Regional genetic structuring and evolutionary history of the impala Aepyceros melampus

Samples of 162 impala antelope (Aepyceros melampus) from throughout its distribution range in sub-Saharan Africa were surveyed using eight polymorphic microsatellite loci. Furthermore, 155 previously published mitochondrial DNA (mtDNA) sequences from the same localities were reanalyzed. Two subspecies of impala are presently recognized--the isolated black-faced impala (Aepyceros melampus petersi) in southwest Africa and the common impala (Aepyceros melampus melampus) abundant in southern and east Africa. All tests performed indicated significant genetic differentiation at the subspecific level. Furthermore, individual-based analyses split the common impala subspecies into two distinct genetic groups, conforming with regional geographic affiliation to southern or east Africa. This was supported by assignment tests, genetic distance measures, pairwise theta values, and analysis of molecular variance. We suggest that the presence of such previously unknown regional structuring within the subspecies reflects a pattern of colonization from a formerly large panmictic population in southern Africa toward east Africa. This scenario was supported by a progressive decline in population diversity indices toward east Africa and a significant increase in the quantity theta/(1 - theta). Both microsatellite and mtDNA data indicated a genetic distinctiveness of the Samburu population in Kenya.     

Phylogeography and conservation of impala and greater kudu

The phylogeography of the bush habituated African bovid species impala (Aepyceros melampus) and greater kudu (Tragelaphus strepsiceros) is investigated using mitochondrial DNA (mtDNA) markers. Combined analysis of individual lineages, relationships and population genetics suggest a colonization process from Southern Africa toward Eastern regions in the greater kudu. Results are less clear for the impala, although remaining consistent with a similar pattern of historical dispersion. The study reveals a similar pattern, that is a marked divergence of lineages from South-western Africa relative to other regions. This pattern is opposed to previously published findings in other African bovid species. In the impala, the genetically isolated region is consistent with morphology because it is recognized as the subspecies A. m. petersi, the black-faced impala. In contrast, the similar split of South-western mitochondrial lineages was not expected in the greater kudu on the basis of morphology. Both species show a significant population genetic differentiation. Beyond their phylogeographical value, our results should raise conservation concerns about South-western populations of both species. The black-faced impala is categorized as vulnerable and our data show indications of hybridization with common impala A. m. melampus. The previously unrecognized genetic status of the South-western kudus could also imply conservation regulations.     

The genetic status of naturally occurring black-nosed impala from northern South Africa

The impala (Aepyceros melampus ssp.) is a widespread antelope species occurring in sub-Saharan Africa. The two recognized subspecies have non-overlapping distribution ranges, with no known natural mixture of these subspecies until human interference. A number of common impala individuals (A. m. melampus) displaying phenotypic characteristics commonly observed in the black-faced impala (A. m. petersi), namely black facial markings, were seen on a farm in the Limpopo Province, South Africa. This farm falls outside the natural distribution range of A. m. petersi. We therefore aimed to identify the taxonomic placement of these individuals (i.e. A. m. melampus or A. m. petersi) through phylogenetic and molecular clock analyses using D-loop and cytochrome subunit b sequence data. Our results showed that these black-nosed impala from Limpopo are in fact A. m. melampus individuals. The existence of the black-nose phenotype in common impala could be more widespread than previously estimated. The occurrence of introgression between the two subspecies in this region could, however, not be fully excluded, and can only be fully assessed through the use of nuclear DNA analysis.     

A morphometric assessment of geographical variation and subspecies in impala

The single species of Aepyceros, Aepyceros melampus (impala), is native to central and southern Africa, from Uganda to South Africa. It inhabits open woodlands, sandy bush country and acacia savannah. This study tested the morphological and geographical variation among specimens of impala and their possible relation with described subspecies. Nineteen skull and horn measurements were taken. A multivariate analysis was used and size and shape were explored. Facial coat patterns were also coded. The results supported four out of five traditionally described subspecies: Aepyceros melampus petersi from Angola/Namibia, Aepyceros melampus melampus from South Africa, Aepyceros melampus suara from East Africa and Aepyceros melampus johnstoni from Zambia and Malawi. We consider Aepyceros melampus rendilis to be a synonym of suara . A revised synopsis is suggested, with comments on the geographical ranges of the subspecies. Skull dimensions also allowed us to estimate the geographical origin and subspecies of some individuals of unknown provenance. Coat patterns showed no clear relation with subspecies or geographic location, with the exception of A. m. petersi .     

