Investigating a simplified method for noninvasive genetic sampling in East African mammals using silica dried scat swabs

Swabbing scat has proved to be an effective noninvasive method to collect DNA from mammals in the field. Previously, this method has relied on preservative liquids or freezing to preserve the DNA collected on swabs. In this study, we determine the effectiveness of using silica to simply dry the swab in field as an alternative way to prevent DNA degredation. Four species were included in the study; reticulated giraffe, impala, fringe‐eared oryx, and lion. Swabs were taken at multiple time points for giraffe and impala scat samples, with the lion and oryx sampled opportunistically. Mitochondrial DNA was successfully amplified and sequenced from scat swabs from all species; however, effectiveness varied between species, with 81.8% amplification success rate from swabs taken from impala scat compared to 25% amplification success rate in giraffe. This variation in success rate was overcome by taking multiple swabs, thus increasing the probability of a successful amplification. The true merit of this method is in its simplicity and cheapness; no preservative liquids were required to be brought into the field, at no stage in the 2 weeks of field sampling were samples frozen, and no commercial kits were used for DNA extraction.     

Antipredator behaviour of African ungulates around human settlements

As human populations grow and come into more frequent contact with wildlife, it is important to understand how anthropogenic disturbance alters wildlife behaviour. Using fine‐scale spatial analyses, we examined how proximity to human settlements affects antipredator responses of ungulates. We studied seven common ungulate species (Kirk's dik‐dik, Thomson's gazelle, impala, common warthog, common wildebeest, common zebra and Masai giraffe) in the Tarangire–Manyara ecosystem in northern Tanzania. In zebra and giraffe, flight responses to humans were significantly more likely when closer to settlements; however, there was a weak relationship between flight responses and distance to settlement in all other species. While there was largely a weak relationship between proximity to human settlements, the distribution of settlements in the landscape appears to affect wildlife behaviour, suggesting that animals perceive and respond to spatial variation in risk exerted by humans.     

Long term monitoring of mammal populations in the Serengeti: census of non-migratory ungulates, 1971*

Long term parallel studies on animal populations can give information on problems concerning population regulation, competition and predation. So far only two populations in the Serengeti have been studied in this way. To initiate long term studies on several ungulate species an aerial transect census was carried out in 1971. The methods of counting and the statistical analysis are described in detail so that future workers can repeat the census. The results are presented for impala, topi, kongoni, giraffe, waterbuck, eland and buffalo male groups. Based upon the confidence limits of the samples, it is suggested that the next census be carried out in June 1974.     

Dynamics of births and juvenile recruitment in Mara–Serengeti ungulates in relation to climatic and land use changes

Natality and recruitment govern animal population dynamics, but their responses to fluctuating resources, competition, predation, shifting habitat conditions, density feedback and diseases are poorly understood. To understand the influences of climatic and land use changes on population dynamics, we monitored monthly changes in births and juvenile recruitment in seven ungulate species for 15 years (1989–2003) in the Masai Mara Reserve of Kenya. Recruitment rates declined for all species but giraffe, likely due to habitat alteration and increasing vulnerability of animals associated with recurrent severe droughts, rising temperatures, unprecedentedly strong and prolonged El Niño-Southern Oscillation (ENSO) episodes, expansion of settlements, cultivation and human population growth in pastoral ranches adjoining the reserve. Birth rate showed strong and humped relationships with moving averages of monthly rainfall, whereas recruitment responded strongly to cumulative past rainfall. Increasing livestock incursions into the reserve depressed recruitment rate for quarter-grown topi. Expansion of pastoral settlements depressed birth rate in impala, zebra and giraffe. Frequent ENSO-related droughts caused progressive habitat desiccation and hence nutritional shortfalls for ungulates. The responses to climatic, land use and resource influences did not reflect body size, migratory or resident lifestyle, dietary guild, digestive physiology or degree of synchrony of breeding of the ungulate species.     

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.     

