Genetic variability in south african blue wildebeest (Connochaetes taurinus)

• 1.1. We used protein gel-electrophoresis to investigate genetic heterogeneity at 33 protein coding loci in a total of 46 blue wildebeest (C. taurinus) kept under different management regimes.
• 2.2. Average heterozygosity ranged from 2.14 to 4.3% and within-population differences accounted for 97.2% of total relative gene diversity.
• 3.3. Comparatively little divergence was found between animals sampled from populations with very diverse population sizes and management histories, with the largest genetic distance estimated between any two populations being only 0.0021.
• 4.4. We discuss our results with particular emphasis on the influence of management history on genetic diversity and divergence in C. taurinus.

     

Isolation and characterization of species-specific microsatellite markers for blue and black wildebeest (<Emphasis Type="BoldItalic">Connochaetes taurinus</Emphasis> and <Emphasis Type="BoldItalic">C. gnou</Emphasis>)

The blue wildebeest (Connochaetes taurinus) is distributed throughout southern and east Africa while the black wildebeest (Connochaetes gnou) is endemic to South Africa and was driven to near extinction in the early 1900s due to hunting pressure and disease outbreaks. Extensive translocation of both species throughout South Africa is threatening the genetic integrity of blue and black wildebeest. To effectively manage these species, genetic tools that can be used to detect hybrid individuals, identify genetically unique subpopulations and determine the levels of genetic diversity are required. In this study, 11 microsatellite markers were developed for wildebeest through next-generation sequencing. The microsatellite loci displayed 2.00–4.14 alleles, unbiased heterozygosity values ranged from 0.32 to 0.60 and observed heterozygosity values ranged from 0.26 to 0.52. The comparatively high level of polymorphism observed in the microsatellite markers indicates that these markers can contribute significantly to our knowledge of population genetic structure, relatedness, genetic diversity and hybridization in these species.     

Trophic ecology of two savanna grazers,blue wildebeest <Emphasis Type="Italic">Connochaetes taurinus</Emphasis> and black wildebeest <Emphasis Type="Italic">Connochaetes gnou</Emphasis>

The feeding niches and trophic ecology of two South African grazers, blue wildebeest Connochaetes taurinus and black wildebeest Connochaetes gnou, are compared using stable carbon and nitrogen isotope data from feces and tooth dentine collagen. As sympatric, closely related taxa predicted to occupy similar trophic positions, the blue and black wildebeest provide a good model for studying the mechanisms of coexistence and macroevolution in mammals. Data from feces collected from a single reserve in the Free State Province reveal different trophic behaviors between two herds of blue wildebeest and between both compared with a single herd of black wildebeest. These data suggest that sympatric coexistence of blue and black wildebeest is facilitated by differential niche occupation at family group or herd levels, rather than between species. However, such separation does not occur over longer time scales: results from dentine collagen support the hypothesis that the two species are indistinct in terms of trophic behavior, although blue wildebeest show more feeding flexibility, probably because of their wider habitat tolerance range. Similarities in premaxillary width of males and females of both species also suggest that both species are adapted to similar feeding styles. Thus, it is unlikely that changes in trophic behavior provided the trigger for divergence of the black from the blue wildebeest lineage in the Middle Pleistocene. We argue that the case of these two species represents an example of speciation that was not driven by resource competition, as is often assumed for many turnover events in mammalian evolution. We briefly discuss a previous suggestion that links black wildebeest evolution to their more territorial breeding behavior associated with Middle-to-Late Pleistocene landscape changes in southern Africa.     