Genetic structure of the common impala (Aepyceros melampus melampus) in South Africa: phylogeography and implications for conservation

We analysed 131 common impala (Aepyceros melampus melampus) samples from two provinces in South Africa (Limpopo and KwaZulu‐Natal) that are separated by the Drakensberg Mountain Range using sequences of the mitochondrial control region and seven polymorphic nuclear microsatellite loci. In line with earlier studies on bovid species, we found very high values of genetic diversity, particularly at the mtDNA locus with an overall nucleotide diversity of 3.6% and an overall haplotype diversity of 0.98. All statistical approaches confirmed a significant population differentiation between Limpopo and KwaZulu‐Natal, suggesting that areas of unsuitable habitat caused by the presence of the Drakensberg Range and the Indian Ocean coastal belt act as a barrier to gene flow. Only few individuals with signs of admixed origin were indicative of translocations or rare migration between the two provinces. Combination of our mtDNA data set with those of previous studies on impala from south‐western, southern and eastern Africa revealed the highest diversity in South Africa. This is in line with the hypothesis of a southern glacial refuge from which various African ungulate species spread northeast during the Holocene, although in the case of impala further analyses based on larger data sets will be necessary to definitively settle this question.     

Skin papillomas in an impala (Aepyceros melampus) and a giraffe (Giraffa camelopardalis).

Viral particles, typical of the papovavirus family, were demonstrated by electronmicroscopy in small papillomas found on the feet of an impala (Aepyceros melampus) and on the face of a giraffe (Giraffa camelopardalis) in Kenya. Histologically the tissues proved to be typical papillomas. The viral particles measured 38 nm and 40 nm in diameter in all tissue sections from the impala and giraffe respectively.     

Hyperthermia in captured impala (Aepyceros melampus): a fright not flight response

To investigate the patterns and mechanisms of capture-induced hyperthermia, we surgically implanted 26 impala (Aepyceros melampus) with miniature thermometric data loggers, which measured body temperatures continuously throughout capture procedures. Four groups of impala, which were habituated to varying levels of handling and boma-housing, were captured by net restraint or by chemical immobilization. The study took place between July 1999 and December 2005. Irrespective of whether impala were chemically captured, net-captured, or disturbed by exposure to a stressor, they developed a precipitous increase in body temperature. This increase in body temperature was not related to activity levels; animals that had low activity levels before immobilization had larger increases in body temperature compared to those that had high activity levels but were not immobilized (t = 3.6, P = 0.001, n = 5). Similarly this increase in body temperature was not related to environmental heat load at the time of darting and immobilization (r = -0.05, P = 0.85). Body temperature increase also did not depend on whether the animals were captured using drugs or not. However, we found that those animals that were habituated more to handling and boma-housing had smaller increases in body temperatures (F = 37, P<0.001) and smaller stress responses, indicated by lower plasma cortisol concentrations (F = 5.5, P<0.05), and less fractious behavior, compared to those animals that were habituated less or not at all. Therefore we believe that capture-induced hyperthermia in impala is caused predominantly by stress, which induces a rapid rise in body temperature.     

Hepatic lesions associated with Cooperioides hepaticae (Nematoda: Trichostrongyloidea) infection in impala (Aepyceros melampus) of the Kruger National Park

Intrahepatic biliary lesions were observed in two of 12 lambs, seven of 12 yearlings and 10 of 25 adult impala (Aepyceros melampus) surveyed in the Kruger National Park, Republic of South Africa. Lesions were associated with the nematode Cooperioides hepaticae, a trichostrongyloid parasite that inhabits the bile ducts of impala, and ranged from a mild chronic-eosinophilic cholangitis to foci of florid hyperplastic cholangitis with duct ectasia. The latter almost always contained viable worms and, after the worms died, the lesions appeared as foreign-body granulomas. Infection was acquired early in life; severe lesions were seen most frequently in yearlings. Adults were less severely infected, which suggested an acquired immunity. Although the incidence of infection was high, cooperiiasis did not appear to be a serious herd-health problem at the time of this study.     

