Next Article in Journal
Deposition of Potentially Toxic Metals in the Soil from Surrounding Cement Plants in a Karst Area of Southeastern Brazil
Previous Article in Journal
Animals Traded for Traditional Medicine Purposes in the Kumasi Central Market, Ghana: Conservation Implications
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:

Taxonomy and Translocations of African Mammals: A Plea for a Cautionary Approach

Spartaco Gippoliti
Jan Robovský
2,3 and
Francesco M. Angelici
IUCN/SSC Primate Specialist Group, Viale Liegi 48A, 00198 Rome, Italy
Department of Zoology, Faculty of Science, University of South Bohemia, Branišovská 1760, 37005 České Budějovice, Czech Republic
Liberec Zoo, Lidové sady 425/1, 46001 Liberec, Czech Republic
Italian Foundation for Vertebrate Zoology (FIZV), Via Marco Aurelio 2, 00184 Rome, Italy
Author to whom correspondence should be addressed.
Conservation 2021, 1(2), 121-136;
Submission received: 6 May 2021 / Revised: 27 May 2021 / Accepted: 8 June 2021 / Published: 14 June 2021


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 This is not always the case, as taxonomy was considered an outdated discipline for almost a century, and some plurality in taxonomic approaches is incorrectly considered as a disadvantage for conservation work.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 taxonomy 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.

1. Introduction

When dealing with Africa’s biodiversity, we should not be tempted to fall into the belief of studying and conserving a piece of Eden that escaped the ruinous consequences of human impact. This is not certainly true, but see [1,2], yet wildlife diversity and abundance had—and still has—an obvious role in shaping Western perceptions of Africa as a unique place on Earth [3,4].
African biomes are also a unique laboratory for scientific research that explores the forces that governed evolutionary history. Dealing with such diverse issue as humans’ evolutionary history or the relationship between tectonic activity and cichlids’ speciation [5], Africa appears as a unique setting to reach a better knowledge of how biodiversity developed. Moreover, African mammals have been a classic subject for scientific inquiries dealing with evolution for more than a century now, e.g., [6,7,8,9,10,11,12,13,14,15,16,17].
Mammal diversity, particularly the richness of relatively large-sized species, fueled at first a growing interest in Africa by hunters from all over the world. Later, a conservation movement developed (aside from a few pioneers such as Carl Akeley), mainly influenced by those researchers—such as George B. Schaller and Ian Douglas-Hamilton—studying charismatic species such as African elephants, lions, gorillas and chimpanzees. The same species and African wildlife biomass and diversity are now the target of a multi-millionaire tourism industry that makes wildlife a highly profitable business for private operators and African countries [18]. This has led to an increased role in active management of some mammal species, including a growing occurrence of translocations as a means to repopulate or restock protected areas that have lost their native stock of some of the most charismatic species [19,20,21,22], to realize the optimal management for threatened taxa (e.g., [23,24] in the case of the Cape mountain zebra) or for the ecological restoration of particular regions (e.g., [25,26]). 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]).
In the present contribution, we aimed to critically review translocations of some mammals in Africa as a conservation tool, partly using giraffes as a case-study because of the current progress in understanding their diversification across Africa and emphasizing some causes of concern relating the possible negative outcomes for the conservation of evolutionary history in a unique continent. Following an increasing emphasis on financial viability, many extralimital species—i.e., species that historically did not occur in an area—or stocks of atypical phenotypes (under “intentional genetic manipulation” [21]) were introduced into private and public reserves to increase public experiences with the intention of increasing ecotourism attractions [33]. Although translocations are not a totally new tool in African conservation, it seems that many current projects are being realized primarily for financial reasons rather than conservation considerations. Another factor that often determines and influences the translocations is the need to create private game reserves dedicated to trophy hunting. This phenomenon is especially widespread in South Africa [34], and, often, the purpose of increasing income is to the detriment of conservation because moving species and subspecies well outside their original ranges increases the risk of hybridization between closely related species or subspecies [35,36]. A similarly questionable type of operation is to create fenced private reserves with the aim of attracting tourists to observe animal species, especially large mammals, including those that had never been locally present there in historic times.
According to [37], there are at least seven types of translocation for which conservation is not the primary aim (note that species conservation may be an associated aim and protection of individual animals of threatened species may be a primary aim): non-lethal management of problem animals, commercial and recreational, biological control, aesthetic, religious, wildlife rehabilitation, and animal rights activism.

2. What Do We Know about Large Mammal Diversity?

Wildlife managers, like most tourists, are convinced that our knowledge of large African mammals is more than satisfactory, as demonstrated from the large number of books existing on the subject e.g., [38,39]. Field guides, in particular, are suspected to vehicle an assuring view concerning our taxonomic knowledge of mammals [40]. Most users are unaware that such tools as field guides are intended to help identify what people see in a given place (i.e., to distinguish a bushbuck, Tragelaphus scriptus (Pallas, 1766) from a sitatunga T. spekii Speke, 1863; taxonomy follows predominantly [41]) but say nothing about the number of taxa, their phyletic relationships, and the rank accorded inside the sitatunga and the bushbuck concepts [42]. Diatribes regarding taxonomic subdivisions of even the most well-known African mammals are widespread (e.g., African elephants [43], ungulates [44], felids [45], and canids [46]), including many species that are commonly relocated through the African continent (see Table 1). Wildlife managers are often convinced that wildlife was ubiquitous before humans exerted a strong pressure on it, which also leads to local extirpations.
Taxonomic research, on the contrary, evidences the existence of discrete morphological (and sometimes genetic) discontinuity inside globally perceived “species,” with geographic patterns that often follow well-known biogeographical subdivisions of Africa, and sometimes an abrupt taxa’s limits congruent with biomes and vegetation types changes are observed [75]. Although we may detect areas of apparent intergradations between clearly different “forms” (for the African buffalo, see [76,77]), there are few doubts about the existence of a long and complicated history of adaptation, separation, retraction, and advancement of geographic ranges of different taxa following climate changes, which must be summarized by a not so simple taxonomy, as is often requested by some stakeholders and researchers [78,79]. Finally, complex taxonomies are also difficult to translate into national and international legislations; yet, this is vital for effective conservation policy to maintain current levels of biodiversity [80].

3. Translocations and Conservation

Table 1 shows some selected cases of translocation of African mammals that caused or may cause serious genetic-conservation, as well as sanitary problems or other undesirable conservation consequences.
Regrettably, in some cases, as with the black rhinoceros Diceros bicornis, the issue remains a highly academic one as extirpation of most rhinoceros populations preceded modern evidence-based conservation management [66,81]. In the latter decades, the increasing attention to tourism by several African countries led to growing attention to protected areas and, if needed, to the reintroductions of species that became historically extinct in these regions [25,82,83]. Hybridization between distinct taxa is a common result of translocations in South Africa, where several genera such as Connochaetes and Aepyceros are involved [21,84]. The same threat is now spreading elsewhere, a case in point being the managed populations of two Taurotragus species in the same area in Senegal; one being the critically endangered Taurotragus derbianus derbianus whose genetic integrity is potentially threatened by the imported T. oryx oryx—fortunately in this case, no hybrids have yet been detected based on microsatellite markers [73]. In the same private “reserves” in Senegal, several species, including Ceratotherium simum simum, Kobus ellipsiprymnus ellipsiprymnus, Tragelaphus strepsiceros strepsiceros, Aepyceros melampus melampus, Oryx gazella gazella, and Giraffa camelopardalis giraffa have been introduced from South Africa [71,72] (see also Table 1), which is a fact that raise some concern, especially in cases of animal escapes. Ironically, translocations may also be responsible for the extinction of pure genetic lineages via hybridization, thereby negatively impacting endangered, indigenous, and rare species. Owing to a general neglect of taxonomy—following the so-called “taxonomic inertia” period described by [40]—it is highly probable that the hybridization problem is greatly undervalued in conservation circles. However, a more subtle danger is the general distortions of the genetic landscape, especially of wild ungulates, as is often reported in North America and Europe [85] but also from Africa [86], which may preclude further research, and which may create management problems that cannot be properly anticipated.
Poor taxonomic knowledge coupled with translocations may also lead to conservation initiatives that have only an aesthetic value and, more seriously, may divert attention from real priorities. On the other hand, we know that funds and conservation interest is not homogeneous in Africa [87], and, therefore, we may accept that the extant of biodiversity we risk to lose is greater than we think, especially outside East and South Africa.