Black wildebeest seek shade less and use solar orientation behavior more than do blue wildebeest

Many ungulates, including wildebeest, seek shade and orient their bodies relative to incoming solar radiation in order to reduce environmental heat loads. Blue (Connochaetes taurinus) and black wildebeest (Connochaetes gnou), which co-exist artificially in some reserves in South Africa, are thought to adopt different thermoregulatory behaviors to mitigate high environmental heat loads. However, whether or not blue and black wildebeest use different behaviors to reduce heat loads in regions where they co-occur has never previously been examined. We compared the shade seeking and solar orientation behavior of free-ranging blue and black wildebeest in summer at three locations in South Africa where both species co-occur. We found that blue wildebeest exhibited more shade seeking behavior than did black wildebeest at all times of day, at all study sites. Black wildebeest remained in the sun but were more likely than blue wildebeest to orient their bodies parallel to the sun at all study sites, a behavior which reduces the amount of surface area exposed to incoming radiation. Black wildebeest were most likely to employ parallel solar orientation during the hottest times of the day when the sun was not directly overhead (i.e., solar noon ± 1 hour). We thus demonstrate that co-occurring blue and black wildebeest use different thermoregulatory behaviors to reduce high heat loads. It is possible that the lack of shade in the historical distribution of black wildebeest led to selective pressure for reliance on solar orientation. Differences in thermoregulatory behavior can affect species-specific heat loads, habitat use, body mass, fitness and grazing activity. Such differences may also allow blue and black wildebeest to inhabit separate microclimates within the same habitat, provided there is sufficient heterogeneity in vegetation structure, potentially facilitating reproductive isolation.     

Assessing introgressive hybridization between blue wildebeest (<Emphasis Type="Italic">Connochaetes taurinus</Emphasis>) and black wildebeest (<Emphasis Type="Italic">Connochaetes gnou</Emphasis>) from South Africa

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.     

Dietary composition of wildebeest (Connochaetes taurinus) kongoni (Alcephalus buselaphus) and cattle (Bos indicus), grazing on a common ranch in south-central Kenya

This paper gives the results of a study conducted at Game Ranching Ltd, situated at the Athi Kapiti Plains, Kenya, to determine the dietary composition of cattle (Bos indicus L), kongoni (Alcephalus buselaphus Pallas) and wildebeest (Connochaetes taurinus Burchell), through two forage growing seasons (wet and dry). The dietary preferences and overlaps between the species were also determined. The diet preference of the three herbivores was determined using a microhistological analysis technique. The three dominant grasses in the ranch: Themeda triandra Fork, Digitaria macroblephara (Hack.) Stapf and Penisetum mezianum Leeke, formed the major diets of the animals. The browse component of the diets increased during the dry season by about 100% for all the animal species, with cattle always having twice as much browse as the wild herbivores. However, the animals selected similar diets in terms of plant species during both seasons but were more selective during the wet season. Dietary overlaps were always above 75%, but more than this during the dry season. The dietary overlaps were lower between cattle and wild herbivores than between the wild herbivores. The diet preference index for the animals differed from season to season, and from plant to plant.     

Rumen Ciliates of Musk-Oxen (Ovibos moschatus Z.) from the Canadian Arctic1

Samples of rumen contents were obtained from 13 musk-oxen living on Banks Island, N.W.T., Canada. Total protozoan numbers ranged from 38.5 to 124.6 times 10⁴ per ml of rumen contents with a mean generic distribution of 52.4%Entodinium, 45.9%Diplodinium, and 1.7%Epidinium. A total of 18 protozoan species were found, six of which have not previously been observed in this host, i.e. D. (Diplodinium) costatum, D. (Ostracodinium) gracile, D. (Ostracodinium) trivesiculatum. D. (Eudiplodinium) medium, Epidinium quadricaudatum, and Epidinium parvicaudatum. Two new species of protozoa are described, Entodinium ovibos n. sp. and Diplodinium (Eudiplodinium) banksi n. sp.     

