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Plant available moisture and plant available nutrients in soils influence forage quality and availability and subsequently affect reproductive performance in herbivores. However, the relationship of soil moisture, soil nutrients and woody forage with reproductive performance indicators is not well understood in mega-browsers yet these three are important in selecting suitable areas for conservation of mega-browsers. Here, the eastern black rhinoceros (Diceros bicornis michaeli), a mega-browser, was studied in seven geographically distinct populations in Kenya to understand the relationships between its reproductive performance indicators and plant available moisture, plant available nutrients and woody cover. Reproductive parameters showed a complex relationship with plant available moisture and plant available nutrients. We found an increase in the predicted yearly percentage of females calving as plant available nutrients decreased in areas of high levels of plant available moisture but no relationship with plant available nutrients in areas of low plant available moisture. Age at first calving was earlier, inter-calving interval was longer and yearly percentage of females calving was higher at higher woody cover. Woody plant cover contributes positively to black rhino reproduction performance indicators, whereas plant available moisture and plant available nutrients add to the selection of conservation areas, in more subtle ways.

The prevention of tick-borne diseases is a major challenge for livestock production globally. Tick control strategies include the use of acaricides, but the prescribed strategies do not achieve the desired results in several countries, including Kenya. To better understand how tick treatment practices, contribute to reported tick treatment failures, we assessed livestock owners' acaricide procurement, level of knowledge about acaricides and tick resistance, and how they apply acaricides. We also assessed the quality of the commonly available acaricides. We focused on three livestock systems in Laikipia County, Kenya: two private ranches; one community ranch whose members communally graze their cattle and acquire and apply acaricides; and individual livestock owners in two pastoral communities who individually graze their cattle and acquire and apply acaricides. Through interviews and focus group discussions we assessed; access to acaricides, livestock owners' knowledge, and acaricide use practices; interview data were triangulated with participant observations (n��=��107). We analysed nine commonly used acaricides to determine the active ingredient concentration and we determined the concentration of active ingredients in acaricide dilutions collected on farms. All livestock owners had access to and used chemical acaricides for tick control, predominantly amitraz-based. Private ranchers bought one amitraz-based acaricide in bulk directly from the manufacturer, while all other livestock owners bought from agrovet shops. The livestock owners acquired knowledge about acaricides from their own experiences and through experience-based recommendations from peers, but not from the technical information provided by the manufacturers and agrovet shops. All pastoral livestock frequently changed acaricide brand and active ingredient class. A large majority of pastoralists (86%) mixed acaricide brands within and across active ingredient classes; a smaller majority (56%) mixed acaricides with crop pesticides and insecticides. Our lab tests confirmed the content description on the labels bought from agrovet shops. However, on-farm acaricide dilutions from all three livestock systems deviated from the level recommended for effective treatment. If too diluted, the acaricide does not kill ticks, promoting resistance development. If too concentrated, this increases environmental contamination and raises public health concerns. Livestock owners lack a technical understanding of the functioning of acaricides, compromising their use and effectiveness. The widely adopted mixing of acaricides with insecticides and pesticides raises serious health concerns.

Wild vertebrate populations all over the globe are in decline, with poaching being the second-most-important cause. The high poaching rate of rhinoceros may drive these species into extinction within the coming decades. Some stakeholders argue to lift the ban on international rhino horn trade to potentially benefit rhino conservation, as current interventions appear to be insufficient. We reviewed scientific and grey literature to scrutinize the validity of reasoning behind the potential benefit of legal horn trade for wild rhino populations. We identified four mechanisms through which legal trade would impact wild rhino populations, of which only the increased revenue for rhino farmers could potentially benefit rhino conservation. Conversely, the global demand for rhino horn is likely to increase to a level that cannot be met solely by legal supply. Moreover, corruption is omnipresent in countries along the trade routes, which has the potential to negatively affect rhino conservation. Finally, programmes aimed at reducing rhino horn demand will be counteracted through trade legalization by removing the stigma on consuming rhino horn. Combining these insights and comparing them with criteria for sustainable wildlife farming, we conclude that legalizing rhino horn trade will likely negatively impact the remaining wild rhino populations. To preserve rhino species, we suggest to prioritize reducing corruption within rhino horn trade, increasing the rhino population within well-protected 'safe havens' and implementing educational programmes and law enforcement targeted at rhino horn consumers. peerReviewed

Weather conditions can impact infectious disease transmission, causing mortalities in humans, wild and domestic animals. Although rainfall in dry tropical regions is highly variable over the year, rainfall is thought to play an important role in the transmission of tick-borne diseases. Whether variation in rainfall affects disease-induced mortalities, is, however, poorly understood. Here, we use long-term data on monthly rainfall and Boran cattle mortality (1998–2017) to investigate associations between within-year variation in rainfall and cattle mortalities due to East Coast fever (ECF), anaplasmosis and babesiosis in Laikipia, Kenya, using ARIMAX modelling. Results show a negative correlation between monthly rainfall and cattle mortality for ECF and anaplasmosis, with a lag effect of 2 and 6 months, respectively. There was no association between babesiosis-induced mortalities and monthly rainfall. The results of this study suggest that control of the tick-borne diseases ECF and anaplasmosis to reduce mortalities should be intensified during rainy periods after the respective estimated time lags following dry periods.

SUMMARYCurrent theories on diversity–disease relationships describe host species diversity and species identity as important factors influencing disease risk, either diluting or amplifying disease prevalence in a community. Whereas the simple term ‘diversity’ embodies a set of animal community characteristics, it is not clear how different measures of species diversity are correlated with disease risk. We therefore tested the effects of species richness, Pielou's evenness and Shannon's diversity on bovine tuberculosis (bTB) risk in cattle in the Afar Region and Awash National Park between November 2013 and April 2015. We also analysed the identity effect of a particular species and the effect of host habitat use overlap on bTB risk. We used the comparative intradermal tuberculin test to assess the number of bTB-infected cattle. Our results suggested a dilution effect through species evenness. We found that the identity effect of greater kudu – a maintenance host – confounded the dilution effect of species diversity on bTB risk. bTB infection was positively correlated with habitat use overlap between greater kudu and cattle. Different diversity indices have to be considered together for assessing diversity–disease relationships, for understanding the underlying causal mechanisms. We posit that unpacking diversity metrics is also relevant for formulating disease control strategies to manage cattle in ecosystems characterized by seasonally limited resources and intense wildlife–livestock interactions.