4. Giraffes as a Case Study

The last decade saw an unparalleled interest in giraffe taxonomy, starting with studies by [82,88,89]. We associate this unprecedented progress with the use of various data types, increased sampling of studies and availability of the data, various expert knowledge, and the tuning of arguments via numerous discussions [44,90,91,92,93,94]. Currently, we have increasing knowledge about giraffe morphological differentiation [44,77,88], phylogenetic structure and timing of differentiations of current or extinct populations [89,95,96,97,98,99,100], gene flow [44,100,101], unique genomic signatures [102], and potential ecological factors responsible for the restricted gene flow [75]. Giraffes are thus becoming the model case study for the testing of species delimitation in mammals, similarly as cetaceans became the extraordinary model for understanding the various aspect of their and general vertebrate evolution in unprecedented detail [103,104].
Departing from the classical monotypic taxonomic account dominating the 20th century, Colin Groves and Peter Grubb considered that an eight species arrangement better interpreted available morphological data [44]; for the basic overview of giraffe taxonomy across the 20th and 21st centuries, see Table 2. This latter work, adopting a phylogenetic species concept, was not always accepted by several researchers, yet, in the Giraffa case, subsequent genetic works were unanimous in always accepting more than one species. Four species were accepted by [99] and more recently by [100]—Giraffa camelopardalis (Linnaeus, 1758); G. reticulata De Winton, 1899; G. giraffa (Boddaert, 1785); and G. tippelskirchi Matschie, 1898—while [97] recognized three species: Giraffa camelopardalis, G. giraffa, and G. tippelskirchi. Although a step forward, several gaps in available data and interpretation were evident, as too often these reviews were based on previous arrangements without a critical examination of geographic and taxonomic gaps. [94] rightly stressed the taxonomic history of the taxon G. g. rothschildi Lydekker, 1903 to evidence the lack of consensus and possible negative setbacks for conservation. A preponderance of scientists have followed the conservative IUCN protocol and continue to refer to the different kinds of giraffes using the one species/nine subspecies account [94], although this view can be quite dangerous, or at least controversial, for conservation. For example, the materials related to [99] specifically posters a “Giraffe conservation guide” and provided a clear and alarming conservation message to the public about the patchy distribution with declining trends in many northern populations. Recently, an important contribution came from [98], which also included genetic data from key museum specimens belonging to now extinct populations. Among the most important findings include the description of a new—historically extinct—taxon from Senegal, G. camelopardalis senegalensis Petzold, Magnant, and Hassanin, 2020, which went extinct around 1970; the revalidation of G. g. wardi Lydekker, 1904; and a stricter geographical delimitation of the nominal G. c. camelopardalis, which make this taxon, the Nubian giraffe, another example of a taxon “allowed to slip into extinction unnoticed” [40]. The better definition of conservation units or ECUs is a greater priority today than fixing the number of species, considering that taxonomy is particularly relevant when translocations are considered a key conservation component [105]. Petzold and collaborators [98] have contributed to better delineate conservation units, especially in the northern continental sector that has traditionally received scarcer conservation attention and where biogeographical barriers, such as the Nile Basin, have been scarcely considered in mammal taxonomic studies [106,107]. The western giraffe, G. c. peralta Thomas, 1898 emerges unanimously as a distinct taxonomic unit and hence as a conservation priority. Translocations to create more populations should be promoted, whereas the creation of breeding populations of extralimital taxa, such as the G. c. giraffa imported from South Africa to Senegal in the Bandia Private Reserve [108], pose more than a question regarding their conservation relevance and possible interference with long-term G. c. peralta conservation. Regrettably, it can be hypothesized that tourism development in West Africa may lead to further attempts to reconstruct a “true African wildlife experience” for naïve tourists through the importation of stocks from South African wildlife reserves, de facto enlarging a problem of genetic pollution that is already widespread in South Africa [19]. In another critical region, the Nile Basin, translocations have a critical role to play in re-establishing species to their former range [109]. It is vital however that each remaining nominal taxon/population is managed separately, avoiding premature lumping based on scattered evidence and a lack of awareness of biogeographical and ecological barriers. For example, [110] had already proposed a close phyletic relationship between the subspecies rothschildi, cottoni Lydekker, 1904 and antiquorum (Swainson, 1835), and although genetic data seems to confirm this, the adaptive significance of some morphological features of rothschildi (dark coloration and two additional posterior horns in the males) are still unknown. Considering the fact that some authors specify in detail some morphological differences between camelopardalis and rothschildi, and other taxa [110,111,112], it would be worthwhile to inspect their validity using modern statistical methods (e.g., multivariate methods, discriminant analysis). Still more controversial seems the inclusion of the taxon congoensis Lydekker, 1903 as a synonym of antiquorum, considering that his original descriptor considered it an intermediate between northern and southern giraffes because of completely spotted limbs (a southern characteristic) and a well-developed frontal horn (a northern characteristic) [110,113]. The putative taxon congoensis is actually restricted to the Garamba National Park on the left side of the White Nile, whereas putative cottoni is found on the other side of the White Nile, which is apparently a non-trivial zoogeographical barrier.
In their recent genetic work [98], Petzold and collaborators evidenced how the nominal G. c. camelopardalis must now be considered extinct. This is further evidence of the relevance of sampling museum specimens from type locality [114] even if we have to correct these authors by circumscribing the type locality of camelopardalis to the Setit River (in present day Sudan and Eritrea), a well-known area for giraffe zoo collectors in the 19th and first half of the 20th centuries [115,116]. According to [98], the residual giraffe populations from the Gambella and Omo NPs in Southwest Ethiopia belong to rothschildi, but, as the only survivors in a complex biogeographical region east of the Nile that is still partially unexplored biologically [117], we recommend affording great priority to their conservation.
In summary, translocations of giraffes have a long tradition [105]. Some represent a helpful conservation management tool, as we mentioned above, but some others are questionable from a conservation perspective, at least until we have a clear understanding of separable conservation units in some key regions that have so far received little attention.
Table 2. Giraffe taxa recognized in the 20th and 21st centuries across several basic sources, which seemed to assess giraffes independently and/or using different data.
Table 2. Giraffe taxa recognized in the 20th and 21st centuries across several basic sources, which seemed to assess giraffes independently and/or using different data.
aethiopica camelopardalis
antiquorumxxx xxx
australiscapensis giraffagiraffa
biturigum camelopardaliscamelopardalis
camelopardalis xxxxxxx
capensisxx giraffagiraffa
congoensisxx camelopardaliscamelopardalis x
cottonixx rothschildirothschildi
giraffa xxxxxx
hagenbecki reticulata reticulatareticulata
infumata x angolensisangolensis
maculata capensis giraffagiraffa
nigrescensxreticulata reticulatareticulata
reticulatax *xxxxxxx
schillingsitippelskirchitippelskirchi tippelskirchitippelskirchi
senaariensis camelopardalis antiquorumantiquorum
thornicroftixx xxxxx
typicaxcamelopardalis camelopardalis
wardixx giraffa
valid taxa (sp.; ssp.)2; 13 2; 131; 81; 81; 91; 91; 91; 8
aethiopica camelopardalis
africana camelopardalis
angolensisgiraffax or giraffa?giraffax *xx
antiquorumxx or camelopardalis?camelopardalisnot sampledxx
australis giraffa
biturigum camelopardalis
camelopardalisxxxnot sampledxx
capensisgiraffa giraffa giraffa
congoensisantiquorum camelopardalis
cottoni rothschildi camelopardalis
giraffaxxxx *xx
hagenbecki reticulata
infumatagiraffa giraffa angolensis
maculata giraffa
nigrescens reticulata
peraltaantiquorumx or camelopardalis?camelopardalisx *xx
renatae camelopardalis
reticulataxxxx *xx
rothschildicamelopardalisxxx *xcamelopardalis
schillingsi tippelskirchi
senaariensis camelopardalis
thornicroftixxxnot sampledxx
tippelskirchixxxx *xx
typica camelopardalis
wardigiraffa giraffa giraffa
valid taxa (sp.; ssp.)1; 61; min. 61; 66; min. 6, up to 111; 91; 8
aethiopica G. c. camelopardalis
angolensisxx *x (as subspecies of G. giraffa)G. g. giraffa
antiquorumxx *x (as subsp. of G. camelopardalis)x (as subsp. of G. camelopardalis)
australis G. g. giraffa
biturigum G. c. camelopardalis
camelopardalisxx *x *x *
capensis giraffa G. g. giraffa
congoensis antiquorum? G. c. antiquorum
cottoni camelopardalis G. c. antiquorum
giraffaxx *x *x *
hagenbecki G. c. reticulata
infumata giraffa G. g. wardi
maculata G. g. giraffa
nigrescens G. c. reticulata
peraltaxx *x (as subsp. of G. camelopardalis)x (as subsp. of G. camelopardalis)
reticulataxx *x *x (as subsp. of G. camelopardalis)
rothschildixcamelopardalisG. c. camelopardalisx (as subsp. of G. camelopardalis)
senegalensis x (as subsp. of G. camelopardalis)
schillingsi G. t. tippelskirchi
senaariensis G. c. antiquorum
thornicroftixx *x (as subsp. of G. tippelskirchi)x (as subsp. of G. tippelskirchi)
tippelskirchixx *x *x *
typica G. c. camelopardalis
wardi giraffa x (as subsp. of G. giraffa)
valid taxa (sp.; ssp.)1; 98; not recognized 4; 73; 10
Taxa recognized as valid are labelled by “x.” Taxa recognized as species are labelled by “*.” Synonyms of specific taxa are specified when they were noted by particular authors. Abbreviations: BSI—body size, C—coloration, G—genomic data, mtDNA—mitochondrial DNA, nDNA—nuclear DNA, sp.—species, ssp.—subspecies, SSH—skull shapes, SSI—skull size, STRs—microsatellites, U—unspecified, U(M)—unspecified (presumably morphology), and U(M+G)—unspecified (presumably morphological and genetic data).

