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.     

Assessing the effect of roads on impala (Aepyceros melampus) stress levels using faecal glucocorticoid metabolites

In Tanzania, a proposed road through the Serengeti National Park has caused international debate as to whether it will cause irreversible damage to the ecosystem. We assessed the effect of current roads and traffic in the Serengeti ecosystem on a model ungulate species, the impala antelope (Aepyceros melampus). We measured how faecal glucocorticoid metabolites (FGM), an indicator of physiological stress levels, were affected by different levels of traffic while accounting for other stress‐related variables such as distance to roads, vegetation type and group size. A total of 196 faecal samples were collected. FGM were measured using an enzyme immunoassay (EIA) validated for several ruminants. FGM levels were significantly elevated near major roads with higher traffic levels compared to roads with less traffic. Predators and illegal hunting might also affect FGM levels in the impala, but the potential effect of these variables was inconsistent with the recorded FGM levels and cannot explain the observed variation. These results show for the first time that an African mammal exhibits significant physiological stress in relation to roads and traffic.     

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.     

Vigilance and group size in impala (Aepyceros melampus Lichtenstein): a study in Nairobi National Park, Kenya

One of the advantages of living in groups is that individuals may need to be less vigilant, allowing them more time for other important activities, such as foraging. This relationship between group size and per cent time spent being vigilant was investigated by observing impala in Nairobi National Park, Kenya. Three types of individual were observed: females, territorial males and bachelor males. Only females showed the predicted negative relationship between per cent vigilance and herd size. Both types of male showed no significant change of vigilance with increasing group size. There was no difference in levels of vigilance in open or closed habitats and no difference in vigilance between herds ‘alone’ and herds with other species that might have provided ‘extra eyes’.     

Fire induced changes in the foraging behaviour of impala Aepyceros melampus in Lake Mburo National Park, Uganda

The impact of fire on the foraging behaviour of impala (Aepyceros melampus), was studied in the Lake Mburo National Park, Uganda. Two indices of foraging efficiency were used to establish differences in feeding behaviour in burnt and unburnt control areas. These are acceptable grass/browse abundance (AGA/ABA), expressed in terms of feeding time achieved per 50 steps and food ingestion rate (FIR), expressed as the cumulative percentage. Accumulation of above ground grass biomass was also measured in burnt and control areas. After burning, the grazing efficiency remains constant throughout the dry season, while the browsing time achieved per 50 steps decreases drastically. In the control areas AGA decreased with increasing length of the dry season while ABA increased. When expressed in terms of cumulative percentage (FIR), the results show a significant increase of 31% in foraging time spent grazing and a 24% decrease in foraging time spent browsing, in burnt areas. The results confirm the classification of impalas as intermediate or mixed feeders with a distinct preference for grass. Impala foraging patterns after burning are discussed and compared with investigations on the effect of burning on the behaviour of impalas and Buffon's kob (Kobus kob kob) in other areas of Africa.     

Feeding ecology and foraging behaviour of impala Aepyceros melampus in Lake Mburo National Park, Uganda

Intersexual and seasonal variation in foraging behaviour of impala (Aepyceros melampus), was studied in the Lake Mburo National Park, Uganda. There was a moderate seasonal difference in foraging efficiency (as measured by ‘acceptable food abundance’), with a minimum in dry season and a maximum in Rainy season. The variation between sexes was more distinct with a pronounced minimum in time spent browsing of males in early wet season. By distinguishing between feeding time spent grazing and feeding time spent browsing the seasonal variation was confirmed. The proportion of foraging time spent feeding (expressed as ‘food ingestion rate’) showed an inverse pattern with a maximum in the late dry season (75.5%), decreasing values throughout the Rainy season and a minimum in early dry season (57.8%). Differences between sexes were explained in terms of reproductive demands and seasonal balance in terms of moderate climate throughout the year. Impala foraging patterns in the bimodal tropics (two Rainy seasons) is discussed and compared with unimodal tropics. The findings are matched against current ideas on optimal foraging.     