Ixodid ticks (Acari: Ixodidae) are important disease vectors for large carnivores, but the composition of the tick communities that parasitize carnivores is poorly understood. We collected ticks from leopards (Panthera pardus) and brown hyenas (Hyaena brunnea) in the Soutpansberg Mountains, South Africa, to determine which species feed on these carnivores. We identified a total of eight tick species belonging to six genera, and recorded Ixodes neitzi and Hyalomma rufipes on P. pardus for the first time.

Wildlife detection dogs are required to correctly discriminate target wildlife species odor from nontarget species odors (specificity), while enabling some degree of target odor variation (generality). Because there is no standardized training protocol, and little knowledge on training efficiency, we conducted a case study to test a dog’s training efficiency in detecting 2-week-old wild otter (Lutra lutra) feces (spraint) odor among feces odors from 6 other large mammal species that often share the otter’s natural habitat, including fox (Vulpes vulpes), hare (Lepus europaeus), roe deer (Capreolus capreolus), and cattle (Bos taurus). The dog was trained using a standard multiple-choice carrousel in a stepwise protocol. We started with odor samples from fresh captive otter spraints and progressed toward 2-week-old spraints from wild otters among other mammalian dung odors and tested for specificity and generality after each training step. We show that training on only 2 variations of spraints from captive otters enabled the dog to detect all desired spraint odor variations in our protocol, indicating a rapid generalization to variations of spraint odor the dog was not trained on, while retaining specificity. Testing such concept formation of target odors should be included in detection dog training and certification and could serve as a quality control measure of detection dog performance.

The scale of resource heterogeneity may influence how resources are locally partitioned between co-existing large and small organisms such as trees and grasses in savannas. Scale-related plant responses may, in turn, influence herbivore use of the vegetation. To examine these scale-dependent bi-trophic interactions, we varied fertilizer [(nitrogen (N)/phosphorus (P)/potassium (K)] applications to patches to create different scales of nutrient patchiness (patch size 2 × 2 m, 10 × 10 m, or whole-plot 50 × 50 m) in a large field experiment in intact African savanna. Within-patch fertilizer concentration and the total fertilizer load per plot were independently varied. We found that fertilization increased the leaf N and P concentrations of trees and grasses, resulting in elevated utilization by browsers and grazers. Herbivory off-take was particularly considerable at higher nutrient concentrations. Scale-dependent effects were weak. The net effect of fertilization and herbivory was that plants in fertilized areas tended to grow less and develop smaller rather than larger standing biomass compared to plants growing in areas that remained unfertilized. When all of these effects were considered together at the community (plot) level, herbivory completely eliminated the positive effects of fertilization on the plant community. While this was true for all scales of fertilization, grasses tended to profit more from coarse-grained fertilization and trees from fine-grained fertilization. We conclude that in herbivore-dominated communities, such as the African savanna, nutrient patchiness results in the herbivore community profiting rather more than the plant community, irrespective of the scale of patchiness. At the community level, the allometric scaling theory’s prediction of plant—and probably also animal—production does not hold or may even be reversed as a result of complex bi-trophic interactions. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3627-0) contains supplementary material, which is available to authorized users.

The microbiota of the mammalian gut is a complex ecosystem, the composition of which is greatly influenced by host genetics and environmental factors. In this study, we aim to investigate the influence of occupancy (a geographical area of habitation), species, age and sex on intestinal microbiota composition of the three lemur species: Eulemur fulvus, E. rubriventer and E. rufifrons. Faecal samples were collected from a total of 138 wild lemurs across Madagascar, and microbial composition was determined using next-generation sequencing of PCR-amplified 16S rRNA gene fragments. Consistent with reports from other primate species, the predominant phyla were Firmicutes (43 ± 6.4% [s.d.]) and Bacteroidetes (30.3 ± 5.3%). The microbial composition was strongly associated with occupancy in the E. fulvus population, with up to 19.9% of the total variation in microbial composition being explained by this factor. In turn, geographical differences observed in faecal microbiota of sympatric lemur species were less pronounced, as was the impact of the factors sex and age. Our findings showed that among the studied factors occupancy had the strongest influence on intestinal microbiota of congeneric lemur species. This suggests adaptation of microbiota to differences in forest composition, climate variations and correspondingly available diet in different geographical locations of Madagascar.

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. 2017 John Wiley & Sons Ltd.

The persistence of tropical rainforests, together with their flora and fauna, is highly threatened by anthropogenic disturbances. In this study, we investigate to what extent selective logging influences the structure and composition of a tropical rainforest in Madagascar and subsequently lemur encounter rates and cluster sizes. We quantified forest structure variables and conducted transect surveys of seven sympatric diurnal lemur species in five protected forest sites with different logging histories. We found that DBH, tree height, the interquartile ranges of DBH and tree height (measure of forest heterogeneity), tree species and family richness were relatively high and tree density was relatively low in less disturbed compared to disturbed sites. Although the disturbed forests have not fully recovered to previous conditions, they seem to have recovered from a functional perspective into suitable lemur habitat, as lemur encounter rates and cluster sizes were similar in disturbed and less disturbed sites. We only found slightly higher encounter rates for Varecia variegata (P = 0.078) and lower encounter rates for Eulemur rufifrons (P = 0.059) in less disturbed forests. This is one of the first studies that report the presence of V. variegata, a species characterised by its drastic decline, in previously logged sites. Lemurs travelling between disturbed and less disturbed sites disperse seeds and hereby facilitate forest regeneration. Therefore, we promote the need for better attention to the value of logged forests for biodiversity conservation in Madagascar and suggest that there is considerable potential for regenerating logged forests to support lemur communities.

Prevention of tick borne diseases is often through tick control practices. This article diagnoses tick control practices, knowledge underlying these practices and how knowledge is shared at the wildlife-livestock interface in Laikipia County, Kenya. It identifies diverse land use and tick control practices by different land and livestock owners from a scientific knowledge-based (techne) and context driven experiential knowledge-based (metis) perspective. Interviews, focus group discussions, observations and documents yielded qualitative data to unravel i) the historical development of tick control in Kenya ii) techne and metis tick control practices within three ranches and among pastoralists in Laikipia County, and iii) status of tick knowledge sharing between stakeholders. Historical tick control measures date back to about 100 years ago, with increasingly strong veterinary measures over the decades under government control. However, the veterinary control system collapsed around 1991 and livestock owners took tick control into their own hands. All respondents indicated having relevant techne available about tick ecology and tick management practices. To adapt to the changing social, economic and institutional context, they further developed metis, integrating the known techne. Metis and techne complemented each other. Our study reveals that metis is developed within stakeholder groups. The data also suggest that metis practices sometimes develop risky effects to animal, human and environmental health. Knowledge on tick control is mainly shared within a social group, not between groups. We esteem, knowledge sharing between different stakeholder groups (ranchers, pastoralists, DVS) may provide opportunities for better informed decision-making based on fruitful combinations of techne and metis for effective and safe tick management.

