benjamin-Fink), (B.K. Reilly).
Contents lists available at ScienceDirect
Global Ecology and Conservation
journal homepage:
2351-9894/©2017 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (
Global Ecology and Conservation 12 (2017) 46e58
1. Introduction
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 nite 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 protability 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
of private
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 dene translocation
as the mediated movement of living organisms from any source (privately managed or wild), with release in another. This
oversimplied denition 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 denition of the
concept of national parks (i.e., a state) acting as conservation units and dene 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 (
content/105/S1/795.full,Anderson, 1949). This may occur as a result of articial 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 tness (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 articially 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 articial over-
lap which may result from translocations. This, in turn, may create articial bottlenecks in nite populations, thereby pro-
moting and sustaining hybridization (Arnold, 1992). For these reasons, translocation to closed and nite 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 rst 2 countries, extensive translocation efforts played
signicant 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
ow when the 2 species are articially 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 rst 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 signicant 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 ve subspecies of blue wildebeest throughout the South African continent, the C.t.taurinus is the solely blue
wildebeest inhabiting South Africa and is articially 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 efciency 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 ve 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
provincial borders.
3. Methods
We documented 6929 legally permitted black and blue wildebeest translocations during ve 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 dened 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-
cation numbers were generated and given to ranches so that cross refrancing may be done condentially.
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. Specically, double blinded identication numbers were issued so that private ranch ownership remains con-
dential. This research was approved by the Tshwane University of Technology's ethics committee, by the research ethics
committee, specically, by the Faculty Higher Degree Committee (FHDC). The double blinded identication 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 efciency 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 quantied 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 identied historical place of habitat of specic 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. Results
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 ve 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 ve different provinces (See ID
number 127, highlighted by a blue triangle, Fig. 4).
4.5. Black and blue wildebeest translocation account for a fth 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. Discussion
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 ndings indicate that sympatric conditions is widely practiced within South African private ranches, and thereby
suggest the need to reevaluate the efciency 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 efciency 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
receiving populations
Wildebeest were translocated within and across provincial borders, and from multiple sources (Figs. 3 and 4). We caution
against risks associated with articially 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 classied 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 identied (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 ndings 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 ndings 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 tness 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 ndings 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 tness 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 signicant disadvantages when translocated back into parental environments. A primary
example of such decreased tness 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
Table 1
Revenue generated in South Africa from all game translocations and wildebeest translocations from 2004 to 2008. Revenue is documented per million South
African Rand.
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 ndings within the context of
genetic laws applicable to nite 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
signicant 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 ndings provide evidence that the wildebeest genetic heritage will benet 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 efciency 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 specic
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 ofces, 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 prot 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 certicate 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 modied to address hybridization in other ungulate species whose
genetic markers have not been identied, 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 eld 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 nal version of this paper.
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... This not clear!! Negative consequences may be in one of the following forms: hybridization and introgression, genetic bottlenecks, inbreeding or outbreeding depression, and shifts in local or regional ecological equilibrium. As a result of such artificial enclosures, game ranch owners are faced with the need to intensely 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) [7]. As such, increasing risks posed to endangered mammals. ...
... South Africa experienced an annual increase of 5.6% in land used for wildlife ranching during 1991-2000 [7]. Diminished economic profitability of livestock, coupled with an increase in stock theft, resulted in a shift from traditional agriculture and cattle farming to ungulate ranching within the private ranching industry. ...
... This sector is emerging in its scope and has the capacity to produce 50-1000% ROIs in full. A primary example is the breeding of a bull golden wildebeest (Connochaetes) with a cow blue wildebeest (Connochaetes taurinus); the offspring is referred to as "Split Golden" and were sold in private auctions during 2014 for 513, 137 rand (a 541% increase in selling price compared to 80,000 rands in 2004) [7]. These Split Golden wildebeests are then mated with golden bulls (i.e., backcrossed) to facilitate the expression of the recessive allele for a golden color coat. ...