5. Discussion

As the current discussion about giraffe taxonomy is too focused on the number of species, we argue that the plurality of taxonomic and conservation approaches might be beneficial in order to unify conservation priorities, contra e.g., [127]; see also the comments on taxonomic instability in [128]. Taxonomists often use some particular species concept (for review see [129]), and population/evolutionary geneticists describe interesting results but often without presenting formal taxonomic actions [130,131] and/or alternatively using some standard conservation units—usually management unit (MU), evolutionary significant unit (ESU), cf. [132], or elemental conservation unit (ECU), cf. [133]. The reasons for diverse approaches are various, from the objective (e.g., lost or inaccessible type material, unsuccessful isolation of DNA from type series, difficulty to have samples from same regions) to the subjective (personal adherence to a particular approach based on various reasons or scarce knowledge of taxonomic practice); moreover, a consensus about the “species” label is often hardly obtainable, albeit units deserving conservation attention could be identical or very similar. This is also the case with giraffes (Table 2), because these populations—angolensis, antiquorum, camelopardalis, giraffa, peralta, reticulata, rothschildi, thornicrofti, and tippelskirchi—have been recognized as valid taxa in at least 13 of 18 reviewed sources since 1904 (Table 2). The concordance about the uniqueness of these populations and the latest assessments of giraffes, e.g., [97,98,99,100,101] are quite considerable. Therefore, we recommend using several criteria to define basic taxonomic and conservation units contemporarily (cf. [66] in the case of Diceros bicornis using MU, ESU, and higher level ESUs) in conservation management plans and programs in order to find the majority consensus on which “lineages” deserve conservation attention. This approach would meet integrative taxonomy standards [134]. Robuchon and collaborators [135] offered an excellent framework for evaluation of the impact of species splitting on species priority-setting that is more than recommended for accommodating new taxonomic knowledge in conservation strategies. Genomic data holds enormous potential to resolve species delimitation and recognize demographic history and adaptive potential in detail [136]; its usage should be recommended for future studies. The common practice (e.g., making sequences and other data available on GenBank and/or dryad data platforms as much as possible) should be continued, because it enables independent data testing and uses different approaches. Additionally, we highly recommend associating photographs and/or basic descriptions of the phenotype/size of the DNA voucher specimens, as proposed by [137].
As some current genomic assessments have recognized significant differentiation of populations, specifically in plains zebra Equus quagga Boddaert, 1785 [138] or tiger Panthera tigris (Linnaeus, 1758) [139], which have been recognized as undifferentiated based on small number of loci (zebras [130]) and some phenotype and ecological features (tigers [140]), we argue for a precautionary principle in conservation management [128,135,141].
Africa is a diverse continent with a growing ecotourism industry. Charismatic megavertebrates are particularly searched by tourists and “big game” hunters, but small-scale tourism may furnish valuable income to communities protecting valuable or appealing populations such as the Niger giraffe G. (camelopardis) peralta, the Bor maneless zebra E. quagga borensis Lönnberg, 1921, or the mountain nyala Tragelaphus buxtoni (Lydekker, 1910); income may also be generated by active promotion of a more sensitive mammal watching activity [142].
A further threat is represented by conservative taxonomies that are proposed by conservation groups without a real review of new and old data, as has been recently the case with the IUCN/SSC Cat Specialist Group [45]. Acceptance of a two-subspecies arrangement largely based on some genetic data seems to overly neglect the great phenotypic diversity still found, for instance, in lion Panthera leo (Linnaeus, 1758) [143,144] and ignores some studies that highlight genetic divergence promoted by ecological discontinuity [145,146] that seems to support a new paradigm to explain microevolutionary divergence in widespread, large-sized carnivores [147]. Therefore, without forgetting or underestimating the composite problems of lion conservation, and of coexistence/interaction with humans involving this highly charismatic species [148,149], taxonomic issues deserve to be included in the future conservation strategies of this charismatic species. Paradoxically, tourists (but also trophy hunters) may be greatly interested in phenotypic lion diversity, and works such as [143] may provide the input for further travels in different regions of Africa and the promotion of new conservation/tourism projects for little-known, overlooked populations such as those of Southwestern Ethiopia. Regrettably, genetic considerations alone have suggested merging quite distinct captive populations of two gazelle subspecies Nanger dama dama (Pallas, 1766) and N. d. mhorr (Bennett, 1833) [150]. Apart from pure scientific considerations [151], this line of action should preclude the possibility for Western Sahara communities to have a part in managing for their own welfare a unique ungulate taxon, which is an important incentive for local ecotourism; this is a view opposed to that recently presented in [152].

6. Conclusions

It should be emphasized that the taxonomically sensitive translocation policies [153] would be important for the preservation of current diversity but also for the ecological restoration of some regions within rewilding, which could be essential for the preservation of some unique (often refugee) species [154,155]. Considering the restricted distribution of some giraffe taxa [98,156] and its important role in communities as flower predators [157], giraffe translocations [105] have a great potential that should be utilized for future generations.
Furthermore, the creation in Africa of fenced private or state wildlife reserves filled with exotic stocks not only have a disputable educative effect on tourists and also on local wildlife managers, actually creating or perpetuating a homogenized idea of “African wildlife,” but even create the possibility of concrete dangers. In fact, some species can escape by crossing the fences, which has already happened, and the potential exists to create free allochthonous extra-range nuclei and also to undermine the genetic integrity of native subspecies possibly present in the areas surrounding the reserves.
It is time that conservation biologists recognize the immense threat that taxonomical oversight coupled with the great economic significance of tourism poses to the diversity and integrity of the “genetic landscape” (that is, the evolutionary landscape) of large mammals in Africa. If we wish to preserve such heritage, keeping it as unmodified as possible to future generations of African people and investigators, we need to act now with an urgent change of attitude toward these issues.

Author Contributions

S.G. coinceived the original idea and wrote a first draft; F.M.A. and J.R. developed the two tables and integrate the original draft. All authors have read and agreed to the published version of the manuscript.


This research received no external founding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.


We wish to thank Ken Kawata for improving the English text. Thomas Kirkwood and two anonymous reviewers provided useful suggestions that greatly improved the final manuscript.

Conflicts of Interest

The authors declare no conflict of interest.