In vitro succinylcholine hydrolysis in plasma of the African elephant (Loxodonta africana) and impala (Aepyceros melampus)

In elephants the time lapsed from i.m. injection of an overdose of the muscle relaxant succinylcholine (SuCh) until death, is significantly longer than in impala. To determine a difference in the rate of SuCh hydrolysis, once the drug enters the circulation, contributes to this phenomenon we have measured the rate of hydrolysis of SuCh in elephant and impala plasma, and by elephant erythrocytes. Rate of hydrolysis was determined by incubating SuCh in plasma or erythrocyte lysate at 37 degrees C and quantifying the choline produced. Plasma SuCh hydrolytic activity in elephant plasma (12.1+/-1.7 Ul(-1) mean+/-S.D.; n=9) was significantly higher than it was in impala plasma (6.6+/-0.6 Ul(-1); n=5), but were approximately 12 and 21 times lower, respectively, than in human plasma. Elephant erythrocyte lysate had no SuCh hydrolytic activity. Applying this data to previous studies, we can show that the ratio of SuCh absorption to SuCh hydrolysis is expected to be 1.25:1 and 1.41:1 for elephants and impala respectively. It will thus take at least 1.7 times longer for elephant to achieve a plasma SuCh concentration similar to that in impala. We conclude that a more rapid hydrolysis of SuCh in elephant plasma is one factor that contributes to the longer time to death compared to impala.     

Observations on the lungworm, Pneumostrongylus calcaratus, in impala (Aepyceros melampus) from Swaziland

The lungworm, Pneumostrongylus calcaratus, was found in 85% (164 of 193) of impala (Aepyceros melampus) collected in Mlawula Nature Reserve in Swaziland. Infection was confirmed at 4.5 mo of age, and the prevalence increased to 100% at 11 mo, with a prevalence of 98% in animals greater than 1 yr of age. Pneumostrongylus calcaratus was usually found in firm, tangrey nodules along the lobar borders of the lungs, although an extensive granulomatous pneumonia with miliary caseous abscesses and calcified nodules was observed in some older animals. In the primary infection in lambs, adult parasites, larvae and eggs were observed in the alveoli and bronchioles within the nodule. There was peribronchial and perivascular mononuclear cuffing, with infiltration of mononuclear cells in the alveolar septum in the vicinity of worms. In lesions in older animals, there was local consolidation with macrophages and giant cells, and foci of parenchymal necrosis associated with degenerating eosinophils, which appeared to lead to the formation of eosinophilic granulomas. Resolving lesions caused interstitial fibrosis, with mineralized nodules. Pneumostrongylosis does not appear to pose a significant threat to the health of impala in Swaziland.     

Loose front teeth: radiological and histological correlation with grooming function in the impala Aepyceros melampus

Casual observations have revealed that the anterior dentition of impala and other antelope is loosely embedded, with the tips of the teeth movable over a distance of 1middot;5 to 2mm. The comb-like anterior dentition of impala Aepyceros melampus is utilized extensively for grooming purposes, and it was hypothesized that the looseness of the teeth might be related to the grooming function. A sample of 12 impala mandibles was obtained from Pilanesberg National Park. Boputhatswana. Six of the incisor canine (IC) complexes were examined macroscopically, radiographically and histologically, while the remaining six were used to determine the alveolar depth relative to total root length, The findings were: (1) wide periodontal ligament spaces, most prominent in the apical region; (2) a loose, highly vascular periodontal ligament; (3) well-developed trans-septal periodontal ligament fibres; and (4) relatively shallow alveoli, with only approximately two-thirds of the roots included within the alveoli. In no case could looseness be ascribed to pathological changes in the periodontal ligament, cementum or alveolar bone. These features suggest that the looseness of the teeth is associated with a see-saw action of the teeth about a fulcrum below the alveolar bone crest. with the maintenance of the closed resting position of the teeth being facilitated by the well-developed trans-septal fibres. It is suggested that the minimal interdental space maintained by this arrangement during grooming assists in the efficient removal of parasites from the pelage by impala.     