Effects of annual rainfall and habitat types on the body mass of impala (Aepyceros melampus) in the Zambezi Valley, Zimbabwe

Body mass is often considered as a good indicator of body condition of individuals in ungulates, hence of their fitness, and thus, may be used as an index to monitor the status of populations subject to harvesting schemes. Here, we report the influence of annual rainfall (a proxy for primary production) and habitat on the body mass of impala in a population cropped for meat in a communal area of Zimbabwe. We analyzed the data from 2 contrasted years for rainfall, in two different habitats. In the good year (i.e. high annual rainfall) impala were heavier than in the poor year, and adult females seemed to be less affected than males by variation in primary production. We show that adult males were suffering from a seasonal decrease in body mass, supposedly linked to the rut, particularly in good habitat. Overall, the habitat effect appeared to be dominated by the rainfall effect, and this may be due to the very high animal densities in the good habitat, i.e. fast resource depletion. Our results also suggest that males (juvenile and adult) are more susceptible to changes in food resource abundance and quality than females, which supports previous studies on sexually dimorphic and polygynous species.     

Differential grooming rate and tick load of territorial male and female impala, Aepyceros melampus

During the breeding season in Zimbabwe territorial male impalawere found to engage in much less self oral grooming and allogroomingthan females, presumably as a reflection of the need to remainvigilant in herding females and repulsing challenging bachelormales. Territorial males spent an average of 11 min engagedin all types of grooming during a 12-h day, compared with 40min grooming by females. Rutting activity and time spent scanningpeaked in May, while self oral grooming and feeding by territorialmales was lowest at this time. The decrement in grooming bymales (relative to females) represented half of all time devotedto rutting behaviors. Territorial males appeared to sacrificefeeding and grooming time in exchange for more time devotedto vigilant activities essential to mating success. In a comparableregion of Zimbabwe, territorial males were also found to harborabout six times as many adult ticks as the females on the sameterritories during the breeding season. Because grooming isunderstood to be effective in removing ticks, the higher tickload of territorial males was attributed to their reduced groomingbehavior. However, testosterone and adrenal cortical steroids,which are elevated in territorial male impala, are known todepress the immune system, and so may be important in controllingparasite infections. Sexually active males of many species aregenerally found to harbor more parasites than females. The differencein tick load between territorial male and female impala mayreflect both behavioral and hormonal parameters.     

Body condition and ruminal morphology responses of free-ranging impala (Aepyceros melampus) to changes in diet

Variation of the intraruminal papillation pattern with diet quality has been described in many ruminant species, but the use of papillation measures as a proxy for habitat quality and nutritional status of animals has not been evaluated. We compared various measures of body condition (body mass, body condition score, kidney fat index, bone marrow fat index, adrenal mass, kidney to adrenal mass ratio), diet quality (%browse, protein and fibre content) and rumen papillation in 106 impalas (Aepyceros melampus) from four different locations in Zimbabwe. The various condition proxies indicated that periods of low diet quality are characterised by a high proportion of browse in the diet of this species. Animals with a high proportion of browse had more voluminous rumens, suggesting a compensation for lower diet quality by increased intake. Macroscopic papillation indices did not yield meaningful significant correlations with diet quality or body condition proxies, and hence, their use for estimating habitat or body condition cannot be advocated. In contrast to previous histological reports, ballooning cells of the Stratum corneum of the ruminal mucosa were more prominent in animals on lower-quality diets. There were significant correlations of the kidney to adrenal mass ratio with other body conditions and with diet quality indices, suggesting that poor body condition and low diet quality represent stressful situations.     

No suggestion of hybridization between the vulnerable black-faced impala (Aepyceros melampus petersi) and the common impala (A. m. melampus) in Etosha National Park, Namibia

There are two recognized subspecies of impala in sub-Saharan Africa: the common impala (Aepyceros melampus melampus) -- widespread in southern and east Africa -- and the vulnerable black-faced impala (A. m. petersi) -- found naturally in only a small enclave in southwest Africa. The Etosha National Park (NP) in Namibia harbours the largest and only protected-area population of black-faced impala, numbering some 1500 individuals. Due to translocations of the exotic common impala to commercial farms in Namibia during the past decades, the black-faced impala in Etosha is faced with the potentially serious threat of hybridization posed by secondary contact with the common impala inhabiting bordering farms. Using eight microsatellite DNA markers, we analysed 127 black-faced impala individuals from the five subpopulations in Etosha NP, to determine the degree, if any, of hybridization within the park. We found that (a) the black-faced impala were highly genetically differentiated from the common impala (pairwise theta-values ranged from 0.18 to 0.39 between subspecies; overall value = 0.27) and (b) black-faced samples showed high levels of genetic variability [average expected heterozygosity (H(E)) = 0.61 +/- 0.01 SE], although not as high as that observed in the common impala (average H(E) = 0.69 +/- 0.02 SE). (c) No hybridization between the subspecies in Etosha was suggested. A Bayesian Markov Chain Monte Carlo approach revealed clear distinction of individuals into groups according to their subspecies of origin, with a zero level of 'genetic admixture' among subspecies.     