... They are sequential to other problems, lack clear solution, and may be easily misrepresented as an isolated concern [2]. Ecological and environmental wicked challenges are often rooted in spillover effects from biodiversity, ecosystem services, social and environmental aspects, management implications, and economic profitability goals [3][4][5][6]. ...
... The wicked case Study of hybridization Hybridization is one of the primary wicked problems of our time [4]. Accelerating at an alarming rate, it is responsible for the genetic extinction of endangered and indigenous species worldwide (Box 1). ...
... Active management operations are often valuable from the conservation point of view, but some of them have been inappropriate or even damaging to the genetic integrity of autochthonous populations of particular species (cf. [27] for mitigation translocation cases), such as in the case of the wildebeest (see [28][29][30]), or whole communities (for evaluation of ungulate translocations, especially in Southern Africa, see [21,31,32]). ...
Full-text available
Ecotourism can fuel an important source of financial income for African countries and can therefore help biodiversity policies in the continent. Translocations can be a powerful tool to spread economic benefits among countries and communities; yet, to be positive for biodiversity conservation, they require a basic knowledge of conservation units through appropriate taxonomic research. This is not always the case, as taxonomy was considered an outdated discipline for almost a century, and some plurality in taxonomic approaches is incorrectly considered as a disadvantage for conservation work. As an example, diversity of the genus Giraffa and its recent taxonomic history illustrate the importance of such knowledge for a sound conservation policy that includes translocations. We argue that a fine-grained conservation perspective that prioritizes all remaining populations along the Nile Basin is needed. Translocations are important tools for giraffe diversity conservation, but more discussion is needed, especially for moving new giraffes to regions where the autochthonous taxa/populations are no longer existent. As the current discussion about the giraffe taxonomy is too focused on the number of giraffe species, we argue that the plurality of taxonomic and conservation approaches might be beneficial, i.e., for defining the number of units requiring separate management using a (majority) consensus across different concepts (e.g., MU-management unit, ESU-evolutionary significant unit, and ECU-elemental conservation unit). The taxonomically sensitive translocation policy/strategy would be important for the preservation of current diversity, while also supporting the ecological restoration of some regions within rewilding. A summary table of the main translocation operations of African mammals that have underlying problems is included. Therefore, we call for increased attention toward the tax-onomy of African mammals not only as the basis for sound conservation but also as a further opportunity to enlarge the geographic scope of ecotourism in Africa.
... The increase in the number of animals threatened by extinction is leading researchers worldwide to increase their efforts to conserve animal species [1]. Among the conservation initiatives, habitat protection and captive breeding for animal translocation/reintroduction are the most applicable since they have direct implications for the survival of the species in a specific location [2][3][4][5]. Animal introduction (i.e., the intentional movement and release of an organism outside its indigenous range), reintroduction (i.e., the intentional movement and release of an organism inside its indigenous range from which it has disappeared), and reinforcement (i.e., the intentional movement and release of an organism into an existing population of conspecifics) are types of animal translocation (all definitions taken from [6]). ...
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The number of animal species threatened with extinction are increasing every year, and biologists are conducting animal translocations, as one strategy, to try to mitigate this situation. Furthermore, researchers are evaluating methods to increase translocation success, and one area that shows promise is the study of animal personality. Animal personality can be defined as behavioral and physiological differences between individuals of the same species, which are stable in time and across different contexts. In the present paper, we discuss how animal personality can increase the success of translocation, as well as in the management of animals intended for translocation by evaluating personality characteristics of the individuals. Studies of the influence of birthplace, parental behavior, stress resilience, and risk assessment can be important to select the most appropriate individuals to be released. Finally, we explain the two methods used to gather personality data.
... This will be even more relevant as interactions between humans and wildlife are increasing across Africa (Hoare 1995;Tchamba 1995;Madden 2004). Furthermore, conservation research in Africa plays a significant role in generating knowledge and strategies for wildlife conservation (Child et al. 2012;de Pinho et al. 2014;Benjamin-Fink and Reilly 2017) in a diversity of landscapes (Myers et al. 2000). Understanding how these key terms are currently characterized and identifying gaps in usage, definition, and measurement is essential to make sure these terms are applied consistently in conservation in Africa and across the globe. ...