Water is a scarce resource in semi-arid savannas where over half of the African elephants (Loxodonta africana) populations occur and may therefore influence their movement pattern. A random search is expected for an animal with no information on the location of the target resource, else, a direction-oriented walk is expected. We hypothesized that elephants movement patterns show a stronger directional orientation towards water sources in the dry season compared to the wet season. We investigated the movement paths of four male and four female elephants with hourly GPS fixes in Tsavo National Park, Kenya in 2012-2013. Consistent with our predictions, the movement paths of elephants had longer step lengths, longer squared net displacements, and were directed towards water sources in the dry season as compared to the wet season. We argue that African elephants know the location of dispersed water resources, enabling them to survive with scarce resources in dry savannas. These results can be used in conservation and management of wildlife, through for instance, protection of preferred water sources.

Tannin concentrations fluctuate spatially and temporally within and among plant species, with consequences for forage quality of herbivores. The extent to which these fluctuations influence foraging activities of goats is not fully understood. While accounting for the effects of the time of the day and season, we tested the hypothesis that goats exposed to high levels of condensed tannins (i) spend less of their foraging time browsing, (ii) spend more time grazing, and (iii) reduce their total foraging time, especially during the dry season when grasses dry out and deciduous trees lose leaves. We orally dosed 15 goats with (i) 20 g of condensed tannins extract dissolved in 50 ml of water (high tannin exposure), another 15 goats (ii) with 20 g of polyethylene glycol dissolved in 50 ml of water, which neutralizes the effects of tannins (low tannin exposure), and the last group of 15 goats (iii) with 50 ml of water (control). We recorded the time spent on grazing, browsing and these two activities together (i.e., foraging) for 30 days in the dry and wet season. As expected, dosing goats with condensed tannins reduced their browsing time and increased the time spent grazing. Goats dosed with polyethylene glycol increased their browsing time and lowered their time spent grazing. Animals dosed with polyethylene glycol foraged for longer than other treatment groups in the dry season, whereas the goats dosed with condensed tannins increased their foraging time in the wet season. Overall, all treatment groups spent a similar amount of time foraging, indicating an instinctive drive by goats to maintain high total foraging time while avoiding over ingestion of tannin-rich forages. We concluded that tannins do not supress total foraging time for free-ranging goats. Instead, they increase the amount of time animals spend on grazing on herbaceous plants and decrease the amount of time animals spend browsing on woody plants. Comparing time spent on different foraging activities by goats is a first step towards understanding how goats utilize feed resources and how tannins influence resource utilization patterns.

Savanna vegetation is characterized by tree–grass co-existence that can experience intense water limitation, yet the water relations of these savanna plants are poorly understood. We examined the water sources for trees and grasses in different seasons and investigated the importance of hydraulic redistribution in three tree species inhabiting a semi-arid savanna in South Africa. We used natural variation in H and O stable isotope composition of source waters to identify the principal water sources for these plants. We conducted an experiment by labelling deep-soil (2.5-m depth) with a deuterium tracer. Seasonal differences in the stable isotope composition of water in trees and grasses indicated that there was water-source use partitioning as well as overlap. Trees and grasses used water from the topsoil after rainfall indicating overlap of water-source use. All tree species shifted to groundwater or subsoil water use when there was no water in the topsoil indicating partitioning of water use. Grasses always used water from the topsoil. The seasonal changes in water-source use by trees and grasses indicated possible shifts in tree–grass interactions during different periods of the year. The tracer experiment confirmed hydraulic redistribution in all the three tree species and water transfer to grasses via the topsoil. However, this occurred only in the dry season. Our observations and experimental results indicate the potential for facilitation effects by trees to their understory grasses and show that dry season hydraulic redistribution from trees to grasses could be an important facilitative mechanism maintaining tree–grass co-existence in savannas.

Species that inhabit marine and intertidal ecosystems face osmoregulatory challenges, risking dehydration and increased ion concentrations in the body. Lizards need to either tolerate or regulate these increased ion concentrations. In this study, we aim to understand how Suter's skink (Oligosoma suteri), an intertidal skink restricted to shoreline habitats, is able to cope with the physiological stress of living in an extreme salt environment. We determined the diet, prey species' salt content, and nasal and cloacal salt excretion on Rangitoto and Motutapu Islands in northern New Zealand, where the skinks have contrasting access to terrestrial invertebrates. Analysis of stable isotopes suggests inter- and intra-population variability in Suter's skink diets. Intertidal invertebrates under washed up seaweed appear to compose a major part of the diet of the Rangitoto population, while the Motutapu population showed evidence for a mixed diet of terrestrial and intertidal invertebrates. Sodium content of prey species decreased with an increasing distance from the seawater. Field secretion of cations through nasal glands consisted primarily of Na(+), which is consistent with other marine and intertidal species. Sodium was also the primary cation in urine. In contrast, fecal cations consisted primarily of K(+). This first study into the salt secretion of an intertidal skink species provides evidence of Suter's skink's plasticity in secreting excess ions through nasal salt glands. This likely enables it to deal with the challenges of living in a semi-marine habitat.

Infectious diseases transmission is strongly determined by who contacts whom. Bovine tuberculosis (bTB) caused by Mycobacterium bovis is a worldwide burden for animal populations. One of the major transmission mechanism between herd is the transfer of infectious animal. In East Africa, pastoralists may receive or bestow livestock to create and strengthen social relationships. Here, we used a network approach to examine the relative importance of such cattle transfer in the transmission of bTB. First, a total of 2550 cattle from 102 herds were tested using the comparative intradermal tuberculin test to assess the presence of bTB infected cattle in the herd. A herd was considered bTB positive if it had at least one tuberculin reactor animal. Next, we calculated the centrality of each herd in the cattle transfer network using four established measures of social network centralization: degree, betweenness, closeness and fragmentation. The relationships between the network centrality measures and bTB infection were examined using generalized linear mixed models (GLMM). We found that a herd's in-degree in the social network was positively correlated with the risk of being infected with bTB (b=4.2, 95%CI=2.1-5.7; p