  1. Faith, J.T. Late Pleistocene and Holocene mammal extinctions on continental Africa. Earth Sci. Rev. 2014, 128, 105–121. [Google Scholar] [CrossRef]
  2. Faith, J.T.; Rowan, J.; Du, A.; Paul, L.; Koch, P.L. Plio-Pleistocene decline of African megaherbivores: No evidence for ancient hominin impacts. Science 2018, 362, 938–941. [Google Scholar] [CrossRef] [Green Version]
  3. Blixen, K.C. Out of Africa; Putnam: London, UK, 1937. [Google Scholar]
  4. Akama, J.S. Western environmental values and nature-based tourism in Kenya. Tour. Manag. 1996, 17, 567–574. [Google Scholar] [CrossRef]
  5. Schwarzer, J.; Swartz, E.R.; Vreven, E.; Snoeks, J.; Cotterill, F.P.D.; Misof, B.; Schliewen, U.K. Repeated trans-watershed hybridization among haplochromine cichlids (Cichlidae) was triggered by Neogene landscape evolution. Proc. R. Soc. B 2012, 279, 4389–4398. [Google Scholar] [CrossRef] [PubMed]
  6. Haas, F.; Schwarz, E. Zur Entwicklung der afrikanischen Stromsysteme. Geol. Rundsch. 1913, 4, 603–607. [Google Scholar] [CrossRef]
  7. Heller, E. The Geographical Barriers to the Distribution of Big Game Animals in Africa. Geogr. Rev. 1918, 6, 297–319. [Google Scholar] [CrossRef]
  8. Schwarz, E. Huftiere aus West-und Zentralafrika. Ergeb. Dtsch. Zent. Afr. Exped. 1920, 1, 831–1044. [Google Scholar]
  9. Lönnberg, E. The development and distribution of the African fauna in connection with and depending upon climate changes. Ark. Zool. 1929, 21, 1–33. [Google Scholar]
  10. Kingdon, J. East African Mammals: An Atlas of Evolution in Africa; Academic Press: London, UK, 1971; Volume 1. [Google Scholar]
  11. Kingdon, J. Island Africa; Collins: London, UK, 1990. [Google Scholar]
  12. Maglio, V.J.; Cooke, H.B.S. Evolution of African Mammals; Harvard University Press: Cambridge, MA, USA, 1978. [Google Scholar]
  13. Vrba, E.S.; Denton, G.H.; Partridge, T.C.; Burckle, L.H. Paleoclimate and Evolution, with Emphasis on Human Origins; Yale University Press: New Haven, CT, USA; London, UK, 1996. [Google Scholar]
  14. Bromage, T.; Schrenk, F. African Biogeography, Climate Change, and Early Hominid Evolution; Oxford University Press: Oxford, UK, 1999. [Google Scholar]
  15. Schikora, T.F. Climate-Linked Temporal and Spatial Patterns in the Evolution of African Bovidae. Ph.D. Thesis, Johann Wolfgang Goethe-Universität, Frankfurt, Germany, 2000. [Google Scholar]
  16. Cotterill, F.P.D. The Evolutionary History and Taxonomy of the Kobus Leche Species Complex of South-Central Africa in the Context of Palaeo-Drainage Dynamics. Ph.D. Thesis, University of Stellenbosch, Stellenbosch, South Africa, 2006. [Google Scholar]
  17. Couvreur, T.L.; Dauby, G.; Blach-Overgaard, A.; Deblauwe, V.; Dessein, S.; Droissart, V.; Hardy, O.J.; Harris, D.J.; Janssens, S.B.; Ley, A.C.; et al. Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna. Biol. Rev. 2021, 96, 16–51. [Google Scholar] [CrossRef]
  18. Price, R.A. The Contribution of Wildlife to the Economies of Sub Saharan Africa: K4D Helpdesk Report; Institute of Development Studies: Brighton, UK, 2017. [Google Scholar]
  19. Castley, J.G.; Boshoff, A.F.; Kerley, G.I.H. Compromising South Africa’s natural biodiversity—Inappropriate herbivore introductions. S. Afr. J. Sci. 2001, 97, 344–348. [Google Scholar]
  20. Spear, D.; Chown, S.L. The extent and impacts of ungulate translocations: South Africa in a global context. Biol. Conserv. 2009, 142, 353–363. [Google Scholar] [CrossRef] [Green Version]
  21. Russo, I.R.M.; Hoban, S.; Bloomer, P.; Kotzé, A.; Segelbacher, G.; Rushworth, I.; Birss, C.; Bruford, M.W. ‘Intentional Genetic Manipulation’ as a conservation threat. Conserv. Genet. Resour. 2019, 11, 237–247. [Google Scholar] [CrossRef] [Green Version]
  22. Berger-Tal, O.; Blumstein, D.T.; Swaisgood, R.R. Conservation translocations: A review of common difficulties and promising directions. Anim. Conserv. 2020, 23, 121–131. [Google Scholar] [CrossRef] [Green Version]
  23. Kotzé, A.; Smith, R.M.; Moodley, Y.; Luikart, G.; Birss, C.; Van Wyk, A.M.; Van Wyk, A.M.; Grobler, J.P.; Dalton, D.L. Lessons for conservation management: Monitoring temporal changes in genetic diversity of Cape mountain zebra (Equus zebra zebra). PLoS ONE 2019, 14, e0220331. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  24. Smith, R.M.; Bhoora, R.V.; Kotzé, A.; Grobler, J.P.; Dalton, D.L. Translocation a potential corridor for equine piroplasms in Cape mountain zebra (Equus zebra zebra). Int. J. Parasitol. Parasites Wildl. 2019, 9, 130–133. [Google Scholar] [CrossRef]
  25. Correia, M.; Timéteo, S.; Rodríguez-Echeverría, S.; Mazars-Simon, A.; Heleno, R. Refaunation and the reinstatement of the seed-dispersal function in Gorongosa National Park. Conserv. Biol. 2017, 31, 76–85. [Google Scholar] [CrossRef]
  26. Cromsigt, J.P.G.M.; te Beest, M.; Kerley, G.I.H.; Landman, M.; le Roux, E.; Smith, F.A. Trophic rewilding as a climate change mitigation strategy? Philos. Trans. R. Soc. B 2018, 373, 20170440. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  27. Germano, J.M.; Field, K.J.; Griffiths, R.A.; Clulow, S.; Foster, J.; Harding, G.; Swaisgood, R.R. Mitigation-driven translocations: Are we moving wildlife in the right direction? Front. Ecol. Environ. 2015, 13, 100–105. [Google Scholar] [CrossRef]
  28. Ackermann, R.R.; Brink, J.S.; Vrahimis, S.; de Klerk, B. Hybrid wildebeest (Artiodactyla: Bovidae) provide further evidence for shared signatures of admixture in mammalian crania. S. Afr. J. Sci. 2010, 106, 1–4. [Google Scholar] [CrossRef]
  29. Grobler, J.P.; Rushworth, I.; Brink, J.S.; Bloomer, P.; Kotze, A.; Reilly, B.; Vrahimis, S. Management of hybridization in an endemic species: Decision making in the face of imperfect information in the case of the black wildebeest—Connochaetes gnou. Eur. J. Wildl. Res. 2011, 57, 997–1006. [Google Scholar] [CrossRef] [Green Version]
  30. Benjamin-Fink, N.; Reilly, B.K. Conservation implications of wildlife translocations; The state’s ability to act as conservation units for wildebeest populations in South Africa. Glob. Ecol. Conserv. 2017, 12, 46–58. [Google Scholar] [CrossRef]
  31. Spear, D.; Chown, S.L. Taxonomic homogenization in ungulates: Patterns and mechanisms at local and global scales. J. Biogeogr. 2008, 35, 1962–1975. [Google Scholar] [CrossRef]
  32. Goss, J.R.; Cumming, G.S. Networks of wildlife translocations in developing countries: An emerging conservation issue. Front. Ecol. Environ. 2013, 11, 243–250. [Google Scholar] [CrossRef]
  33. Maciejewski, K.; Kerley, G.I.H. Understanding tourists’ preference for mammal species in private protected areas: Is there a case for extralimital species for ecotourism? PLoS ONE 2014, 9, e88192. [Google Scholar] [CrossRef]
  34. van Hoven, W. Private game reserves in Southern Africa. In Institutional Arrangements for Conservation, Development and Tourism in Eastern and Southern Africa: A Dynamic Perspective; Springer: Dordrecht, The Netherlands, 2015; pp. 101–118. [Google Scholar] [CrossRef]
  35. Lindsey, P.A.; Roulet, P.A.; Romañach, S.S. Economic and conservation significance of the trophy hunting industry in sub-Saharan Africa. Biol. Conserv. 2007, 134, 455–469. [Google Scholar] [CrossRef]
  36. Muposhi, V.K.; Gandiwa, E.; Makuza, S.M.; Bartels, P. Ecological, physiological, genetic trade-offs and socio-economic implications of trophy hunting as a conservation tool: A narrative review. J. Anim. Plant Sci. 2017, 27, 1–14. [Google Scholar]
  37. Seddon, P.J.; Strauss, W.M.; Innes, J. Animal translocations: What are they and why do we do them? In A Reintroduction Biology: Integrating Science and Management, 1st ed.; Ewen, J.G., Armstrong, D.P., Parker, K.A., Seddon, P.J., Eds.; Wiley-Blackwell: Oxford, UK, 2012; pp. 1–32. [Google Scholar] [CrossRef]