Rumen Ciliate Protozoa of the Blue Duiker (Cephalophus monticola), with Observations on Morphological Variation Lines Within the Species Entodinium dubardi

ABSTRACT. Protozoal concentrations were determined in rumen and cecal contents of 20 blue duikers ( Cephalophus monticola ). Ten animals of each sex were fed either a high concentrate or high roughage diet. Rumen protozoa were present in 19 of the 20 animals and concentrations ranged from 4.5 to 33.7 × 10⁶ per g of rumen contents. At the higher concentrations, protozoal cells equaled between 30–40% of the total rumen contents volume. No protozoa were found in cecal contents. Weight of rumen contents was higher in females than in males ( P < 0.01), and rumen protozoa concentrations were higher in males ( P < 0.05) and in those animals fed the high concentrate diet ( P < 0.05). All the protozoa were identified as belonging to a single species, Entodinium dubardi . However, an average of about 30% of the E. dubardi cells varied from the typical morphology of this species. These cells appeared to be on variation lines leading toward 7–10 other non-caudate species of Entodinium . The present data were used to evaluate and discuss the concept of variation lines within E. dubardi .     

The development of genome-wide single nucleotide polymorphisms in blue wildebeest using the DArTseq platform

Blue wildebeest (Connochaetes taurinus taurinus) are economically important antelope that are widely utilised in the South African wildlife industry. However, very few genomic resources are available for blue wildebeest that can assist in breeding management and facilitate research. This study aimed to develop a set of genome-wide single nucleotide polymorphism (SNP) markers for blue wildebeest. The DArTseq genotyping platform, commonly used in polyploid plant species, was selected for SNP discovery. A limited number of published articles have described the use of the DArTseq platform in animals and, therefore, this study also provided a unique opportunity to assess the performance of the DArTseq platform in an animal species. A total of 20,563 SNPs, each located within a 69 bp sequence, were generated. The developed SNP markers had a high average scoring reproducibility (>99%) and a low percentage missing data (~9.21%) compared to other reduced representation sequencing approaches that have been used in animal studies. Furthermore, the number of candidate SNPs per nucleotide position decreased towards the 3′ end of sequence reads, and the ratio of transitions (Ts) to transversions (Tv) remained similar for each read position. These observations indicate that there was no read position bias, such as the identification of false SNPs due to low sequencing quality, towards the tail-end of sequencing reads. The DArTseq platform was also successful in identifying a large number of informative SNPs with desirable polymorphism parameters such as a high minor allele frequency (MAF). The Bos taurus genome was used for the in silico mapping of the marker sequences and a total of 6020 (29.28%) sequences were successfully mapped against the bovine genome. The marker sequences mapped to all of the bovine chromosomes establishing the genome-wide distribution of the SNPs. Moreover, the high observed Ts:Tv ratio (2.84:1) indicate that the DArTseq platform targeted gene-rich regions of the blue wildebeest genome. Finally, functional annotation of the marker sequences revealed a wide range of different putative functions indicating that these SNP markers can be useful in functional gene studies. The DArTseq platform, therefore, represents a high-throughput, robust and cost-effective genotyping platform, which may find adoption in several other African antelope and animal species.     

Occurrence of the Rumen Ciliate Oligoisotricha bubali in Domestic Cattle (Bos taurus)

Oligoisotricha bubali, previously observed twice in water buffalo, was detected in rumen contents of domestic cattle (Bos taurus) in two different areas of Tennessee. Concentrations ranged from <1 to 35% of the total protozoa in unweaned calves and up to 72% in older animals in feedlot. In contrast to the other genera of holotrichs, both total numbers and percent composition of O. bubali increased when animals were fed a corn silage-concentrate diet.     