Myths and metaphors that occur in media frames play an important role in influencing public perceptions of an issue in times of war, political conflict, crisis and disaster. This, in turn, influences policy makers and (inter)national assistance and aid programmes. We investigated whether a metaphoric spill-over of frames used in connection with political events could explain the misrepresentation in the framing of wildlife conservation. Zimbabwe experienced a severe political conflict and economic downturn in 2000 when land reforms took place. We analyzed newspaper articles on Zimbabwe's wildlife conservation published between 1989 and 2010 from newspapers in Zimbabwe, the United Kingdom and the United States of America. We selected three issues about wildlife conservation in Zimbabwe in the local and international media, namely, the ivory ban, rhino protection, and Communal Areas Management Programme for Indigenous Resources to investigate the spill-over effect. Our results show that in the 1990s, the majority of newspaper articles highlighted that wildlife conservation in Zimbabwe was largely successful. However, two major changes occurred after 2000 following the land reforms in Zimbabwe. First, the international media showed little interest in wildlife conservation in Zimbabwe as evidenced by a sharp decline in published articles and second, the frames changed in the international media with the “political unrest and land reform” blame frame becoming more dominant. This transition in reporting, frames, and low frame parity shows that there was a spill-over effect of political frames into wildlife conservation following Zimbabwe's land reforms in 2000. Metaphoric spill-over effects may thus create myths in the readership, in turn influencing policy-derived actions in a sector that is not or poorly related to the actual disaster. Keywords Framing; Land reform; Metaphors; Nature conservation

Illegal hunting of wildlife, or top-down harvesting, is a major issue in today's society, particularly in tropical ecosystems. There has been widespread concern about increasing illegal hunting of wildlife in most conservation areas in Zimbabwe following the political instability and economic decline the country faced since 2000. In this study, we focused on the northern Gonarezhou National Park (GNP), a large and unfenced protected area, and adjacent communal areas in southern Zimbabwe. We hypothesised that illegal hunting activities would (1) be perceived to have increased due to economic collapse and (2) vary with law enforcement efforts. A total of 236 local residents from eight villages adjacent to the northern GNP were interviewed using semi-structured questionnaires from December 2010 to May 2011, and law enforcement data for northern GNP between 2000 and 2010 were retrieved from the park law enforcement database. A total of 26 animal species were reportedly hunted. Bushmeat consumption and the need for local trade to raise income were reported as the main reasons behind illegal hunting. Contrary to the first hypothesis, the majority of respondents ( n = 156, 66%) reported that illegal hunting activities had declined between 2000 and 2010 largely due to increased park protection as also supported by law enforcement data. A total of 22 animal species were recorded as having been illegally hunted in northern GNP. The number of illegal hunters arrested declined with increased law enforcement efforts although the number of wire snares recovered and hunting dogs shot appeared to increase following increased law enforcement efforts. These results partly support the second hypothesis that illegal hunting activities would vary with law enforcement efforts.

Many small Parks inKenyaare being fenced to control human-wildlife conflict. Some of these Parks have a diversity of large herbivores which might increase in density in the wake of fencing and subsequent compression of their home ranges due to closure of their migratory routes. It is important to understand the consequences of such an increase on the structuring of insularised herbivore assemblages in such Parks. We studied seasonal resource segregation and niche breadth variation as mechanisms of coexistence in a high density grazer assemblage inLakeNakuruNational Parkwhich is small and completely fenced. Diet composition and habitat use were considered as variables of resource use. We predicted that overlap in resource use and niche breadth would be the smallest among grazers with similar body weights in the dry season which is the most resource limiting for grazers in East Africa. Our results were contrary to the predictions because of lack of seasonal differentiation in the overlap of diet composition and habitat use, and in niche breadth. Overlaps in resource use were consistently high during both the wet and dry seasons, and niche breadth contraction during the dry season was not possible probably because of lack of species-specific niches during the dry season. Our results suggest that there might be competitive interactions in this grazer assemblage which is an important parameter to consider in the management of the Park.

Annual rural incomes in Southern Africa show large rainfall-induced fluctuations. Variable rainfall has serious implications for agro-pastoral activities (crop cultivation and livestock keeping), whereas wildlife and tourism are less affected. The aim of this paper is to investigate the role of wildlife income in reducing rainfall-induced fluctuations in households’ annual incomes. We analyse costs and benefits from agro-pastoral systems in southeastern Zimbabwe by means of a two-tier longitudinal survey and wildlife benefits through analysis of wildlife revenues. We use the portfolio theory framework to investigate whether wildlife conservation has the potential for farmers to reduce risk associated with agricultural production. Results show that even though wildlife income is small, it tends to be less volatile than income from the agro-pastoral system. Furthermore, the addition of wildlife as an asset to the rural farmers’ portfolio of assets showed that wildlife can be used as a hedge asset to offset risk from agricultural production without compromising on return. The potential of diversification using wildlife is, however, limited since agriculture and wildlife assets are positively correlated. We conclude that revenues from wildlife have some potential to reduce annual household income fluctuations, but only to a limited extent. We argue that if wildlife is organised on a more commercial basis, a more substantial role can be played by wildlife income in reducing variations in rural households’ incomes.

Bovine tuberculosis (bTB) infection is generally correlated with individual cattle's age, sex, body condition, and with husbandry practices such as herd composition, cattle movement, herd size, production system and proximity to wildlife - including bTB maintenance hosts. We tested the correlation between those factors and the prevalence of bTB, which is endemic in Ethiopia's highland cattle, in the Afar Region and Awash National Park between November 2013 and April 2015. A total of 2550 cattle from 102 herds were tested for bTB presence using the comparative intradermal tuberculin test (CITT). Data on herd structure, herd movement, management and production system, livestock transfer, and contact with wildlife were collected using semi-structured interviews with cattle herders and herd owners. The individual overall prevalence of cattle bTB was 5.5%, with a herd prevalence of 46%. Generalized Linear Mixed Models with a random herd-effect were used to analyse risk factors of cattle reactors within each herd. The older the age of the cattle and the lower the body condition the higher the chance of a positive bTB test result, but sex, lactation status and reproductive status were not correlated with bTB status. At herd level, General Linear Models showed that pastoral production systems with transhumant herds had a higher bTB prevalence than sedentary herds. A model averaging analysis identified herd size, contact with wildlife, and the interaction of herd size and contact with wildlife as significant risk factors for bTB prevalence in cattle. A subsequent Structural Equation Model showed that the probability of contact with wildlife was influenced by herd size, through herd movement. Larger herds moved more and grazed in larger areas, hence the probability of grazing in an area with wildlife and contact with either infected cattle or infected wildlife hosts increased, enhancing the chances for bTB infection. Therefore, future bTB control strategies in cattle in pastoral areas should consider herd size and movement as important risk factors.