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.     

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.     

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.     

Foraging behaviour of wild impala (Aepyceros melampus) and Burchell's zebra (Equus burchelli) in relation to sward height

Foraging behaviour plays a key role in the interaction between herbivores and vegetation, their predominant food source. Understanding this interaction is crucial to providing information that is useful for conservation of herbivores. The objective of this study was to determine how sward height influences functional response and movement patterns of free ranging wild impala and zebra at the Kenya Wildlife Service Training Institute. The study was conducted for 3 months. Sward height is an important parameter that indicates how intensive a sward has been grazed and it influences intake rate through its effect on bite size. Bite size, instantaneous intake rate, specific mass intake rate and feeding station interval for impala and zebra increased with sward height. Sward height in combination with an animal's sex was found to have a profound effect on specific mass intake rate in impala. Zebra had a longer feeding station interval and lower stepping rate in tall swards compared to impala. Despite differences in their specific body mass and digestive strategies, impala and zebra maximized their intake rates in tall swards as a trade‐off among the swards. Tall swards are therefore critical in the study area and should be protected from bush encroachment which is a persistent problem.     

Observations on the life-cycle and larval morphogenesis of Cooperia fuelleborni (Nematoda: Trichostrongyloidea) parasitic in impala, Aepyceros melampus

The previously unknown life-cycle of Cooperia fuelleborni, parasitic in impala, is described. The morphology of the developmental stages are compared to those of other Cooperia spp. The similarity between C. fuelleborni and C. curticei is discussed as well as the differences between those two species. The morphological studies showed that C. fuelleborni is closely related to C. curticei of sheep, but also has some affinity with the cattle parasite C. punctata and C. oncophora.     

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.     

Class I antigens of the bovine major histocompatibility system are weakly associated with variation in faecal worm egg counts in naturally infected cattle

Three bulls selected for high faecal worm egg counts and three bulls selected for low faecal worm egg counts were mated to Africander-Hereford cross cows. Faecal worm egg counts were taken on four occasions from the 132 offspring. Also, each animal was typed for 32 class I antigens of the bovine major histocompatibility system (BoLA). Least squares analysis of variance showed that line, sex and some of the antigens were associated with differences in worm egg output in the faeces. After adjusting for the effects of line and sex, cattle with antigen W9 had about twice as many worm eggs in their faeces as cattle without W9; cattle with antigen CA45 had about half the concentration of faecal worm eggs as cattle without CA45. However, the antigen associations were of borderline significance at the 5% level and more work in additional populations is necessary to confirm these associations.     

Estimating density of Kirk’s dik‐dik (Madoqua kirkii Günther), impala (Aepyceros melampus Lichtenstein) and common zebra (Equus burchelli Gray) at Mpala,Laikipia District,Kenya

This study compares two methods of estimating the density of three ungulate species: Kirk’s dik‐dik, impala and common zebra, in a dry savannah ecosystem. Fixed strip width and distance sampling involving direct animal counts were used in parallel, and tested for bias and precision in two habitats. Distance sampling was the method that achieved a better balance between accuracy and precision. The differences in sightability of ungulates, according to body size, colour and habitat characteristics, are taken into account by the distance sampling method, but not by the strip method, which produced, what we suspect are, underestimates of animal density.     

Seasonal parasitism by egg parasites of the yellow rice borer,Scirpophaga incertulas [Lepidoptera: Pyralidae]

Scirpophaga incertulas (Walker) [Lep.: Pyralidae] is a common pest of rice in West Bengal, India. A three year study (1978–1981) revealed that egg parasites were important in reducing the borer population in an uninterrupted rice cultivation practice. The parasite activity showed periodical fluctuation coinciding with the moth emergence. It was evident thatTelenomus spp. were most common and most dominating in the early summer crop and became highly adaptive under diverse crop growing situations. Among three species ofTelenomus, T. rowani Gahan was the most abundant durint the summer and winter seasons.Tetrastichus schoenobii Ferriere was second in importance in regulating the borer population and its attack was heaviest on the later broods in rice crops cultivated during winter.Trichogramma japonicum Ashmead was very sporadic and had a particular preference for the dry period. It usually occurred in association with other two parasites in the later part of the year. Parasitism of egg masses by two or more species of the parasites was also common during the peak period of parasite activity.