黑斑羚粪便中碳同位素揭示的食性变化

利用稳定碳同位素数据(δ13C)分析了南非克鲁格国家公园混食性黑斑羚(Aepyceros melampus)时间和空间尺度上的食性变化,验证了两个假说,即有蹄类食性变化是由生境中木本植物与草本植物的相对配比导致;降雨控制有蹄类生态。结果表明:黑斑羚的食性涵盖了精食者-粗食者采食谱系,且食性中木本与草本比例在不同月间、季节、年度和区域间存在很大变化。栖息于开放性热带稀树草原和草原中的黑斑羚通常采食比生境中更高比例的草本,但在时间尺度上并不恒定。在克鲁格北部的一个区域(Punda Maria) ,黑斑羚采食的草本比克鲁格国家公园中其它任何区域都多。与其它生境相比,在河边的黑斑羚采食草本数量更少,尤其是在食性空间变化更为明显的旱季。因此,我们的数据不支持有蹄类食性组成变化是由生境中木本与草本比例不同造成的假说,食性与降雨量间也无明显的关系。我们的结果支持草本中蛋白含量增加引起黑斑羚采食比例的增加这一模型。粪便中氮含量在时间和空间上的变化很小,揭示在可利用食物中,无论木本还是草本,黑斑羚进行选择采食以保证最好的食物质量。基于这些结果,我们认为更具体的食物选择和可利用性最适采食理论能够更好地解释这种生态学变化。     

Characterization of nine microsatellite loci in impala (Aepyceros melampus)

Nine microsatellite loci were isolated from a genomic DNA library created from impala (Aepyceros melampus). Observed and expected levels of heterozygosity were computed utilizing 25 individuals from a population in central Kenya. Tests for Hardy–Weinberg equilibria were conducted and found that three of the nine loci deviated from equilibrium in this population. These markers were developed to analyse the genetic effects of culling and isolation on a game preserve in Kenya.     

Territorial behaviour in southern impala rams (Aepyceros melampus Lichtenstein)

The role of territoriality was investigated by studying 25 impala rams at a reserve in the Waterberg region of South Africa (23°45′S, 28°23′E). Mean territorial tenure was 67.25 days (range 23–99), with a mean territory size of 21.0 ± 11.27 ha, compared with home ranges of 34.1 ± 9.03 ha for territorial rams and 58.8 ± 33.35 ha for bachelor rams, using the fixed kernel method. Territory boundaries remained constant, whilst the area surrounding important features such as water holes, appears to be neutral in terms of territoriality. The rut, as evidenced from peaks in chasing and roaring, lasted for 2 months from 10 April to 10 June 2001, with intensified behaviour including matings observed from 16 May to 4 June 2001. Territorial rams chase and roar more than bachelors. Flehmen and display behaviours are performed by all rams, whilst fights and other reproductive behaviours are generally rare. Bachelors browse more than territorial rams. Only bachelors spar and allogroom, and orally groom themselves more than territorial rams.     

Geratodontology and horn growth of the impala (Aepyceros melampus)

Examination of the jaws of an impala population showed advanced wear on mandibular M1 compared with other ungulates which have been examined. This could lead to an erroneous interpretation of age if based upon mandibular tooth wear alone. Explanations are offered for this pattern in terms of the apparent pattern of wear of the impala molar teeth. Suggestions are also put forward for a method of determining specific age, from a conceptual wear model, when only extreme parameters are known. Horn growth in the male is also described.     

Seasonal variation in pituitary-gonadal function in free-ranging impala (Aepyceros melampus).

Blood, testicular biopsies and electroejaculates were collected from adult male impala, free-ranging in the Kruger National Park (Republic of South Africa), during the breeding (rut; April-May) and nonbreeding (September-October) seasons. Blood samples were collected at 5-min intervals for 120 min from anaesthetized males (n = 7 impala/group) treated intravenously with saline, gonadotrophin-releasing hormone (GnRH: 1 microgram/kg body weight) or human chorionic gonadotrophin (hCG: 10 or 30 iu/kg). Semen was collected from six more animals during the breeding season and 12 animals during the nonbreeding season using a standardized electroejaculation protocol. Ejaculates obtained during the nonbreeding season were of inferior quality to those collected during the breeding season, and were characterized by lower sperm concentrations, poorer sperm motility and more morphologically abnormal sperm forms. Within season, there were no differences in testosterone secretion between the two hCG doses, and these responses were similar to those observed after GnRH, but during the rut, testosterone secretion stimulated by both GnRH and hCG was approximately nine times greater than during the nonbreeding season. This seasonal increase in testosterone production was associated with a doubling in testicular volume and concentrations of luteinizing hormone (LH) receptors. Although concentrations of testicular follicle-stimulating hormone (FSH) receptors were similar between seasons, receptor content increased during rut as a result of increased testicular volume. In contrast to testosterone secretion, basal LH and FSH secretions were unaffected by season and GnRH-induced gonadotrophin secretion was reduced during rut.(ABSTRACT TRUNCATED AT 250 WORDS)     