Very little is known about marking behavior of the endangered Andean bear (Tremarctos ornatus). Here, we present a first detailed description of Andean bear marking behavior obtained using camera traps. From November 2012 to April 2013, we inspected 16 bear trails in the Napo province of eastern Ecuador, and installed camera traps (n = 3) at marking sites to document their marking behavior. We obtained 22 video recordings of Andean bears, all of which were captured during daytime. Almost all recordings (n = 18) contained behavior associated with marking. Tree-rubbing was the main behavioral display at marking sites, and consisted of 4 common activities: (1) tree-sniffing, (2) rubbing the neck and/or shoulders, (3) rubbing the flanks, and (4) rubbing the back. Bears also claw-marked and urinated while rubbing trees. We only observed scent-marking from males. Consistent with other bear species, we suggest that Andean bears communicate intra-specifically through their marking behavior.

Savannas support mixed tree-grass communities and interactions between these are typically viewed as being competitive based on studies that focused on grass aboveground production. However, an important plant response to competition and resource limitation is an increase in root reserves. We investigated root characteristics of perennial grasses in the presence and absence of trees as a proxy of competition in South African savannas in three sites that differed in rainfall. We based our study on the hypothesis that competition from trees and water limitation will result in increased storage in roots of grasses under trees. Results indicate no significant effect of variation in rainfall of the different study locations on root characteristics of grasses. Furthermore, trees did not significantly influence most grass root characteristics that we measured. The only exception was nitrogen-content that showed an increase with rainfall and tree presence through potentially higher mineralization rates and nitrogen availability in the under-tree canopy environment. As the study sites are in the drier rainfall range in South Africa, it is likely that trees and grasses in these dry savannas may have a positive relationship conforming to the stress-gradient hypothesis. Alternatively, grasses and trees may be using complementary water and nutritional resources.

Understanding a species’ foraging habits and preferences is fundamental to understanding its overall ecology and essential for its management and conservation. In general, large herbivores are classified as either grazers, browsers , or mixed feeders , and a species’ diet preference is related to its body mass and digestive trait syndrome. Here, we analyze feeding strategies of large herbivores in South and Southeast Asia (SSEA) as related to their body mass and digestive trait syndromes. Overall, our results are similar to patterns observed on other continents. The majority of large herbivore species in SSEA are mixed feeders. Browsers and frugivores dominate the smallest body mass classes, while bulk feeders , predominantly grazers, dominate the largest body mass classes. There is an absence of hindgut fermenters in the lower body mass classes, and an absence of ruminants in the megaherbivore class (>1000 kg). Cervids in SSEA do not get as large as Bovids, and in both Cervids and Bovids the greatest number of species are found in the smaller body mass classes. Although large herbivores in SSEA occur across a wide range of different habitat types, there are discernible habitat associations with different groups of species. While this chapter sheds light on this important facet of large herbivore ecology in the region, there remains an acute lack of data on the foraging ecology of the majority of species in SSEA.

Elephant inhabiting arid and semi-arid savannas often experience periods of drought, which, if prolonged, may cause mortality. During dry periods, elephant aggregate around water sources and deplete local forage availability. However, the relationships between adult elephant mortality and both high local elephant density and forage availability close to water during dry periods remain unexplored. We hypothesized that elephant mortality is higher: a) when dry periods are longer, b) closer to water points, and c) in areas with higher local elephant density. Using nine years of elephant carcass data from Tsavo Conservation Area in Kenya, we analysed the probability of adult elephant mortality using maximum entropy modelling (MaxEnt). We found that elephant carcasses were aggregated and elephant mortality was negatively correlated with four months cumulative precipitation prior to death (which contributed 41% to the model), Normalised Difference Vegetation Index (NDVI) (19%) and distance to water (6%), while local elephant density (19%) showed a positive correlation. Three seasons (long dry, short dry and short wet seasons) showed high probability of elephant mortality, whereas low probability was found during long wet seasons. Our results strongly suggest that elephants starve to death in prolonged drought. Artificial water holes may lead to lower mortality, but also to larger populations with subsequent high browsing pressure on the vegetation. Our results suggest that elephant populations in arid and semi-arid savannas appear to be regulated by drought-induced mortalities, which may be the best way of controlling elephant numbers without having to cull.

Large herbivores are purported to continue consuming toxin-containing forages as long as their capacity to neutralize, detoxify and excrete dietary toxins is not exceeded. This capacity depends on the availability of liver enzymes, energy and amino acid precursors. While this may explain increased intake of toxin-rich forages by herbivores supplemented with nutrients, a different effect may emerge in rangelands dominated by forages that are rich in condensed tannins which are not as degradable and readily absorbable as toxins. In a field experiment, we investigated the effects of supplementing animals with a high-energy source (yellow maize grain) and a high-protein source (soybean meal) on browse intake, foraging behaviour and diet composition of goats in a semi-arid savanna. In line with our prediction, nutrient supplementation increased the percentage time spent by goats on browsing and subsequently enhanced browse intake. Supplemented goats consumed more condensed tannins than goats that were not supplemented. Goats from supplemented groups tended to compose different diets from goats that received no supplement. Supplementation may have modified animals’ requirements for nutrients in ways that influenced feeding behaviour. We contend that supplements likely replaced the nutrients that are routinely bound and rendered indigestible by condensed tannins and thus increased the intake of tannin-rich forages through delaying a negative post-ingestive feedback (aversion) from dietary tannins. In conclusion, we showed that nutrient supplementation can be used to increase defoliation of tannin-rich shrubs by mixed feeders which suggests a potential for browsers and mixed-feeders to serve as biological bush control agents. Furthermore, these results have serious implication for herbivores and the environment.

1. Scaling theory predicts that organisms respond to different scales of resource patchiness in relation to their own size. We tested the hypothesis that the scale of nutrient patchiness mediates resource partitioning between large trees and small grasses in a semi-arid savanna. 2. In a factorial field experiment, Colophospermum mopane trees and associated grasses were fertilized at either a fine or coarse scale of patchiness with nitrogen (N), phosphorus (P) or N + P. The growth of marked tree shoots, herbaceous biomass and leafNand P concentrations were monitored for 2 years following fertilization. 3. Responses of trees were partly scale dependent. Tree leaf N concentration and shoot length relatively increased with fertilization at a coarse scale. Tree leaf mass decreased when P was supplied at a fine scale of patchiness, suggesting intensified grass competition. 4. Phosphorus fertilization increased leaf P concentrations more in grasses than trees, whereas N fertilization increased leaf N concentration moderately in both trees and grasses. Herbaceous above-ground biomass around focal trees was negatively correlated with tree size when fertilized with N, suggesting intensified tree competition. 5. Synthesis. Our results support the hypothesis that trees benefit more from nutrients supplied at a relatively coarse scale of patchiness. No direct responses of grasses to scale were detected. In trees, the scale effect was surpassed by the effect of sample year, when rainfall varied