Subspecific identity and a comparison of genetic diversity between wild and ex situ wildebeest

The original North American ex situ wildebeest population was believed to originate from the white-bearded wildebeest (Connochaetes taurinus albojubatus), which is both morphologically distinct and geographically separated from the brindled wildebeest (C. t. taurinus). However, after an import of wildebeest into North America in 2001, managers have suspected that white-bearded and brindled wildebeest were mixed in herds at multiple institutions. We sequenced the mitochondrial control region (d-loop) from a portion of the managed North American population and compared our sequences with previously published sequences from wild individuals to determine the subspecific identity and genetic diversity of our ex situ population. We were able to confidently identify C. t. albojubatus as the subspecies identity of the sampled portion of our population. Within our population, haplotype and nucleotide diversity were low (0.169 and 0.001, respectively) with a single common haplotype (H1) containing 41 of the 45 individuals sequenced, while two rare haplotypes (H2 and H3) were derived from three individuals and a single individual, respectively. Nucleotide and haplotype diversity were greater overall in the wild populations compared with our managed population. However, C. t. albojubatus was found to exhibit lower nucleotide diversity in both wild and ex situ populations when compared to other wild subspecies. Though the overall goal of the North American wildebeest population is for public education and not reintroduction, maintaining genetic diversity is vital for the long-term viability of this managed population, which may benefit from periodic supplementation of wild animals.     

Grass greenness and grass height promote the resource partitioning among reintroduced Burchell's zebra and blue wildebeest in southern Mozambique

Differences in the selection of habitat and specific dietary items support resource partitioning and coexistence of sympatric African grazing herbivores, such as zebra and wildebeest. In Maputo Special Reserve (MSR), southern Mozambique, these two species were extirpated during the civil war (1977–1992); since 2010, they have been reintroduced into the Reserve. Identifying the resource selection by reintroduced species and how these species coexist, while utilising the same resources, is both of ecological interest and important for the management of wildlife communities and parks. This is a key application of our research. Therefore, the present study investigated resource partitioning between Burchell's zebra (Equus burchelli, Smuts 1832) and blue wildebeest (Connochaetes taurinus, Burchell 1823) in the MSR. We conducted the study from July 2016 to June 2017. The data were collected by direct observation, driving the vehicle along the reserve's roads that covered the vegetation communities where zebras and wildebeest are known to commonly occur. The composition of the diet and specific features of the grass grazed by the two species, including greenness, height, and the number of stems, were assessed. The widely available grass, Aristida barbicollis, contributed most to the diet of both herbivores. The dietary overlap between the two herbivores was higher during the dry season (95%) than wet season (86%). Resources partitioning appears to be determined, principally, by the height and greenness of the grass, with the zebra grazing taller grass, which may facilitate the access of the wildebeest to the greener, lower proportion of the forage. That results follow the expectation that, among native herbivores, overlap in resource use is not expected based on evolutionary segregation.     

Ciliate Protozoa in the Rumen of Brazilian Water Buffalo,Bubalus bubalis Linnaeus*

SYNOPSIS Total numbers and distribution of genera, subgenera and species were determined for the ciliate protozoa in rumen contents of 4 Brazilian water buffalo Bubalus bubalis Linnaeus. The fauna of one animal, housed in close proximity to European and zebu-type cattle, differed considerably from that of the remaining animals, which were somewhat isolated on a large ranch. Several of the protozoan species observed in the semi-isolated animals were first described in rumen contents from humped Indian cattle, and their subsequent occurrence in other hosts and geographic locations has been limited or absent. In all, 49 different species of protozoa were found, 8 of which have not been previously described. Three of the new species belong to the genus Entodinium: E. ciculum sp. n., E. spinonucleatum sp. n. and E. triangulum sp. n.: 4 to Diplodinium (Ostracodinium): D. (O.) brazili sp. n., D. (O.) esalqum sp. n., D. (O.) nucleolobum sp. n., and D. (O.) tiete sp. n.; and one to Diplodinium (Eudiplodinium): D. (E.) bubalus sp. n.     

Ciliate protozoa in the rumen of Brazilian water buffalo, Bubalus bubalis Linnaeus.