  38. Dorst, J.; Dandelot, P. A Field Guide to the Larger Mammals of Africa; Collins: London, UK, 1970. [Google Scholar]
  39. Kingdon, J. The Kingdon Field Guide to African Mammals; Academic Press: San Diego, CA, USA, 1997. [Google Scholar]
  40. Gippoliti, S.; Cotterill, F.P.D.; Zinner, D.; Groves, C.P. Impacts of taxonomic inertia for the conservation of African ungulate diversity: An overview. Biol. Rev. 2018, 93, 115–130. [Google Scholar] [CrossRef]
  41. Grubb, P. Artiodactyla. In Mammal Species of the World: A Taxonomic and Geographic Reference; Wilson, D.E., Reeder, D.M., Eds.; The Johns Hopkins University Press: Baltimore, MD, USA, 2005; pp. 637–722. [Google Scholar]
  42. Gippoliti, S.; Groves, C.P. Cryptic problematic species and troublesome taxonomists: A tale of the Apennine bear and the Nile white rhinoceros. In Problematic Wildlife II; Angelici, F.M., Rossi, L., Eds.; Springer: Cham, Switzerland, 2020; pp. 509–527. [Google Scholar] [CrossRef]
  43. Grubb, P.; Groves, C.P.; Dudley, J.P.; Shoshani, J. Living African elephants belong to two species: Loxodonta africana (Blumenbach, 1797) and Loxodonta cyclotis (Matschie, 1900). Elephant 2000, 2, 1–4. [Google Scholar] [CrossRef]
  44. Groves, C.P.; Grubb, P. Ungulate Taxonomy; The Johns Hopkins University Press: Baltimore, MD, USA, 2011. [Google Scholar]
  45. Kitchener, A.C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A.V.; Christiansen, P.; Driscoll, C.; et al. A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN/SSC Cat Specialist Group. Cat News 2017, Special Issue 11, 1–80. [Google Scholar]
  46. Gippoliti, S.; Lupi, L. A note on the wild canids (Carnivora: Canidae) of the Horn of Africa, with the first evidence of a new–forgotten–species for Ethiopia Canis mengesi Noack, 1897. Bonn Zool. Bull. 2020, 69, 111–115. [Google Scholar] [CrossRef]
  47. Walker, J.F. Will Secret Wildlife Imports Doom Ultra-Rare Giant Sable? Available online: (accessed on 22 May 2021).
  48. van Wyk, A.M.; Dalton, D.L.; Hoban, S.; Bruford, M.W.; Russo, I.M.; Birss, C.; Grobler, P.; van Vuuren, B.J.; Kotzé, A. Quantitative evaluation of hybridization and the impact on biodiversity conservation. Ecol. Evol. 2017, 7, 320–330. [Google Scholar] [CrossRef] [Green Version]
  49. Moodley, Y.; Harley, E.H. Population structuring in mountain zebras (Equus zebra): The molecular consequences of divergent demographic histories. Conserv. Genet. 2005, 6, 953–968. [Google Scholar] [CrossRef]
  50. Cordingley, J.E.; Sundaresan, S.R.; Fischhoff, I.R.; Shapiro, B.; Ruskey, J.; Rubenstein, D.I. Is the endangered Grevy’s zebra threatened by hybridization? Anim. Conserv. 2009, 12, 505–513. [Google Scholar] [CrossRef]
  51. Schieltz, J.M.; Rubenstein, D.I. Caught between two worlds: Genes and environment influence behaviour of plains × Grevy’s zebra hybrids in central Kenya. Anim. Behav. 2015, 106, 17–26. [Google Scholar] [CrossRef] [Green Version]
  52. Green, W.C.H.; Rothstein, A. Translocation, hybridisation and the endangered black-faced impala. Conserv. Biol. 1998, 12, 475–480. [Google Scholar] [CrossRef]
  53. Measey, J.; Hui, C.; Somers, M.J. Terrestrial Vertebrate Invasions in South Africa. In Biological Invasions in South Africa; van Wilgen, B., Measey, J., Richardson, D., Wilson, J., Zengeya, T., Eds.; Springer: Cham, Switzerland, 2020; pp. 115–151. [Google Scholar] [CrossRef] [Green Version]
  54. Taylor, A.; Avenant, N.; Schulze, E.; Viljoen, P.; Child, M.F. A conservation assessment of Redunca fulvorufula fulvorufula. In The Red List of Mammals of South Africa, Swaziland and Lesotho; Child, M.F., Roxburgh, L., Do Linh San, E., Raimondo, D., Davies-Mostert, H.T., Eds.; South African National Biodiversity Institute and Endangered Wildlife Trust: Pretoria and Gauteng, South Africa, 2016; pp. 1–7. [Google Scholar]
  55. Furstenburg, D. Nyala Tragelaphus angasii. In The New Game Rancher; Oberem, P., Oberem, P.T., Eds.; Briza Publications: Queenswood, South Africa, 2016; pp. 1–14. [Google Scholar]
  56. Matthee, C.A.; Robinson, T.J. Mitochondrial DNA population structure of roan and sable antelope: Implications for the translocation and conservation of the species. Mol. Ecol. 1999, 8, 227–238. [Google Scholar] [CrossRef]
  57. Furstenburg, D. Sable Antelope Hippotragus niger. In The New Game Rancher; Oberem, P., Oberem, P., Eds.; Briza Publications: Queenswood, South Africa, 2016; pp. 1–13. [Google Scholar]
  58. Alpers, D.L.; Van Vuuren, B.J.; Arctander, P.; Robinson, T.J. Population genetics of the roan antelope (Hippotragus equinus) with suggestions for conservation. Mol. Ecol. 2004, 13, 1771–1784. [Google Scholar] [CrossRef] [PubMed]
  59. Barrie, A. Translocation of Roan Antelope in South Africa and the Effect This Has Had on the Genetic Diversity of the Species. Mini Dissertation, University of Johannesburg, Johannesburg, South Africa, 2015. Available online: (accessed on 22 May 2021).
  60. Furstenburg, D. Springbok Antidorcas marsupialis. In The New Game Rancher; Oberem, P., Oberem, P.T., Eds.; Briza Publications: Queenswood, South Africa, 2016; pp. 1–18. [Google Scholar]
  61. Starin, E.D. Notes on sitatunga in The Gambia. Afr. J. Ecol. 2000, 38, 339–342. [Google Scholar] [CrossRef]
  62. Carter, N. Arm’d Rhinoceros; Andre Deutsch: London, UK, 1965. [Google Scholar]
  63. Penzhorn, B.L. A summary of the re-introduction of ungulates into South African National Parks (to December 1970). Koedoe 1971, 14, 145–159. [Google Scholar] [CrossRef]
  64. Knight, M.H.; Kerley, G.I.H. Black rhino translocations within Africa. Afr. Insight 2009, 39, 70–83. [Google Scholar] [CrossRef]
  65. Fyumagwa, R.D.; Nyahongo, J.W. Black rhino conservation in Tanzania: Translocation efforts and further challenges. Pachyderm 2010, 47, 59–65. [Google Scholar]
  66. Moodley, Y.; Russo, I.-R.M.; Dalton, D.L.; Kotzé, A.; Muya, S.; Haubensak, P.; Bálint, B.; Munimanda, G.K.; Deimel, C.; Setzer, A.; et al. Extinctions, genetic erosion and conservation options for the black rhinoceros (Diceros bicornis). Sci. Rep. 2017, 7, 41417. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  67. Sheil, D.; Kirkby, A.E. Observations on Southern white rhinoceros Ceratotherium simum simum translocated to Uganda. Trop. Conserv. Sci. 2018, 11, 1–7. [Google Scholar] [CrossRef] [Green Version]
  68. Emslie, R.; Brooks, M. African Rhino: Status Survey and Conservation Action Plan; IUCN/SSC African Rhino Specialist Group: Gland, Switzerland; Cambridge, UK, 1999. [Google Scholar]
  69. Grimwood, I.R. Airlift for Hunter’s antelope: Rescue operation in Kenya. Oryx 1964, 7, 164–167. [Google Scholar] [CrossRef]
  70. Hofmann, R.R. Hirola: Translocation to Tsavo NP and new scientific information. Gnusletter 1996, 15, 2–5. [Google Scholar]
  71. Vincke, X.; Hornick, J.-L.; Njikam, N.I.; Leroy, P. Gestion de la faune sauvage au Sénégal: Comparaison du Parc National du Niokolo Koba et de la Réserve privée de Bandia. Ann. Med. Vet. 2005, 149, 232–237. [Google Scholar]
  72. Vermeulen, C. La réserve faunique de Bandia (Sénégal): Modèle ou contre-modèle pour l’Afrique de l’Ouest ? Parcs et Réserves 2010, 65, 23–27. [Google Scholar]
  73. Kubátová, A.; Štochlová, K.; Brandlová, K.; Jůnková Vymyslická, P.; Bolfíková, B.C. Comparison of divergent breeding management strategies in two species of semi-captive eland in Senegal. Sci. Rep. 2020, 10, 8841. [Google Scholar] [CrossRef]
  74. Hall, S. The king of the jungle is back. Rwanda welcomes lions at Akagera National Park after 21 years’ absence. Eye Mag. 2015, 32–33. [Google Scholar]