There is a growing concern that the feeding habits of the African elephant, which include pushing over, uprooting and snapping trees, may have a negative impact on other herbivores. Browsed trees are known to respond by either increasing production (shoots and leaves) or defence (secondary compounds). It is not clear, however, what proportion of the browsed biomass can be made available at lower feeding heights after a tree is pushed over or snapped; thus, it is also unclear how the forage quality is affected. In a field survey in Kruger National Park, South Africa, 708 Mopane trees were measured over four elephant utilization categories: snapped trees, pushed-over trees, uprooted trees and control trees. The elephants’ impact on the leaf biomass distribution was quantified, and the forage quality (Ca, P, K and Mg, N, digestibility and condensed tannin [CT] concentrations) were analyzed. Pushed-over and uprooted trees had the maximum leaf biomass at lower height s( o1m), snapped trees at medium heights (1‐2m) and control trees at higher heights (42m). In all three utilization categories, the minimum leaf biomass was seven times higher than it was for control trees at a height of below 1m. Leaf nitrogen content increased in all three categories and was significantly higher in snapped trees. CT concentrations increased slightly in all trees that were utilized by elephants, especially on granitic soils in the dry season. The results provide the insight that elephants facilitate the redistribution and availability of browse and improve the quality, which may positively affect small browsing herbivores.

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.

The goal of this study was to test whether body-mass based foraging principles, guided by plant available moisture (PAM) and plant available nutrients (PAN), could explain large mammalian herbivore species distribution and richness in India. We tested (1) whether the occurrence of larger-bodied herbivore species increases with PAM, but is independent of PAN, (2) whether the occurrence of smaller-bodied herbivore species decreases with PAM, but increases with PAN, and (3) whether herbivore species richness is highest in areas with intermediate PAM and high PAN. We analyzed the distribution and richness of the 16 large (>10 kg) herbivore species found in sub-Himalayan mainland India. Since the distributions of large herbivores in India have been altered by historic human activity, we only used India's largest 76 protected areas as data points, with respect to PAM (log10(rainfall/potential evapotranspiration)), PAN (soil cation exchange capacity), elevation, tree cover, and fire frequency. Using regression and null models to analyze the data, we found positive relations between PAM and the occurrences of the larger-bodied species (elephant and gaur), and negative relations between PAM and the occurrences of smaller-bodied species (chinkara, four-horned antelope and blackbuck). We also found positive relations between the occurrence of the smaller-bodied species and PAN. Large herbivore species richness in India is highest in Kanha and Indravati, areas with high PAN and intermediate PAM. We found that elevation, tree cover and fire frequency were insignificant predictors of herbivore species richness, although elevation and tree cover explained the distribution of a few species. Based on our null model analyses results, we conclude that moisture and soil nutrients are important in determining large herbivore species distribution and richness in sub-Himalayan India.

Information about the distribution of grass foliar nitrogen (N) and phosphorus (P) is important for understanding rangeland vitality and for facilitating the effective management of wildlife and livestock. Water absorption effects in the near-infrared (NIR) and shortwave-infrared (SWIR) regions pose a challenge for nutrient estimation using remote sensing. The aim of this study was to test the utility of water-removed (WR) spectra in combination with partial least-squares regression (PLSR) and stepwise multiple linear regression (SMLR) to estimate foliar N and P, compared to spectral transformation techniques such as first derivative, continuum removal and log-transformed (Log( 1 / R )) spectra. The study was based on a greenhouse experiment with a savanna grass species ( Digitaria eriantha ). Spectral measurements were made using a spectrometer. The D. eriantha was cut, dried and chemically analyzed for foliar N and P concentrations. WR spectra were determined by calculating the residual from the modelled leaf water spectra using a nonlinear spectral matching technique and observed leaf spectra. Results indicated that the WR spectra yielded a higher N retrieval accuracy than a traditional first derivative transformation ( R 2 = 0.84 , RMSE = 0.28) compared to R 2 = 0.59 , RMSE = 0.45 for PLSR. Similar trends were observed for SMLR. The highest P retrieval accuracy was derived from WR spectra using SMLR ( R 2 = 0.64 , RMSE = 0.067), while the traditional first derivative and continuum removal resulted in lower accuracy. Only when using PLSR did the first derivative result in a higher P retrieval accuracy ( R 2 = 0.47 , RMSE = 0.07) than the WR spectra ( R 2 = 0.43 , RMSE = 0.070). It was concluded that the water removal technique is a promising technique to minimize the perturbing effect of foliar water content when estimating grass nutrient concentrations.

1. Understanding and accurately predicting the spatial patterns of habitat use by organisms is important for ecological research, biodiversity conservation and ecosystem management. However, this understanding is complicated by the effects of spatial scale, because the scale of analysis affects the quantification of species-environment relationships. 2. We therefore assessed the influence of environmental context (i.e. the characteristics of the landscape surrounding a site), varied over a large range of scales (i.e. ambit radii around focal sites), on the analysis and prediction of habitat selection by African elephants in Kruger National Park, South Africa. 3. We focused on the spatial scaling of the elephants' response to their main resources, forage and water, and found that the quantification of habitat selection strongly depended on the scales at which environmental context was considered. Moreover, the inclusion of environmental context at characteristic scales (i.e. those at which habitat selectivity was maximized) increased the predictive capacity of habitat suitability models. 4. The elephants responded to their environment in a scale-dependent and perhaps hierarchical manner, with forage characteristics driving habitat selection at coarse spatial scales, and surface water at fine spatial scales. 5. Furthermore, the elephants exhibited sexual habitat segregation, mainly in relation to vegetation characteristics. Male elephants preferred areas with high tree cover and low herbaceous biomass, whereas this pattern was reversed for female elephants. 6. We show that the spatial distribution of elephants can be better understood and predicted when scale-dependent species-environment relationships are explicitly considered. This demonstrates the importance of considering the influence of spatial scale on the analysis of spatial patterning in ecological phenomena.

In this first detailed analysis of gaur Bos gaurus life-history traits, data were collected from a 20-month field study in South India and from captive gaur populations. Mean age of females at first parturition was 3 years; females remained fertile beyond the age of 15 years. Adult females were three times more abundant than adult males in the wild; survival of females was greater than males beyond three years of age. Life span of both sexes has not exceeded 24 years in captivity. Gaur life-history traits are similar to those of other similar-sized Bovini species.