Hunting differentially affects mixed-sex and bachelor-herds in a gregarious ungulate, the impala (Aepyceros melampus: Bovidae)

We investigated herd-sizes and herd-compositions of Impala ( Aepyceros melampus ) inside a protected area [Lake Mburo National Park (LMNP) in western Uganda] and the unprotected adjacent ranchland [the Ankole Ranching Scheme (ARS)]. Impala experience intense hunting and poaching in the study area, and poaching is especially strong on the ARS. We found evidence for changes in overall group-sizes in both mixed-sex and pure bachelor herds between areas in and outside LMNP. Mixed-sex herds strongly decreased in size outside the National Park, but bachelor herds even slightly increased in size. While the group-composition of mixed-sex herds was very similar in areas in and outside LMNP, bachelor herds comprised more yearlings and subadult males on the ARS. Our study suggests that effects of hunting and other human nuisance may differ between herd types: mixed herds probably decrease in size because females are more strongly hunted. Around LMNP, impala are usually hunted using nets and spears, thereby increasing the hunters' chance of being injured. Poachers therefore prefer hornless females (and their calves), as it is less dangerous to handle net-caught females than males. As a result, males are less hunted, but increased vigilance and, therefore, reduced aggression among the members of a bachelor herd, may account for the observed increase in herd sizes and changes in group-compositions.     

Sites of attachment and density assessment of ixodid ticks (Acari: Ixodidae) on impala (Aepyceros melampus)

During 1994 and 1995, 45 impala ewes, Aepyceros melampus were examined at Letaba Ranch in the Northern Province. Tick counts were made from 15 animals on three occasions, in July, October and February. The objective was to determine whether the total body tick counts can be estimated from counts done on specific, selected sampling sites on the skin. Twelve sites were shaved within ten predilection sites and the parasite numbers counted from these samples. These counts were compared to the total body parasite count as determined by the scrub and digestion techniques. More than 80% of all the ticks were present on the muzzle, the head, the pinna and the legs. Larger numbers of ticks were collected in October than in July or February. Two mathematical models were tested for the tick counts. The first model was made up of tick counts from a single shaved sampling site, the pinna. The correlation between the tick counts on the pinna and the total tick counts was highly significant (p values ranging between 0.0208 and 0.0001). The second model was developed based on tick counts from four regions, namely the head, the pinna and the front and hind feet. A less significant correlation was obtained between the number of ticks counted on the four sites and the total tick count.     

An examination of the infracommunities and component communities from impala (Aepyceros melampus) in the Kruger National Park, South Africa

The intestinal helminth parasites of the impala from the Kruger National Park, South Africa, were examined to describe the parasite community structure. Demographic variation and the associated differences in behavior were used to further investigate the patterns of community composition. Monte Carlo simulations were performed to test for differences in species richness and mean abundance between the various demographic groups, and nonmetric multidimensional scaling ordination was used to compare community composition. Seventeen species of nematodes, totaling more than 1.3 million worms, were recovered. Males harbored a greater number of nematode species than did females, but adult females were more heavily infected than their male counterparts. Lambs acquired infections early in life, and their parasite community composition rapidly approached that of the older animals. The parasite community in the juvenile and adult males was significantly different from the community of the adult females. These data suggest that social and feeding behavior of the different age-sex classes structure the parasite component community of impala. Additionally, the distinction between common and rare parasites, and their classification in other herbivores, implies complex transmission dynamics that includes extensive species sharing within the Kruger National Park.     

Stable isotope evidence for impala Aepyceros melampus diets at Akagera National Park,Rwanda

Stable isotope analysis of tooth enamel was used to investigate the relative proportions of grass and browse in seasonal and overall diets of impala Aepyceros melampus at Akagera National Park, Rwanda. Bulk enamel samples suggest that on average, impala ate c. 86% C₄ grass year‐round, far more than in most previously studied impala populations across Africa. Intra‐tooth samples show that seasonal changes in the proportion of C₄ grass versus C₃ browse are minimal (c. 10%), the diet being dominated by C₄ grass year‐round in contrast to other impala populations that consume ≥50% browse during the dry season. Intra‐tooth oxygen isotope values track carbon isotope changes to a moderate degree, but are not patterned clearly enough to permit identification of wet versus dry seasons. As other studies have shown that impala select high‐protein diets, the foraging behaviour at Akagera is probably because of the availability of palatable grass for much of the year in the edaphic grasslands around the lacustrine environments of the eastern portions of Akagera National Park.