The terms ‘coexistence’, ‘tolerance,’ and ‘acceptance’ appear frequently in conservation literature, but lack consistent characterization, making them difficult to apply across intervention frameworks. This review aims to describe the common characterizations of these three terms using Africa-based research as a case study. Through systematic lexical searches, we identified 392 papers containing one or more of the three terms. We assessed their usage, definition, and measurement (or lack thereof) in wildlife conservation. Coexistence was used in 46% of papers, but was defined in only 2% and measured in 4%. Tolerance and acceptance were used in 63% and 61% of the papers in which they appeared, respectively, defined in 4% and 2%, and measured in 19% and 5%. These results confirm the lack of clear understanding of these concepts and evidence the need for a precise lexicon. This would allow conservationists to cohesively describe their work and increase replicability of research across contexts.
... Among these are ungulates and red-legged partridge (Alectoris rufa) in Italy (Randi, 2005;Baratti et al., 2005), lesser white-fronted geese (Anser erythropus) and grey partridge (Perdix perdix) in Scandinavia (Ruokonen et al., 2000;Nijman et al., 2010;Liukkonen, 2006), black wildebeest (Connochaetes gnou) and blue wildebeest (C. taurinus) in South Africa (Benjamin-Fink and Reilly, 2017), and slow lorises (Nycticebus spp) in Indonesia (Nekaris and Starr, 2015). Such practises can cause ecological and taxonomic havoc. ...
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The illegal cage bird trade is increasingly recognised as a major impediment to the survival of a large number of songbirds. Indeed some bird species are now more common in captive private hands than they are in the wild. This includes the black-winged mynas (Acridotheres melanopterus, A. tricolor and A. tertius), three species of Critically Endangered songbirds endemic to Indonesia. Only 20 years ago these species were not considered globally threatened but high levels of trapping from the wild for the largely domestic cage bird trade has brought all three species to the brink of extinction. It is estimated that less than 500 black-winged mynas remain in the wild. Here we investigate the trade in black-winged mynas, online and in bird markets, and make an assessment of the role captive breeding played in the conservation and management of the species over the period 2009–2018. We found prices peaked in 2014 at ∼ US$140 (∼75% of the monthly minimum wage) per bird and have subsequently gone down to US$85 (now ∼33% of the minimum wage), possibly indicating higher supply and lower demand. In 2015–2018 we surveyed seven bird markets in western Java and in 127/145 visits observed 1253 black-winged mynas for sale. Turnover was high, with ∼50% of birds sold after one week upon arrival in the market. We estimate that between 1300 and 2300 mynas (retail value ∼ US$170,000) are sold annually in these seven bird markets. Few birds had closed leg-rings, but were in all likelihood a combination of captive-bred, first-generation captive-born and wild-caught individuals; some appeared to be crossbreeds between the three recognised species. With additional bird markets in Java and Bali and a thriving online trade, we estimate that the number of black-winged mynas in private ownership in Indonesia is in the order of 40,000 birds. Without proper registration and regulation in the trade of captive-bred mynas, even a small amount of wild-caught birds entering this now substantial trade will act as a serious impediment to the conservation of black-winged mynas. With the species already being ecologically extinct, we anticipate that it soon will join the ranks of species like Père David's deer Elaphurus davidianus and scimitar-horned oryx Oryx dammah that are extinct in the wild but that have captive populations in the tens of thousands. In order to prevent the imminent extinction of black-winged mynas in the wild, we recommend (1) that the Indonesian authorities invest in more effective law enforcement and prosecution of lawbreakers; (2) establishing a multi-stakeholder three species black-winged myna management plan, in which commercial captive breeders participate; and (3) better coordination of reintroduction programmes.
... Historically, the main threats to black wildebeest included hunting pressure, habitat loss and disease outbreaks (Vrahimis 2013). However, the only current significant threat to this species is hybridization with its congener (Benjamin-Fink and Reilly 2017;East 1999;Grobler et al. 2011). ...