Browsing intensity influences a plant’s response to herbivory. Plants face a trade-off between investment in the production of secondary compounds and investment in growth. To elucidate this trade-off, we simulated four browsing intensities (0%, 50%, 75% and 100%) on mopane saplings, Colophospermum mopane (J.Kirk ex Benth.) J.Leonard, in a greenhouse experiment. This showed that, with increasing defoliation intensity, plants change their investment strategy. At intermediate levels of defoliation (50%), mopane saplings increased the synthesis of condensed tannins, so that tannin concentrations followed a hump-shaped relation with defoliation intensity, with significantly higher tannin concentration at intermediate defoliation levels. When defoliated heavily (75% and 100%), tannin concentrations dropped, and plants were carbon stressed as indicated by a reduced growth rate of the stem diameter, and leaf production and mean individual leaf mass were reduced. This suggests that, at intermediate defoliation intensity, the strategy of the plants is towards induced chemical defences. With increasing defoliation, the relative costs of the secondary metabolite synthesis become too high, and therefore, the plants change their growing strategy. Hence, browsers should be able to benefit from earlier browsing by either adopting a low or a relatively high browsing pressure.

Wild herbivore diversity and abundance have declined in African savannas for the past 20 years. Competition for forage resources between wild herbivore and livestock species might contribute to this decline, given habitat and diet overlap under conditions of resource limitation. Development of conservation and management strategies rely on understanding the spatial distribution of resources for livestock and wildlife, especially since the combination of transhumance pastoralism and wildlife conservation is common over Africa. We studied the distribution patterns of wild grazers in relation to cattle abundance, and distance to permanent and seasonal water bodies in semi-arid Kenya between 1983 and 2000. Cattle were abundant far from permanent water bodies during dry and wet years. Wild grazers less dependent on water also concentrated far from water during all years. Only wild grazer species more dependent on water remained concentrated close to water. Hence, wild grazers and cattle show spatial partitioning in the use of permanent and seasonal water bodies, but not in their forage resources. Our study provides no strong evidence of spatial displacement of wild grazers by cattle. We suggest that pastoralist decisions on cattle distribution do not need to negatively affect wildlife distribution and that coexistence can be possible through spatial partitioning. Our results show that pastoralist decisions play an important role in the interactions between livestock and wildlife in African savannas and that herd mobility is a key component in supporting sustainable use of resources for both wildlife and livestock.

The global decrease of vulture populations has been attributed to several factors, such as food availability, poisoning, human disturbance, or habitat suitability. We studied the effect of factors that vary both spatially and temporally on the nest site distribution of the Griffon Vulture Gyps fulvus in northeast Portugal, and influence the population dynamics of these cliff-dwelling birds. Several demographic parameters were studied in the field, and the age structure of the population was determined. Additionally we investigated how food availability affected population fluctuations. The vulture population was most sensitive to juvenile mortality according to our model results. Our models showed that a decrease in the present food supply resulted in a decrease in vulture abundance and, moreover, negatively affected the spatial distribution of the species by decreasing the number of breeding pairs per colony. The total quantity of sheep and goat biomass, a greater distance to the nearest unpaved road, and the presence of Egyptian Vultures Neophron percnopterus at the breeding colonies showed a significant positive relationship to the numbers of Griffon Vulture breeding pairs. It is recommended that the available food supply in the most important foraging areas should be monitored and protected. Furthermore, to manage the community of cliff breeding raptors adequately, future research should identify and protect the most suitable breeding areas for each species. Human disturbance should be kept to a minimum by protecting breeding areas, especially in an area like Riba-Côa where species such as the Egyptian Vulture, Bonelli's Eagle Hieraaetus fasciatus and Griffon Vulture breed so close to each other.

Patches of wooded vegetation in nutrient-poor grassland characteristically contain high soil moisture and nutrient availability compared with surrounding grassland. These ‘islands of fertility’ appear stable in size, suggesting that tree recruitment at the patch boundary is limited. We hypothesize that tree establishment in adjacent grassland is limited by (1) competition for resources, (2) fire or (3) herbivory. In a South African grassland, we measured moisture availability and conducted a bioassay experiment to analyse whether soil nutrient limits tree recruitment at the patch boundary. We thus measured nutrient concentrations of maize plants grown in patch, boundary and grassland soil. To investigate whether browsing or fire affected tree seedlings at the patch boundary, we burned plots including patches, and used fences to exclude browsers. Neither soil moisture nor nutrient availability at the boundary differed from inside the patches, suggesting that tree recruitment at the boundary is not resource limited. Both fire and browsing combined caused a significantly lower tree seedling growth at the patch boundary, suggesting that these two factors can impede tree recruitment. The balance between positive feedback mechanisms facilitating tree recruitment, and the negative impact of fire and browsing can explain the apparent stability of these islands of fertility.

Amongst many ongoing initiatives to preserve biodiversity, the Millennium Ecosystem Assessment again shows the importance to slow down the loss of biological diversity. However, there is still a gap in the overview of global patterns of species distributions. This paper reviews how remote sensing has been used to assess terrestrial faunal diversity, with emphasis on proxies and methodologies, while exploring prospective challenges for the conservation and sustainable use of biodiversity. We grouped and discussed papers dealing with the faunal taxa mammals, birds, reptiles, amphibians, and invertebrates into five classes of surrogates of animal diversity: (1) habitat suitability, (2) photosynthetic productivity, (3) multi-temporal patterns, (4) structural properties of habitat, and (5) forage quality. It is concluded that the most promising approach for the assessment, monitoring, prediction, and conservation of faunal diversity appears to be the synergy of remote sensing products and auxiliary data with ecological biodiversity models, and a subsequent validation of the results using traditional observation techniques.

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.

Little quantitative information is available about the role of stems as structural foraging deterrents for large grazers and the actual mechanisms by which such deterrents affect foraging behaviour. We measured bite size, bite rate and the rate of forage intake of cattle foraging on artificial micro-swards of the tropical, broad-leaved guinea grass. These micro-swards varied in total forage mass density, in forage quality, defined as the proportion of high-quality plant parts (leaves) and of foraging deterrents (stems), and in the spatial pattern of plant parts. We hypothesized that stems interfere with the process of grasping of leaves and predicted that decreasing forage quality, by reducing bite size and bite rate, depresses the slope and the asymptotic maximum of the functional response curve. Further we hypothesized that increasing cluster size of leaves increases the accessibility of leaves to cattle, thus alleviating the negative effects of decreasing forage quality, and predicted that increasing leaf cluster size has positive effects on bite size and on the rate of forage intake. The slope of the functional response curve decreased with decreasing forage quality, mainly because of depressed bite size. These effects were also found when the amount of leaves in the grass sward was kept constant while only the amount of stems increased. Thus, the observed effects are not merely the result of decreased availability of leaves, but at least in part caused by the increasing interference of stems with the grasping of leaves. Leaf cluster size had a positive effect on bite size. However, this effect did not show in the rate of forage intake, because for small leaf clusters high bite rates compensated for decreased bite sizes. Instead of the familiar negative relationship of bite rate with bite size, we found a positive relationship. We speculate that this effect is the result of decreased chewing times for small bites and increased time needed to grasp these bites, effectively changing the limitation of bite rate from chewing time to a limitation by the time needed to grasp bites.