Total numbers and distribution of genera, subgenera and species were determined for the ciliate protozoa in rumen contents of 4 Brazilian water buffalo Bubalus bubalis Linnaeus. The fauna of one animal, housed in close proximity to European and zebu-type cattle, differed considerably from that of the remaining animals, which were somewhat isolated on a large ranch. Several of the protozoan species observed in the semi-isolated animals were first described in rumen contents from humped Indian cattle, and their subsequent occurrence in other hosts and geographic locations has been limited or absent. In all, 49 different species of protozoa were found, 8 of which have not been previously described. Three of the new species belong to the genus Entodinium: E. ciculum sp. n., E. spinonucleatum sp. n. and E. triangulum sp. n.; 4 to Diplodinium (Ostracodinium): D. (O.) brazili sp. n., D. (O.) esalqum sp. n., D. (O.) nucleolobum sp. n., and D. (O.) tiete sp. n.; and one to Diplodinium (Eudiplodinium): D. (E.) bubalus sp. n.     

Management of hybridization in an endemic species: decision making in the face of imperfect information in the case of the black wildebeest—<Emphasis Type="Italic">Connochaetes gnou</Emphasis>

Hybridization between introduced and endemic ungulates, resulting from anthropogenic actions, has been reported for several species. Several studies of such events contain the common themes of extralimital movements, problematic phenotypic and genetic detection, and imperfect management. In southern Africa, the endemic black wildebeest (Connochaetes gnou) currently faces a serious threat of hybridization and introgression. This species survived near extinction and consequent genetic bottlenecks in the late 1800s and in the 1930s. Initiatives by private farmers followed by conservation authorities led to a dramatic recovery in numbers of this species. However, in an ironic twist, the very same advances in conservation and commercial utilisation which led to the recovery of numbers are now themselves threatening the species. Injudicious translocation has brought the species into contact with its congener, the blue wildebeest (Connochaetes taurinus), and in recent times, hybridization between the species has occurred at numerous localities in South Africa. Consequently, a significant proportion of the national black wildebeest population potentially carries a proportion of introgressed blue wildebeest genetic material. We discuss completed and ongoing attempts to find molecular markers to detect hybrids and highlight the difficulty of detecting advanced backcrosses. Additional avenues of research, such as work on morphology (cranial and postcranial elements), estimating of the probability of introgression and modelling of diffusion rates are also introduced. In addition to the difficulty in detecting hybrid animals or herds, the lack of consensus on the fate of hybrid herds is discussed. Finally, in an environment of imperfect information, we caution against implementation of management responses that will potentially induce a new genetic bottleneck in C. gnou.     

Effect of Absence of Ciliate Protozoa From the Rumen on Microbial Activity and Growth of Lambs

A survey of the components of the rumen ciliate population in a series of adult sheep, raised in the Faculty of Agriculture, University of Alexandria, has shown that a mixture of Entodinium, Isotricha, Ophryoscolex, Diplodinium, and Polyplastron species was found in the rumen contents of Egyptian sheep; no Epidinium and a negligible number of Dasytricha ruminantium were also observed. The microbial population, reducing sugars, ammonia, volatile fatty acids (VFA) production, and growth rate of 14 lambs inoculated with whole rumen contents from a mature sheep were compared over a 6-month period with those of 13 lambs maintained under the same conditions, except that they were strictly isolated from other ruminants. Certain large oval organisms and large numbers of flagellates and Oscillospira were frequently observed in the rumen contents of the isolated lambs. The reducing sugars, ammonia, and VFA levels, measured before and at intervals after feeding, in the inoculated lambs showed a pronounced rise above the values found in the ciliate-free animals. The propionic acid-acetic acid ratio in the rumen contents of the faunated lambs was considerably higher than in the nonfaunated controls. The inoculated lambs grew faster than the isolated lambs. Differences in weight gain which ranged from 15 to 17% were statistically significant. The inoculated animals impressed the observers by their good appearance which was superior to that of the ciliate-free lambs. It was, therefore, concluded that the rumen ciliate protozoa are essential for the metabolism and growth of young lambs.     