Transposon impala, a novel tool for gene tagging in the rice blast fungus Magnaporthe grisea

impala, a Tc1-mariner transposable element from Fusarium oxysporum, was introduced into the rice blast fungus Magnaporthe grisea to develop transposon-based insertional mutagenesis. A construct (pNIL160) containing an autonomous impala copy inserted in the promoter of niaD encoding Aspergillus nidulans nitrate reductase was introduced by transformation into a M. grisea nitrate reductase-deficient mutant. impala excision was monitored by restoration of prototrophy for nitrate. Southern analysis of niaD+ revertants revealed that impala was able to excise and reinsert at new loci in M. grisea. As observed for its host Fusarium oxysporum, impala inserted at a TA site left a typical excision footprint of 5 bp. We have shown that a defective impala copy was inactive in M. grisea, yet it can be activated by a functional impala transposase. A transformant carrying a single copy of pNIL160 was used to generate a collection of 350 revertants. Mutants either altered for their mycelial growth (Rev2) or nonpathogenic (Rev77) were obtained. Complementation of Rev77 with a 3-kb genomic fragment from a wild-type locus was successful, demonstrating the tagging of a pathogenicity gene by impala. This gene, called ORP1, is essential for penetration of host leaves by M. grisea and has no sequence homology to known genes.     

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.     

A review of the social behaviour of the giraffe Giraffa camelopardalis: a misunderstood but socially complex species

1. Until around 2000, giraffe Giraffa camelopardalis were believed to have no social structure. Despite a resurgence of interest in giraffe behaviour in around 2010, most studies are of isolated populations, making it difficult to draw general conclusions. Although it is now well established that giraffe social organisation is non-random, there is little consensus as to what influences preferred and avoided associations or the underpinning mechanisms.
2. We test two hypotheses: first, giraffe have a complex cooperative social system, exhibited by 1) stable groups of females, 2) offspring that stay in their natal group for part or all of their lives, 3) support by non-mothers in rearing young, and 4) non-reproductive females in the group; and second, giraffe form matrilineal societies, evidenced by 1) male dispersal, 2) female philopatry, 3) assistance in raising or protecting offspring, and 4) individual benefits gained from social foraging.
3. We reviewed 404 papers on giraffe behaviour and social organisation; captive studies were included where they supplemented information from free-living populations.
4. We show that giraffe exhibit many of the features typical of mammals with complex cooperative social systems and matrilineal societies. However, the social complexity hypothesis posits that such species also require complex communication systems to regulate interactions and relations among group members; giraffe communication systems are poorly understood.
5. Quantifying the fitness and survival benefits of the giraffe’s social organisation is necessary to ensure its long-term survival. Giraffe numbers have declined by 40% since 1985, they have been declared extinct in seven (possibly nine) countries and are listed as Vulnerable by the International Union for Conservation of Nature. We identify research areas that will advance our understanding of giraffe behaviour and conservation requirements.

     

Vegetation factors influencing density and distribution of wild large herbivores in a southern African savannah

Understanding factors influencing large herbivore densities and distribution in terrestrial ecosystems is a fundamental goal of ecology. This study examined environmental factors influencing the density and distribution of wild large herbivores in Gonarezhou National Park, Zimbabwe. Vegetation and surface water were predicted to have a stronger influence than anthropogenic‐related disturbances (livestock grazing, fires, settlements and poaching) on the density and distribution of wild large herbivores. Aerial survey data for seven common wild large herbivores conducted in 2007 and 2009 and environmental data were collected. Only grass cover explained a significant proportion of the variation in large herbivore densities and distribution. Moreover, only two species densities significantly differed across the Gonarezhou, namely impala and zebra. In contrast, buffalo, elephant, giraffe, kudu and nyala densities did not differ significantly across the Gonarezhou. Overall, the findings only partly support the study prediction. The study results suggest the need to further investigate the roles of environmental factors at smaller scales in order to tease out their relative strengths in influencing density and distribution of large herbivores.     

A note on foetal growth and development of the giraffe Giraffa camelopardalis giraffa

The growth of the foetus of the southern giraffe is discussed. Gestation period is calculated as 457 days being the mean of 48 observations of other workers. Birthmass is taken as 102 kg from measurements on wild giraffe. This is almost twice as great as some published figures for captive giraffe. However, subspecies also differ. Foetal growth followed a typical "J" shaped curve when plotted from the mass of 24 foetuses. The increase in vertebral column length followed a straight line. Crown/rump measurements could not always be accurately taken.     