  75. Thomassen, H.A.; Freedman, A.H.; Brown, D.M.; Buermann, W.; Jacobs, D.K. Regional differences in seasonal timing of rainfall discriminate between genetically distinct East African giraffe taxa. PLoS ONE 2013, 8, e77191. [Google Scholar] [CrossRef] [Green Version]
  76. Grubb, P. Variation and incipient speciation in the African buffalo. Z. Säugetierkd. 1972, 37, 121–144. [Google Scholar]
  77. Grubb, P. Morphoclinal evolution in ungulates. In Antelopes, Deer, and Relatives; Vrba, E.S., Schaller, G.B., Eds.; Yale University Press: New Haven, CT, USA; London, UK, 2000; pp. 156–170. [Google Scholar]
  78. Taylor, P.J.; Denys, C.; Cotterill, F.P.D. Taxonomic anarchy or an inconvenient truth for conservation? Accelerated species discovery reveals evolutionary patterns and heightened extinction threat in Afro-Malagasy small mammals. Mammalia 2019, 38, 313–329. [Google Scholar] [CrossRef] [Green Version]
  79. Gippoliti, S. Species delimitation in mammals: A comment on Zachos (2018). Mamm. Biol. 2019, 94, 127–131. [Google Scholar] [CrossRef]
  80. Gippoliti, S.; Capula, M.; Ficetola, G.F.; Salvi, D.; Andreone, F. Threatened by legislative conservationism? The case of the critically endangered Aeolian lizard. Front. Ecol. Evol. 2017, 5, 130. [Google Scholar] [CrossRef] [Green Version]
  81. Rookmaaker, K. The black rhino needs a taxonomic revision for sound conservation. Int. Zoo News 2005, 52, 280–282. [Google Scholar]
  82. Brown, D.M.; Brenneman, R.A.; Koepfli, K.-P.; Pollinger, J.P.; Milá, B.; Georgiadis, N.J.; Louis, E.E., Jr.; Grether, G.F.; Jacobs, D.K.; Wayne, R.K. Extensive population genetic structure in the giraffe. BMC Biol. 2007, 5, 57. [Google Scholar] [CrossRef] [Green Version]
  83. Huntley, B.J.; Russo, V.; Lages, F.; Ferrand, N. Biodiversity of Angola Science & Conservation: A Modern Synthesis; Springer International Publishing: Cham, Switzerland, 2019. [Google Scholar] [CrossRef] [Green Version]
  84. Miller, S.M.; Moeller, C.-H.; Harper, C.K.; Bloomer, P. Anthropogenic movement results in hybridisation in impala in southern Africa. Conserv. Genet. 2020, 21, 653–663. [Google Scholar] [CrossRef]
  85. de Jong, J.F.; van Hooft, P.; Megens, H.-J.; Crooijmans, R.P.M.A.; de Groot, G.A.; Pemberton, J.M.; Huisman, J.; Bartoš, L.; Iacolina, L.; van Wieren, S.E.; et al. Fragmentation and Translocation Distort the Genetic Landscape of Ungulates: Red Deer in the Netherlands. Front. Ecol. Evol. 2020, 8, 535715. [Google Scholar] [CrossRef]
  86. Heller, R.; Okello, J.B.A.; Siegismund, H. Can small wildlife conservancies maintain genetically stable populations of large mammals? Evidence for increased genetic drift in geographically restricted populations of Cape buffalo in East Africa. Mol. Ecol. 2010, 19, 1324–1334. [Google Scholar] [CrossRef] [PubMed]
  87. Brockington, D.; Scholfield, K. Expenditure by conservation nongovernmental organizations in sub-Saharan Africa. Conserv. Lett. 2010, 3, 106–113. [Google Scholar] [CrossRef]
  88. Seymour, R. Patterns of Subspecies Diversity in the Giraffe, Giraffa camelopardalis (L. 1758): Comparison of Systematic Methods and Their Implications for Conservation Policy. Ph.D. Thesis, University of Kent, Canterbury, UK, 2001. [Google Scholar]
  89. Hassanin, A.; Ropiquet, A.; Gourmand, A.-L.; Chardonnet, B.; Rigoulet, J. Mitochondrial DNA variability in Giraffa camelopardalis: Consequences for taxonomy, phylogeography and conservation of giraffes in West and central Africa. C. R. Biol. 2007, 330, 265–274. [Google Scholar] [CrossRef] [PubMed]
  90. Mitchell, G.; Skinner, J.D. On the origin, evolution and phylogeny of giraffes Giraffa camelopardalis. Trans. R. Soc. S. Afr. 2003, 58, 51–73. [Google Scholar] [CrossRef]
  91. Groves, C. Giraffe taxonomy: Where are we now? Giraffid 2015, 9, 8–9. [Google Scholar]
  92. Bercovitch, F.B.; Berry, P.S.M.; Dagg, A.; Deacon, F.; Doherty, J.B.; Lee, D.E.; Mineur, F.; Muller, Z.; Ogden, R.; Seymour, R.; et al. How many species of giraffe are there? Curr. Biol. 2017, 27, R123–R138. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  93. Fennessy, J.; Winter, S.; Reuss, F.; Kumar, V.; Nilsson, M.A.; Vamberger, M.; Fritz, U.; Janke, A. Response to “How many species of giraffe are there?”. Curr. Biol. 2017, 27, R123–R138. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  94. Bercovitch, F.B. Giraffe taxonomy, geographic distribution and conservation. Afr. J. Ecol. 2020, 58, 150–158. [Google Scholar] [CrossRef]
  95. Bock, F.; Fennessy, J.; Bidon, T.; Tutching, A.; Marais, A.; Deacon, F.; Janke, A. Mitochondrial sequences reveal a clear separation between Angolan and South African giraffe along a cryptic rift valley. BMC Evol. Biol. 2014, 14, 219. [Google Scholar] [CrossRef] [Green Version]
  96. Stanton, D.W.G.; Hart, J.; Galbusera, P.; Helsen, P.; Shephard, J.; Kümpel, N.F.; Wang, J.; Ewen, J.G.; Bruford, M.W. Distinct and diverse: Range-wide phylogeography reveals ancient lineages and high genetic variation in the endangered Okapi (Okapia johnstoni). PLoS ONE 2014, 9, e101081. [Google Scholar] [CrossRef]
  97. Petzold, A.; Hassanin, A. A comparative approach for species delimitation based on multiple methods of multi-locus DNA sequence analysis: A case study of the genus Giraffa (Mammalia, Cetartiodactyla). PLoS ONE 2020, 15, e0217956. [Google Scholar] [CrossRef]
  98. Petzold, A.; Magnant, A.-S.; Edderai, D.; Chardonnet, B.; Rigoulet, J.; Saint-Jalme, M.; Hassanin, A. First insights into past biodiversity of giraffes based on mitochondrial sequences from museum specimens. Eur. J. Taxon. 2020, 703, 1–33. [Google Scholar] [CrossRef]
  99. Fennessy, J.; Bidon, T.; Reuss, F.; Kumar, V.; Elkan, P.; Nilsson, M.A.; Vamberger, M.; Fritz, U.; Janke, A. Multi-locus analyses reveal four giraffe species instead of one. Curr. Biol. 2016, 26, 2543–2549. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  100. Coimbra, R.T.F.; Winter, S.; Kumar, V.; Koepfli, K.-P.; Gooley, R.M.; Dobrynin, P.; Fennessy, J.; Janke, A. Whole-genome analysis of giraffe supports four distinct species. Curr. Biol. 2021, 31, 1–10. [Google Scholar] [CrossRef]
  101. Winter, S.; Fennessy, J.; Janke, A. Limited introgression supports division of giraffe into four species. Ecol. Evol. 2018, 8, 10156–10165. [Google Scholar] [CrossRef]
  102. Agaba, M.; Ishengoma, E.; Miller, W.; McGrath, B.C.; Hudson, C.N.; Bedoya Reina, O.C.; Ratan, A.; Burhans, R.; Chikhi, R.; Medvedev, P.; et al. Giraffe genome sequence reveals clues to its unique morphology and physiology. Nat. Commun. 2016, 7, 11519. [Google Scholar] [CrossRef] [Green Version]
  103. Gatesy, J.; Geisler, J.H.; Chang, J.; Buell, C.; Berta, A.; Meredith, R.W.; Springer, M.S.; McGowen, M.R. A phylogenetic blueprint for a modern whale. Mol. Phylogenet. Evol. 2013, 66, 479–506. [Google Scholar] [CrossRef] [PubMed]
  104. McGowen, M.R.; Gatesy, J.; Wildman, D.E. Molecular evolution tracks macroevolutionary transitions in Cetacea. Trends Ecol. Evol. 2014, 29, 336–346. [Google Scholar] [CrossRef] [PubMed]
  105. Muller, Z.; Lee, D.E.; Scheijen, C.P.J.; Megan, K.L.; Strauss, M.K.L.; Kerryn, D.; Carter, K.D.; Deacon, F. Giraffe translocations: A review and discussion of considerations. Afr. J. Ecol. 2020, 58, 159–171. [Google Scholar] [CrossRef]
  106. Kostin, D.S.; Martynov, A.A.; Komarova, V.A.; Alexandrov, D.Y.; Yihune, M.; Kasso, M.; Bryja, J.; Lavrrenchenko, L.A. Rodents of Choke Mountain and surrounding areas (Ethiopia): The Blue Nile gorge as a strong biogeographic barrier. J. Vertebr. Biol. 2020, 69, 20016. [Google Scholar] [CrossRef]
  107. Angelici, F.M.; Colangelo, P.; Gippoliti, S. Out of Europe: Investigating Hystrix cristata (Rodentia: Hystricidae) skull morphometric geographic variability in Africa. Biogeographia 2021, 36, a001. [Google Scholar] [CrossRef]