Resource manipulation, such as the creation and maintenance of grazing lawns, may shape the structure of herbivore communities. We tested the hypothesis that grazing lawns contribute towards the subsistence of the Kobus kob kob in a dystrophic West African savanna, where kob and Hippopotamus amphibius both occur. Comparison of the foliage of grazing lawns and ungrazed swards shows that hippo lawns are more nutritious with regard to both structure and nutrients; kob lawns are higher in nutrients only. Up to the early dry season, hippo lawns meet kob energy and protein demand, thereafter, the shortness of the sward limits intake. Kob lawns always provide sub-maintenance values. Grazing on ungrazed swards is least profitable. We suggest that grazing lawns are essential for the daily subsistence of mesoherbivores, particularly on nutrient-poor soil, and that megaherbivores facilitate their food supply, for at least part of the year.

In temperate regions climate change has led to advances in plant phenology which may disrupt the synchrony between food availability and reproductive requirements of higher trophic levels. Because leaf quality generally drops with leaf maturation, for herbivorous animals a stoichiometric effect of climate change may be of greater importance than a shift in food quantity. We hypothesized that such a climate-related stoichiometric effect caused the low reproductive success of a beaver Castor fiber population that was translocated westward in Europe at the time of a rapid increase in spring temperatures. The staple food of these beavers was willow Salix spp. In 2003 willow bud burst occurred approximately one month earlier at the release site than at the site of origin. After bud burst, leaf quality, measured as nitrogen and phosphorus concentration, decreased in the course of spring in a predictable manner with accumulated degree-days. Nitrogen and phosphorus concentrations calculated from accumulated degree-days were, at a given date, around 20% lower at the release site during the 15 years after the translocation than at the site of origin during the 15 years prior to the translocation. Previous work had identified phosphorus as a likely limiting resource for beaver reproduction, and in most post-translocation years phosphorus concentrations at the release site were below the required level towards the end of gestation. In addition to an increase over time, annual reproductive rate of the translocated beavers was highest in cold springs. We conclude that it was initially low due to a combination of low food quality and factors not related to climate.

Germination of Acacia tortilis seeds strongly depends on micro-site conditions. In Lake Manyara National Park, Tanzania, Acacia tortilis occurs abundantly in recently abandoned arable fields and in elephant-mediated gaps in acacia woodland, but does not regenerate in grass swards or beneath canopies. We examined the germination of Acacia tortilis using field and laboratory experiments. Seeds placed on top of the soil rarely germinated, while seeds covered with elephant dung or buried under the soil surface had a germination success between 23–43%. On bare soil 39% of both the dung-covered and buried seeds germinated, in perennial grass swards 24–43%, and under tree canopies 10–24% respectively. In laboratory experiments, seed water absorption correlated positively with temperature up to 41 °C, while subsequent germination was optimal at lower (21–23 °C) temperatures. Seeds that had absorbed water lost their viability when kept above 35.5 °C. The absence of light did not significantly influence germination success. Acacia tortilis does not actively disperse its seeds, but regeneration outside tree canopies was substantial. The regeneration potential thus strongly depends on the physiognomy of the vegetation.

1. The issue of competition between livestock and wild herbivores has remained contentious. We studied the diets and population structures of the mountain ungulate bharal Pseudois nayaur and seven species of livestock to evaluate whether or not they compete for forage. The study was conducted in the high altitude Spiti Valley, Indian Trans-Himalaya. 2. We compared resource (forage) availability and bharal population structures between rangelands differing in livestock density. Forage availability was estimated by clipping the standing graminoid biomass in sample plots. Livestock and bharal population structures were quantified through annual censuses. Seasonal diets of livestock were studied by direct observations, while those of bharal were quantified through feeding signs on vegetation. 3. We found that livestock grazing causes a significant reduction in the standing crop of forage. Graminoid availability per unit livestock biomass was three times greater in a moderately grazed rangeland compared with an intensively grazed one. 4. There was considerable diet overlap among the herbivore species. In summer, bharal, yak Bos grunniens, horse Equus caballus, cow Bos indicus, and dzomo (yak-cow hybrids) fed predominantly on graminoids, while donkey E. asinus, sheep Ovis aries, and goat Capra hircus, consumed both graminoids and herbs. The summer diet of bharal was a subset of the diets of three livestock species. In winter, depleted graminoid availability caused bharal, yak and horse to consume relatively more herbs, while the remaining livestock species fed predominantly on graminoids. Diet overlap was less in winter but, in both seasons, all important forage species in the bharal diet were consumed in substantial amounts by one or more species of livestock. 5. Comparison of the population structures of bharal between two rangelands differing in livestock density by c. 30% yielded evidence of resource competition. In the intensively grazed rangeland, bharal density was 63% lower, and bharal population showed poorer performance (lower young : adult female ratios). 6. Synthesis and applications. High diet overlap between livestock and bharal, together with density-dependent forage limitation, results in resource competition and a decline in bharal density. Under the present conditions of high livestock density and supplemental feeding, restricting livestock numbers and creating livestock-free areas are necessary measures for conserving Trans-Himalayan wild herbivores. Mediating competitive effects on bharal through supplemental feeding is not a feasible option.

Understanding livestock grazing impacts on native wildlife is difficult when grazing is pervasive and has a long history. Spiti Valley in the Indian Trans-Himalaya is characterized by overstocking of rangelands and a grazing history of over 3 millennia. An intriguing aspect of the wild large-herbivore assemblage in Spiti Valley is its low diversity. In the present theoretical exploration, data on body masses of wild large-herbivore species in the Trans-Himalaya are examined against the backdrop of competition theory, to evaluate the possible role of competition in structuring the herbivore assemblage. The analysis is then expanded to include the livestock assemblage, and the possibility of the low wild herbivore diversity in Spiti Valley being a consequence of the high livestock diversity is explored. Null model analyses suggest that competitive interactions may have played a role in structuring the Trans-Himalayan wild herbivore assemblage. This is reflected in a proportional regularity in species body masses, with each species, on average, being a constant proportion larger than the nearest smaller one. Such a proportional regularity is absent in the livestock assemblage. The analyses make a case for the competitive exclusion of at least four wild grazers in Spiti over the last 3 millennia. The present grazer assemblage conforms reasonably to the theoretical predictions ‐ these four species are among the six wild herbivores presently missing from Spiti. The implications of the analyses for conservation management in the Trans-Himalaya are discussed.