Influence of diet transition on serum calcium and phosphorus and fatty acids in zoo giraffe (Giraffa camelopardalis)

In response to new recommendations for feeding giraffe in zoos, giraffe (n = 6) were transitioned from a typical hoofstock diet to diets containing reduced starch, protein, Ca and P and added n3 fatty acids. This diet was fed as a 50:50 mix with alfalfa and grass hay. Over the next 4 years, serum Ca, P, and fatty acids were measured every 6 months (summer and winter). Serum Ca was not affected by season (P = 0.67) or by diet (P = 0.12). Serum P was not affected season (P = 0.14), but was reduced by diet (P<0.01), and serum Ca:P was also increased by diet (P<0.01). The ratio of serum Ca:P tended to be affected by season (P = 0.07), in which animals tended to have greater Ca:P during the summer vs. the winter. The diet transition resulted in reduced serum saturated fatty acids (including lauric, myristic, palmitic, arachidic, and behenic acids), and increases in n6 fatty acids (including linolenic and arachidonic acids) and n3 fatty acids (docosahexaenoic acid) (P<0.05 for each). Overall, this diet transition resulted in blood nutrient profiles that more closely match that of values found in free-ranging giraffe.     

Seasonal diet changes in elephant and impala in mopane woodland

Elephant and impala as intermediate feeders, having a mixed diet of grass and browse, respond to seasonal fluctuations of forage quality by changing their diet composition. We tested the hypotheses that (1) the decrease in forage quality is accompanied by a change in diet from more monocots in the wet season to more dicots in the dry season and that that change is more pronounced and faster in impala than in elephant; (2) mopane (Colophospermum mopane), the most abundant dicot species, is the most important species in the elephant diet in mopane woodland, whereas impala feed relatively less on mopane due to the high condensed tannin concentration; and (3) impala on nutrient-rich soils have a diet consisting of more grass and change later to diet of more browse than impala on nutrient-poor soils. The phosphorus content and in vitro digestibility of monocots decreased and the NDF content increased significantly towards the end of the wet season, whereas in dicots no significant trend could be detected. We argue that this decreasing monocot quality caused elephant and impala to consume more dicots in the dry season. Elephant changed their diet gradually over a 16-week period from 70% to 25% monocots, whereas impala changed diets rapidly (2?C4?weeks) from 95% to 70% monocots. For both elephants and impala, there was a positive correlation between percentage of monocots and dicots in the diet and the in vitro digestibility of these forage items. Mopane was the most important dicot species in the elephant diet and its contribution to the diet increased significantly in the dry season, whereas impala selected other dicot species. On nutrient-rich gabbroic soils, impala ate significantly more monocots than impala from nutrient-poor granitic soils, which was related to the higher in vitro digestibility of the monocots on gabbroic soil. Digestibility of food items appears to be an important determinant of diet change from the wet to the dry season in impala and elephants.     

Virus associated papillomas in colobus monkeys (Colobus guereza)

Numerous papillomas ranging in size from 1--10 mm were seen in four colobus monkeys (Colobus guereza). The lesions were confined to the fingers and toes and to the dorsal-lateral aspects of the hands and feet distal to the carpus and tarsus. Electron microscopy of two of the lesions revealed compactly arranged crystalline arrays of intranuclear virus particles in the epidermis. The particles were seen only in cells of the stratum granulosum and stratum corneum. By the negative stain technique, the non-enveloped spherical particles measured 52--58 mm in diameter. The arrangement of the capsomeres was consistent with an icosahedral symmetry. The observations suggested a viral etiology for the cutaneous papillomas similar to that of other mammalian species.     