Orientation to solar radiation in black wildebeest (<Emphasis Type="Italic">Connochaetes gnou</Emphasis>)

We recorded the body axis orientation of free-living black wildebeest relative to incident solar radiation and wind. Observations were made on three consecutive days, on six occasions over the course of 1 year, in a treeless, predominantly cloudless habitat. Frequency of orientation parallel to incident solar radiation increased, and perpendicular to incident solar radiation decreased, as ambient dry-bulb temperature or solar radiation intensity increased, or wind speed decreased. We believe these changes were mediated via their effect on skin temperature. Parallel orientation behavior was more prominent when the wildebeest were standing without feeding than it was when they were feeding. We calculate that a black wildebeest adopting parallel orientation throughout the diurnal period would absorb 30% less radiant heat than the same animal adopting perpendicular orientation. Parallel orientation was reduced at times when water was freely available, possibly reflecting a shift from behavioral to autonomic thermoregulatory mechanisms. The use of orientation behavior by black wildebeest is well developed and forms part of the suite of adaptations that help them to maintain heat balance while living in a shadeless, often hot, environment.     

Water provision alters wildebeest adaptive habitat selection and resilience in the Central Kalahari

Wildlife populations in semi‐arid regions require unrestricted mobility along ecological gradients and across large landscapes to enable adaptive responses to seasonal variability and patchy resources. In the Kalahari region of Botswana, herbivore populations historically depended on seasonal access to the nutrient‐rich Schwelle area in the wet season and to water from the Boteti River during drought periods. Blue wildebeest Connochaetes taurinus in the Central Kalahari Game Reserve (CKGR) have lost access to these key habitats due to fences and encroachment of livestock and humans. We deployed satellite collars onto 10 female wildebeest in the CKGR to examine seasonal movements and habitat selection in relation to the environmental conditions and fragmented ecosystem. Wildebeest favoured open, short‐grass pan habitats in all seasons, probably in response to better forage quality and lower predation risk. The ability to remain in pan habitats during the dry season was a result of artificial water provision. A wildebeest herd that had no artificial water in its home range survived the dry season, whereas those wildebeest that were accustomed to water provision died when their water points failed in the dry season. Thus, water provision altered adaptive behaviour and reduced resilience of the population to the arid environment.     

Antibodies to Brucella spp. among blue wildebeest and African buffalo in Kenya

A serologic survey of blue wildebeest (Connochaetes taurinus Burchell) and African buffalo (Syncerus caffer Sparrman) in the Masai Mara area was conducted. Antibodies to Brucella spp. were found in 18% of the blue wildebeest and 30% of the African buffalo examined. There were titers in all age groups and in both sexes. Hygromata were seen in both species. The increase in numbers of blue wildebeest and African buffalo which share grazing and watering areas with cattle of the Masai people, makes the presence of infections by Brucella spp. in wildlife an important consideration in any program for control of brucellosis.     

Prey selection and prey preferences of spotted hyenas <Emphasis Type="Italic">Crocuta crocuta</Emphasis> in the Etosha National Park,Namibia

The feeding ecology of the spotted hyena Crocuta crocuta was studied in the central and eastern part of the Etosha National Park, Namibia. Hyenas mainly hunted migratory ungulates such as springbok Antidorcas marsupialis, zebra Equus burchelli, and blue wildebeest Connochaetes taurinus, but also resident species such as the greater kudu Tragelaphus strepsiceros and gemsbok Oryx gazella. There were, however, major differences in the species most frequently killed by hyenas in central and eastern Etosha. The preferred prey species of spotted hyenas in central Etosha was springbok, whereas regarding the abundance of zebra and wildebeest, these two species were rather avoided. In contrast, the prey species preferred by hyenas in eastern Etosha was kudu. Zebra and gemsbok were taken in proportion to their abundance, whereas wildebeest and springbok seemed to be rather avoided. Differences in prey selection and preferences were also reflected in differences in hunting group sizes. In eastern Etosha, where spotted hyenas frequently hunted larger prey, hunting group sizes were significantly larger compared to those in the center of the park.