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Introgressive hybridization poses a threat to the genetic integrity of black wildebeest (Connochaetes gnou) and blue wildebeest (Connochaetes taurinus) populations in South Africa. Black wildebeest is endemic to South Africa and was driven to near extinction in the early 1900s due to habitat destruction, hunting pressure and disease outbreaks. Blue wildebeest on the other hand are widely distributed in southern and east Africa. In South Africa the natural distribution ranges of both species overlap, however, extensive translocation of black wildebeest outside of its normal distribution range in South Africa have led to potential hybridization between the two species. The molecular identification of pure and admixed populations is necessary to design viable and sustainable conservation strategies, since phenotypic evidence of hybridization is inconclusive after successive generations of backcrossing. The aim of this study was to assess levels of hybridization in wildebeest using both species-specific and cross-species microsatellite markers. Black wildebeest (157) and blue wildebeest (122) from provincial and national parks and private localities were included as reference material, with 180 putative hybrid animals also screened. A molecular marker panel consisting of 13 cross-species and 11 species-specific microsatellite markers was developed. We used a Bayesian clustering model to confirm the uniqueness of blue- and black wildebeest reference groups, assign individuals to each of the two clusters, and determine levels of admixture. Results indicated a clear partition between black wildebeest and blue wildebeest (the average proportions of membership to black wildebeest and blue wildebeest clusters were QI = 0.994 and QI = 0.955 respectively). From the putative hybrid samples, only five hybrid individuals were confirmed. However, high levels of linkage disequilibrium were observed in the putative hybrid populations which may indicate historical hybridization. Measures of genetic diversity in the black wildebeest populations were found to be lower than that of the blue wildebeest. The observed lower level of genetic diversity was expected due to the demographic history of the specie. This study will make a significant contribution to inform a national conservation strategy to conserve the genetic integrity of both species.
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In this letter, we revisit a study we published in 2017, following comment in a paper by Marchesini et al. published in this volume. We provide some further analyses that help us to reinforce the original conclusions of our earlier paper, and to address the points raised by Marchesini et al. We conclude that the concerns raised in their review do not alter the inference we presented earlier, and we identify issues with analyses presented by Marchesini et al. that limit their utility. The key points of inference remain that this species in Europe shows remarkably low levels of diversity within populations and strong structure among populations which can be explained by a combination of natural and anthropogenic processes.
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Hybridization is threatening many endangered, indigenous and rare mammal species worldwide. At the same time, wildlife managers are forced to make management decisions with scarce data and limited resources. Anthropogenic-driven landscape change is expanding at an exponential rate and may play a significant role in increasing hybridization rates. Understanding the ecological context in which hybridization occurs is crucial to increasing conservation and management effectiveness on both regional and global scales. This review explores the biological and spatial means by which anthropogenic landscape disturbances, often resulting from management practices, affect mammalian hybridization rates. We caution that the literature provides evidence that shifts in habitat characteristics may be attributed to accelerating anthropogenic disturbance, and is especially problematic for closely related mammalian species that recently diverged, and therefore have the potential to hybridize when ecological conditions are favorable. While direct impacts lead to the expansion or contraction of the distribution of wild species, indirect impacts result in expansion of the distribution of non-native species. Additionally, management and conservation guidelines strongly depend on the degree to which anthropogenic-driven landscape change alters ecological processes and the time scale at which hybridization occurs; anthropogenic activities induce hybrid-ization by directly and indirectly changing species composition in a given landscape. Therefore, we suggest that conservationists and wildlife decision-makers should approach hybridization as a global threat to biodiversity with site-specific solutions following our suggested guidelines. We provide a summary of the impact of various management practices on hybridization and put forward a decision tree for management. Effective management and conservation objectives should focus on reducing anthropogenic drivers that promote hybridization and introgression rates. Integrating the implications of ecological shifts due to anthropogenic landscape change into species conservation and landscape management guidelines has the potential to increase conservation and management effectiveness in light of increasing environmental change and degradation worldwide.