  108. Malyjurkova, L.; Hejzlarova, M.; Vymyslicka, P.J.; Brandlova, K. Social Preferences of Translocated Giraffes (Giraffa camelopardalis giraffa) in Senegal: Evidence for Friendship among Females? Agric. Trop. Subtrop. 2014, 47, 5–13. [Google Scholar] [CrossRef] [Green Version]
  109. Brown, M.B.; Bolger, D.T.; Fennessy, J. All the eggs in one basket: A countrywide assessment of current and historical giraffe population distribution in Uganda. Glob. Ecol. Conserv. 2019, 19, e00612. [Google Scholar] [CrossRef]
  110. Lydekker, R. On the Subspecies of Giraffa camelopardalis. Proc. Zool. Soc. Lond. 1904, 74, 202–227. [Google Scholar] [CrossRef]
  111. Krumbiegel, I. Die Giraffe: Unter Besonderer Berücksichtigung der Rassen; Monographien der Wildsäugetiere; Verlag Dr. Paul Schöps: Leipzig, Germany, 1939; Volume VIII, pp. 1–98. [Google Scholar]
  112. MacClintock, D.; Mochi, U. A Natural History of Giraffes; Charles Scribner´s Sons: New York, NY, USA, 1973. [Google Scholar]
  113. Schouteden, H. Note sur la giraffe du Congo. Rev. Zool. Afr. 1913, 2, 134–137. [Google Scholar]
  114. Gippoliti, S. Everything mammal conservation biologists always wanted to know about taxonomy (but were afraid to ask). J. Nat. Conserv. 2020, 54, 125793. [Google Scholar] [CrossRef]
  115. Tedesco Zammarano, V. Fauna e Caccia; Ministero delle Colonie: Rome, Italy, 1930. [Google Scholar]
  116. Gippoliti, S.; Hagos, F.; Angelici, F.M. Eritrean ungulates in Italian museums as benchmark for taxonomy and conservation planning. Hystrix 2018, 29 (XI ATIt Congress Suppl.), 30. [Google Scholar]
  117. Gippoliti, S. On the Taxonomy of Erythrocebus with a re-evaluation of Erythrocebus poliophaeus (Reichenbach, 1862) from the Blue Nile Region of Sudan and Ethiopia. Primate Conserv. 2017, 31, 53–59. [Google Scholar]
  118. Lydekker, R. Two undescribed giraffe. Nature 1911, 87, 484. [Google Scholar] [CrossRef]
  119. Haltenorth, T. Klassifikation der Säugetiere: Artiodactyla I (18). In Handbuch der Zoologie; Walter de Gruyter & Co.: Berlin, Germany, 1963; Volume 8, pp. 1–167. [Google Scholar]
  120. Spinage, C.A. The Book of the Giraffe; Collins: London, UK, 1968. [Google Scholar]
  121. Ansell, W.F.H. Artiodactyla (Excluding the Genus Gazella). In The Mammals of Africa—An Identification Manual for African Mammals; Meester, J.A., Setzer, H.W., Eds.; Smithsonian Institution Press: Washington, DC, USA, 1971; pp. 1–84. [Google Scholar]
  122. Dagg, A.I.  Giraffa camelopardalis . Mamm. Species 1971, 5, 1–8. [Google Scholar] [CrossRef]
  123. East, R. African Antelope Database 1998; IUCN/SSC Antelope Specialist Group IUCN: Gland, Switzerland; Cambridge, UK, 1999. [Google Scholar]
  124. Skinner, J.D.; Mitchell, G. Family Giraffidae (Giraffe and Okapi). In Handbook of the Mammals of the World; Wilson, D.E., Mittermeier, R.A., Eds.; Lynx Edicions: Barcelona, Spain, 2011; Volume 2, pp. 788–802. [Google Scholar]
  125. Ciofolo, C.; Le Pendu, Y. Giraffa camelopardalis Giraffe. In Mammals of Africa; Kingdon, J., Hoffmann, M., Eds.; Bloomsbury Publishing: London, UK, 2013; Volume VI, pp. 98–109. [Google Scholar]
  126. Shorrocks, B.; Bates, W. The Biology of African Savannahs, 2nd ed.; Oxford University Press: Oxford, UK, 2015. [Google Scholar]
  127. Garnett, S.T.; Christidis, L. Taxonomy anarchy hampers conservation. Nature 2017, 546, 25–27. [Google Scholar] [CrossRef]
  128. Robovský, J.; Melichar, L.; Gippoliti, S. Zoos and conservation in the Anthropocene: Opportunities and problems. In Problematic Wildlife II; Angelici, F.M., Rossi, L., Eds.; Springer: Cham, Switzerland, 2020; pp. 451–484. [Google Scholar] [CrossRef]
  129. Groves, C.P.; Cotterill, F.P.D.; Gippoliti, S.; Robovský, J.; Roos, C.; Taylor, P.J.; Zinners, D. Species definitions and conservation: A review and case studies from African mammals. Conserv. Genet. 2017, 18, 1247–1256. [Google Scholar] [CrossRef]
  130. Lorenzen, E.D.; Arctander, P.; Siegismund, H.R. High variation and very low differentiation in wide ranging plains zebra (Equus quagga): Insights from mtDNA and microsatellites. Mol. Ecol. 2008, 17, 2812–2824. [Google Scholar] [CrossRef] [PubMed]
  131. Moritz, C. Defining ‘Evolutionarily Significant Units’ for Conservation. Trends Ecol. Evol. 1994, 9, 373–375. [Google Scholar] [CrossRef]
  132. Minelli, A. The galaxy of the non-Linnaean nomenclature. Hist. Philos. Life Sci. 2019, 41, 31. [Google Scholar] [CrossRef]
  133. Wood, C.C.; Gross, M.R. Elemental Conservation Units: Communicating extinction risk without dictating targets for protection. Conserv. Biol. 2008, 22, 36–47. [Google Scholar] [CrossRef]
  134. Schlick-Steiner, B.C.; Steiner, F.M.; Seifert, B.; Stauffer, C.; Christian, E.; Crozier, R.H. Integrative taxonomy: A multisource approach to exploring biodiversity. Annu. Rev. Entomol. 2010, 55, 421–438. [Google Scholar] [CrossRef] [PubMed]
  135. Robuchon, M.; Faith, D.P.; Julliard, R.; Leroy, B.; Pellens, R.; Robert, A.; Thévenin, C.; Véron, S.; Pavoine, S. Species splitting increases estimates of evolutionary history at risk. Biol. Conserv. 2019, 235, 27–35. [Google Scholar] [CrossRef]
  136. Stanton, D.W.G.; Frandsen, P.; Waples, R.K.; Heller, R.; Russo, I.-R.; Orozco-terWengel, P.A.; Pedersen, C.E.T.; Siegismund, H.R.; Bruford, M.W. More grist for the mill? Species delimitation in the genomic era and its implications for conservation. Conserv. Genet. 2019, 20, 101–113. [Google Scholar] [CrossRef] [Green Version]
  137. Groves, C. The genus Cervus in eastern Eurasia. Eur. J. Wildl. Res. 2006, 52, 14–22. [Google Scholar] [CrossRef]
  138. Pedersen, C.E.T.; Albrechtsen, A.; Etter, P.D.; Johnson, E.A.; Orland, L.; Chikhi, L.; Siegismund, H.R.; Heller, R. A southern African origin and cryptic structure in the highly mobile plains zebra. Nat. Ecol. Evol. 2018, 2, 491–498. [Google Scholar] [CrossRef]
  139. Liu, Y.-C.; Sun, X.; Driscoll, C.; Miquelle, D.G.; Xu, X.; Martelli, P.; Uphyrkina, O.; Smith, J.L.D.; O´Brien, S.J.O.; Luo, S.-J. Genome-Wide Evolutionary Analysis of Natural History and Adaptation in the World’s Tigers. Curr. Biol. 2018, 28, 1–10. [Google Scholar] [CrossRef] [Green Version]
  140. Wilting, A.; Courtiol, A.; Christiansen, P.; Niedballa, J.; Scharf, A.K.; Orlando, L.; Balkenhol, N.; Hofer, H.; Kramer-Schadt, S.; Fickel, J.; et al. Planning tiger recovery: Understanding intraspecific variation for effective conservation. Sci. Adv. 2015, 1, e1400175. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  141. Gippoliti, S.; Cotterill, F.P.D.; Groves, C.P.; Zinner, D. Poor taxonomy and genetic rescue are possible co-agents of silent extinction and biogeographic homogenization among ungulate mammals. Biogeographia 2018, 33, 41–54. [Google Scholar] [CrossRef] [Green Version]
  142. Dinets, V.; Hall, J. Mammalwatching: A new source of support for science and conservation. Int. J. Biodivers. Conserv. 2018, 10, 154–160. [Google Scholar] [CrossRef] [Green Version]
  143. Lupták, P.; Csurma, L. The external variability and taxonomy of recent and extinct subspecies of lion (Panthera leo). Gazella 2009, 36, 33–150. [Google Scholar]
  144. Zamudio, K.R.; Bell, R.C.; Mason, N.A. Phenotypes in phylogeography: Species’ traits, environmental variation, and vertebrate diversification. Proc. Natl. Acad. Sci. USA 2016, 113, 8041–8048. [Google Scholar] [CrossRef] [Green Version]
  145. Moore, A.E.; Cotterill, F.P.D.; Winterbach, C.W.; Winterbach, H.E.K.; Antunes, A.; O’Brien, S.J. Genetic Evidence for Contrasting Wetland and Savannah Habitat Specializations in Different Populations of Lions (Panthera leo). J. Hered. 2016, 107, 101–103. [Google Scholar] [CrossRef] [Green Version]