The biomass of game animals in Nairobi National Park, 1960–66

The large animals of Nairobi National Park have been censused intermittently once a month from mid-1960 until the end of 1966. Of the 23 species discussed, the density per kilometre has been determined accurately for the following: Grant's gazelle, Thomson's gazelle, water-buck, impala, wildebeest, hartebeest, eland, buffalo, warthog, giraffe, zebra and ostrich. From these figures the biomass has been calculated for each species each year. After a peak biomass of 12,775 kg/sq. km during the drought of 1960–61 the population declined to and remained fairly level at about 5690 kg/sq. km from 1962 to 1966. This figure includes about 1050 kg/sq. km of cattle and sheep. The large predators account for only about 1.4 % of the total biomass of wild animals while removing about 15.5% from this group annually.     

Reciprocal Allogrooming in Dam-reared and Hand-reared Impala Fawns

Among adult females and males of African antelope impala are unique in their performance of reciprocal allogrooming. The occurrence of this behaviour in neonatal impala fawns was explored in a free-ranging impala herd at the San Diego Wild Animal Park where 5 dam-reared fawns were observed from birth through 10 weeks of age. One-way maternal grooming and reciprocal allogrooming with the dam and non dam partners emerged as distinct behavioural systems. Maternal grooming, directed mostly to the anogenital area, was typical of that seen in other ungulates, and sharply declined over the first two weeks. Reciprocal allogrooming, characterized by alternate exchanges of grooming bouts with a partner in the same manner as in adults, was seen as early as 3–8 d after birth. All fawns were grooming with unrelated adult females by the end of the second week. By week 2 virtually every measure of reciprocal allogrooming by fawns (grooming delivered per hour, reciprocity, and percent of encounters initiated) was as high as for adults. The appearance of this reciprocal allogrooming pattern, especially at such an early age, appears to be unique among ungulates, and possibly mammals in general. Three hand-reared impala fawns, deprived of the opportunity to interact with older herdmates, but having access to impala fawns and heterospecific fawns, were observed from 1–3 mo of age. The hand-reared impala showed no alteration in the occurrence of reciprocal allogrooming behaviour compared with the dam-reared control fawns, indicating that allogrooming experience with older animals was not required for the appearance of reciprocal allogrooming at an early age. Interestingly, hand-reared fawns persisted in grooming heterospecific fawns despite the fact that heterospecifics rarely reciprocated grooming. We postulate that the strong predisposition for impala young to groom others may be related to the threat of tick infestation in the impala's ecotone habitat.     

Rothschild’s giraffe Giraffa camelopardalis rothschildi (Linnaeus, 1758) in East Africa: A review of population trends,taxonomy and conservation status

Giraffe populations have suffered a 40% decline in the past thirty years, making them a new priority for conservation and there are considerable uncertainty and disagreement over the taxonomic classification of giraffes. Consequently, there has never been a more critical time to fully understand the global population size and distribution of all giraffe subspecies. The Rothschild's giraffe Giraffa camelopardalis rothschildi (Linnaeus, 1758) is arguably one of the most imperilled giraffe subspecies. Once widespread across southern Sudan, Uganda and Kenya, the Rothschild's giraffe is now confined to a few, isolated and enclosed populations throughout Kenya and Uganda, with only one natural population remaining. Information about Rothschild's giraffe population size, distribution, and conservation history is patchy, and confined to recondite or inaccessible resources. Here, I present a review of taxonomy, distribution, conservation status and population size of the Rothschild's giraffe and present the most recent estimate of population sizes based on direct census data. Although the Latin name affixed to Rothschild's giraffe is subject to discussion, continued reference to this subspecies as G. c. rothschildi is strongly recommended.     

Gazing at a giraffe gyroscope: where are we going?

Giraffe are popular animals to watch while on wildlife safaris, and feature prominently in zoos, advertisements, toys and cartoons. Yet, until recently, few field studies have focused on giraffe. We introduce this giraffe topic issue with a review essay that explores five primary questions: How many (sub) species of giraffe exist? What are the dynamics of giraffe herds? How do giraffe communicate? What is the role of sexual selection in giraffe reproduction? How many giraffe reside in Africa? A confluence of causes has produced drastic declines in giraffe population numbers in Africa, and we conclude that guiding giraffe conservation plans depends upon evaluation of the five key quandaries that we pose.     

Immunization with viruslike particles from cottontail rabbit papillomavirus (CRPV) can protect against experimental CRPV infection.