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Anthropogenic hybridization is an increasing conservation threat worldwide. In South Africa, recent hybridization is threatening numerous ungulate taxa. For example, the genetic integrity of the near‐threatened bontebok (Damaliscus pygargus pygargus) is threatened by hybridization with the more common blesbok (D. p. phillipsi). Identifying nonadmixed parental and admixed individuals is challenging based on the morphological traits alone; however, molecular analyses may allow for accurate detection. Once hybrids are identified, population simulation software may assist in determining the optimal conservation management strategy, although quantitative evaluation of hybrid management is rarely performed. In this study, our objectives were to describe species‐wide and localized rates of hybridization in nearly 3,000 individuals based on 12 microsatellite loci, quantify the accuracy of hybrid assignment software (STRUCTURE and NEWHYBRIDS), and determine an optimal threshold of bontebok ancestry for management purposes. According to multiple methods, we identified 2,051 bontebok, 657 hybrids, and 29 blesbok. More than two‐thirds of locations contained at least some hybrid individuals, with populations varying in the degree of introgression. HYBRIDLAB was used to simulate four generations of coexistence between bontebok and blesbok, and to optimize a threshold of ancestry, where most hybrids will be detected and removed, and the fewest nonadmixed bontebok individuals misclassified as hybrids. Overall, a threshold Q‐value (admixture coefficient) of 0.90 would remove 94% of hybrid animals, while a threshold of 0.95 would remove 98% of hybrid animals but also 8% of nonadmixed bontebok. To this end, a threshold of 0.90 was identified as optimal and has since been implemented in formal policy by a provincial nature conservation agency. Due to widespread hybridization, effective conservation plans should be established and enforced to conserve native populations that are genetically unique.
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Over the last two decades across eastern and southern Africa a range of novel institutional arrangements for tourism, conservation and development have emerged. In this chapter we clarify how we attempt to understand these innovative institutional arrangements and explain the key questions that run through the chapters of this book as well as their relevance. We further elaborate on how different contemporary institutional arrangements are framed from instrumentally and critically oriented views and clarify the middle position that we take by providing a stage for reflection on these different views. The chapter closes with a concise outline of the contribution each chapter makes to this book.
The territorial behaviour of the black wildebeest was studied from 1968 to 1970 in the Willem Pretorius Game Reserve and other reserves in South Africa.Territoriality is a prerequisite for reproduction. Non-territorial males are barred from partaking in the rut. Territorial males may occupy a territory from their fourth year on, but the majority of territorial owners were five years old or more.Territorial males formed a network of territories, although single territorial males could also be observed. The spacing between individuals varied between 180-450 m in the Willem Pretorius Game Reserve; spacing of territorial black wildebeest as well as the number of territorial males depend on the density of the population.Territorial black wildebeest showed a strong and lasting attachment to their territories. The species was territorial throughout the year although absenteeism became prevalent after the rut.Territorial black wildebeest advertised their territories and defended them against intruding conspecific males. Advertising behaviour included demonstrative-threat advertising derived from marking as well as acoustic, static-optic and dynamic-optic advertising. The species did not demarcate a territory with either faeces or urine, or pre-orbital or interdigital glands.The encounters of territorial males were governed by highly ritualised motor patterns (Challenge Ritual). The ritual includes both aggressive and non-aggressive behaviour patterns.The Challenge Ritual ensures that only the fittest male can maintain his territory and thus partake in the rut. Furthermore, it serves to “satisfy“ the social needs of the solitary bulls in the highly gregarious species. Other functions of territorial behaviour include the separation of sexes and the spacing out of the population.The known agonistic and territorial behaviour of the various species of the tribe Alcelaphini is compared and discussed. The differences and similarities in behaviour do not follow the taxonomic separation of the various species but are indicative of different habitats.
Black wildebeest fossils from the interior of South Africa and the Cape coastal zone are compared to modern specimens in order to trace the pattern of morphological change and the distribution of the species through time. Measurements taken on selected postcranial skeletal elements suggest that the evolution of the black wildebeest was marked by a general reduction in body size. It appears that the evolution of Connochaetes gnou from a blue wildebeest-like (C. taurinus) ancestor is best documented in areas to the south of the Vaal River. -from Author