  146. Dures, S.G.; Carbone, C.; Savolainen, V.; Maude, G.; Gotelli, D. Ecology rather than people restrict gene flow in Okavango-Kalahari lions. Anim. Conserv. 2020, 23, 505–515. [Google Scholar] [CrossRef]
  147. Shafer, A.B.A.; Wolf, J.B.W. Widespread evidence for incipient ecological speciation: A meta-analysis of isolation-by-ecology. Ecol. Lett. 2013, 16, 940–950. [Google Scholar] [CrossRef] [PubMed]
  148. Bauer, H.; Chapron, G.; Nowell, K.; Henschel, P.; Funston, P.; Hunter, L.T.B.; Macdonald, D.W.; Packer, C. Lion (Panthera leo) populations are declining rapidly across Africa, except in intensively managed areas. Proc. Natl. Acad. Sci. USA 2015, 112, 14894–14899. [Google Scholar] [CrossRef] [Green Version]
  149. Trinkel, M.; Angelici, F.M. The Decline in the lion population in Africa and possible mitigation measures. In Problematic Wildlife: A Cross-Disciplinary Approach; Angelici, F.M., Ed.; Springer: Cham, Switzerland, 2016; pp. 45–68. [Google Scholar]
  150. Senn, H.; Banfield, L.; Wacher, T.; Newby, J.; Rabeil, T.; Kaden, J.; Kitchener, A.; Abaigar, T.; Luisa Silva, T.; Maunder, M.; et al. Splitting or lumping? A conservation dilemma exemplified by the critically endangered Dama Gazelle (Nanger dama). PLoS ONE 2014, 9, e98693. [Google Scholar] [CrossRef] [Green Version]
  151. Schreiber, A.; Moreno, E.; Groves, C.; Robovský, J. Systematics and management units of Dama Gazelle Nanger dama. Gnusletter 2018, 35, 8–12. [Google Scholar]
  152. Garnett, S.T.; Thomson, S.A. Are the implications for conservation of a major taxonomic revision of the world’s birds’ simply serendipity? Anim. Conserv. 2020, 23, 355–356. [Google Scholar] [CrossRef]
  153. IUCN/SSC. Guidelines for Reintroductions and Other Conservation Translocations: Version 1.0; IUCN Species Survival Commission: Gland, Switzerland, 2013. [Google Scholar]
  154. Kerley, G.I.H.; Kowalczyk, R.; Cromsigt, J.P.G.M. Conservation implications of the refugee species concept and the European bison: King of the forest or refugee in a marginal habitat? Ecography 2012, 35, 519–529. [Google Scholar] [CrossRef]
  155. Ali, A.H.; Amin, R.; Evans, J.S.; Fischer, M.; Ford, A.T.; Kibara, A.; Goheen, J.R. Evaluating support for rangeland-restoration practices by rural Somalis: An unlikely win-win for local livelihoods and hirola antelope? Anim. Conserv. 2018, 22, 144–156. [Google Scholar] [CrossRef]
  156. O’Connor, D.; Stacy-Dawes, J.; Muneza, A.; Fennessy, J.; Gobush, K.; Chase, M.J.; Brown, M.B.; Bracis, C.; Elkan, P.; Rabeil, T.; et al. Updated geographic range maps for giraffe, Giraffa spp., throughout sub-Saharan Africa, and implications of changing distributions for conservation. Mamm. Rev. 2019, 49, 285–299. [Google Scholar] [CrossRef] [Green Version]
  157. Fleming, P.; Hofmeyr, S.; Nicolson, S.; Du Toit, J. Are giraffes pollinators or flower predators of Acacia nigrescens in Kruger National Park, South Africa? J. Trop. Ecol. 2006, 22, 247–253. [Google Scholar] [CrossRef] [Green Version]
Table 1. Some selected cases of mammalian translocations in Africa, with uncertain outcomes, including goals and consequent problems. Taxonomy follows predominantly [41].
Table 1. Some selected cases of mammalian translocations in Africa, with uncertain outcomes, including goals and consequent problems. Taxonomy follows predominantly [41].
SpeciesOriginal RangeTranslocated toPurposeAftermathReferences
Damaliscus pygargus phillipsi Harper, 1939 Northeast South AfricaSouthwest South Africa, Botswana, Mozambique, Namibia, Swaziland, Zimbabwe, AngolaHunting restocking, conservation, eco-tourismGenetic integrity of D. p. pygargus (Pallas, 1767); invasive in Angola[19,47,48]
Connochaetes taurinus (Burchell, 1824)Northern South AfricaSouthern South AfricaHunting, restockingGenetic integrity of Connochaetes gnou (Zimmermann, 1780)[30]
Equus zebra hartmannae Matschie, 1898NamibiaWestern Cape, Eastern Cape (South Africa)IntroductionGenetic integrity of E. z. zebra Linnaeus, 1758[49]
Equus grevyi Oustalet, 1882KenyaKenya (outside natural range)Conservation (favor range expansion)Genetic integrity due to hybridization with E. quagga boehmi Matschie, 1892[50,51]
Aepyceros melampus petersi Bocage, 1879Namibia, AngolaAngola, NamibiaConservationPossible hybridization with A.m. melampus (Lichtenstein, 1812)[52]
Hippotragus equinus koba (Gray, 1872)West AfricaSouth AfricaHunting, restockinggenetic integrity of H. e. equinus (É. Geoffroy Saint-Hilaire, 1803)[19,53]
Redunca fulvorufula fulvorufula (Afzelius, 1815)South AfricaNamibiaConservationIntroduction in a new area outside the historic range; possible competition with other species, sanitary problems, etc.[54]
Tragelaphus angasii Angas, 1849 South AfricaBotswana, Namibia, AngolaHuntingCompetition and hybridization with
T. strepsiceros (Pallas, 1766); competition with T. scriptus (Angola)
Hippotragus niger (Harris, 1838) ssp.Tanzania, Zambia, Mozambique, MalawiSouth AfricaConservationGenetic integrity of different subspecies; possible hybridization with H. equinus[56,57]
Hippotragus equinus ssp.Namibia, Botswana, South AfricaSouth AfricaHunting, conservationGenetic integrity of different subspecies[58,59]
Antidorcas marsupialis marsupialis (Zimmermann, 1780)South AfricaSouth AfricaHunting, introductionPossible hybridization with A. m. hofmeyri Thomas, 1926[60]
Tragelaphus spekii gratus P. L. Sclater, 1880 (?)Unknown localityThe GambiaTourism, aesthetic reasonsIntroduction? reintroduction? population probably extinct[61]
Diceros bicornis michaeli Zukowski, 1965KenyaSouth AfricaConservation restocking Genetic integrity of D. b. bicornis (Linnaeus, 1758)[62,63]
Diceros bicornis michaeliSouth Africa (ranches), EAZA zoosTanzaniaConservation restocking, and reintroductionGenetic integrity of D. b. michaeli [64,65,66] (see reference [66] in case of ex situ stock)
Diceros bicornis ssp.All areasEast AfricaConservationPossible genetic erosion of Masai Mara population with recognized traces of gene pool of D. b. longipes Zukowski, 1949[66]
Ceratotherium simum simum (Burchell, 1817)Kenya (ranch)UgandaConservation?Introduction of one allochthonous subspecies into previous range of extinct C. s. cottoni (Lydekker, 1908) [67]
Ceratotherium simum simumSouth AfricaKenya, ZambiaConservationcreation of new nuclei outside the historic range[68]
Beatragus hunteri (Sclater, 1889)Southeast Kenya Kenya (Tsavo East NP)Conservation Creation of a new population outside the natural range[69,70]
Taurotragus oryx oryx (Pallas, 1766)South Africa SenegalTourism, aesthetic reasonsPossible competition, and genetic integrity of T. derbianus derbianus (Gray, 1847)[71,72,73]
Kobus ellipsiprymnus ellipsiprymnus (Ogilby, 1833)South AfricaSenegal Tourism, aesthetic reasonsGenetic integrity of K. e. unctuosus (Laurillard, 1842) [71,72]
Oryx gazella gazella (Linnaeus, 1758)South Africa SenegalTourism, aesthetic reasonsIntroduction in an area where it has never been present[71,72]
Panthera leo cf. melanochaita (H. Smith, 1842) (ex P. l. krugeri (Roberts, 1929))South Africa Rwanda“Reintroduction”Introduction outside the natural range. In Rwanda, was formerly present as P. l. azandica (Allen, 1924) [74]
Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Share and Cite

MDPI and ACS Style

Gippoliti, S.; Robovský, J.; Angelici, F.M. Taxonomy and Translocations of African Mammals: A Plea for a Cautionary Approach. Conservation 2021, 1, 121-136.

AMA Style

Gippoliti S, Robovský J, Angelici FM. Taxonomy and Translocations of African Mammals: A Plea for a Cautionary Approach. Conservation. 2021; 1(2):121-136.

Chicago/Turabian Style

Gippoliti, Spartaco, Jan Robovský, and Francesco M. Angelici. 2021. "Taxonomy and Translocations of African Mammals: A Plea for a Cautionary Approach" Conservation 1, no. 2: 121-136.

Article Metrics

Back to TopTop