We tested the ability of vaccination with virus-like particles (VLPs) to protect domestic rabbits against papillomas induced by the cottontail rabbit papillomavirus (CRPV). A recombinant baculovirus system that expressed only the L1 major papillomavirus structural protein or L1 plus the minor L2 protein was used in insect cells as the source of VLPs. Groups of 10 rabbits were immunized with native or denatured VLPs from CRPV or type 1 bovine papillomavirus by using Freund's adjuvant. Alum was used as the adjuvant for an additional group immunized with CRPV L1-L2 VLPs. Animals were challenged with 5 x 10(10) and 2 x 10(11) particles on opposing flanks. No protection was seen in rabbits immunized with native or denatured bovine papillomavirus L1-L2 or with denatured CRPV L1-L2. In these groups, the lower and higher challenge doses resulted in 27 of 30 animals with extensive papillomas, with each of the remaining animals having a smaller number of persistent papillomas. Progression to carcinoma developed in 20 rabbits. Animals inoculated with native CRPV VLPs composed of L1 alone or L1-L2 developed many fewer lesions; the lower and higher challenge doses resulted in 17 of 29 and 5 of 29 rabbits, respectively, with no lesions, and the remainder developed only one to eight papillomas, which all regressed except for those on 1 rabbit. None developed cancer within 1 year of infection. Rabbits vaccinated with native CRPV VLPs developed high-titer antibodies in an enzyme-linked immunosorbent assay based on native VLPs, and passive transfer of serum or immunoglobulin G from rabbits immunized with CRPV VLPs protected against CRPV challenge. We conclude that native VLPs can induce antibody-mediated, type-specific protection against experimental papillomavirus infection.     

Rainfall influences on ungulate population abundance in the Mara-Serengeti ecosystem

1. Rainfall is the prime climatic factor underpinning the dynamics of African savanna ungulates, but no study has analysed its influence on the abundance of these ungulates at monthly to multiannual time scales. 2. We report relationships between rainfall and changes in age- and sex-structured abundances of seven ungulate species monitored monthly for 15 years using vehicle ground counts in the Maasai Mara National Reserve, Kenya. 3. Abundance showed strong and curvilinear relationships with current and cumulative rainfall, with older topi, Damaliscus korrigum (Ogilby); warthog, Phacochoerus aethiopicus (Pallas); waterbuck, Kobus ellipsyprimnus (Ogilby); and impala, Aepyceros melampus (Lichtenstein) responding to longer lags than younger animals, portraying carryover effects of prior habitat conditions. 4. The abundances of newborn calves were best correlated with monthly rainfall averaged over the preceding 5-6 months for topi, waterbuck, warthog, and 2 months for the migratory zebra Equus burchelli (Gray), but with seasonal rainfall averaged over 2-5 years for giraffe, Giraffa camelopardalis (L.); impala; and kongoni, Alcelaphus busephalus (Pallas). The cumulative late wet-season rainfall was the best predictor of abundance for quarter- to full-grown animals for most species. Monthly rainfall exerted both negative and positive effects on the abundances of zebra, impala and waterbuck. Ignoring age, both sexes responded similarly to rainfall. 5. Births were strongly seasonal only for warthog and topi, but peaked between August and December for most species. Hence abundance was strongly seasonal for young topi and warthog and the migratory zebra. Pronounced seasonality in births for warthog and topi obliterated otherwise strong relationships between abundance and rainfall when both month and rainfall were included in the same model. Aggregated density produced relationships with rainfall similar to those for fully grown animals, emphasizing the necessity of demographic monitoring to reliably reveal rainfall influences on ungulate abundance in the Mara. 6. Strong relationships between abundance and rainfall suggest that rainfall underpins the dynamics of African savanna ungulates, and that changes in rainfall due to global warming may markedly alter the abundance and diversity of these mammals. Ungulates respond to rainfall fluctuations through movements, reproduction or survival, and the responses appear independent of breeding phenology and synchrony, dietary guild, or degree of water dependence. Newborns and adults have contrasting responses to rainfall. Males and females respond similarly to rainfall when age is ignored.     

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.