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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.

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.

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.

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

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.

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.

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.

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.

A key question in savanna ecology is how trees and grasses coexist under N limitation. We used N stable isotopes and N content to study N source partitioning across seasons from trees and associated grasses in a semi-arid savanna. We also used (15)N tracer additions to investigate possible redistribution of N by trees to grasses. Foliar stable N isotope ratio (δ(15)N) values were consistent with trees and grasses using mycorrhiza-supplied N in all seasons except in the wet season when they switched to microbially fixed N. The dependence of trees and grasses on mineralized soil N seemed highly unlikely based on seasonal variation in mineralization rates in the Kruger Park region. Remarkably, foliar δ(15)N values were similar for all three tree species differing in the potential for N fixation through nodulation. The tracer experiment showed that N was redistributed by trees to understory grasses in all seasons. Our results suggest that the redistribution of N from trees to grasses and uptake of N was independent of water redistribution. Although there is overlap of N sources between trees and grasses, dependence on biological sources of N coupled with redistribution of subsoil N by trees may contribute to the coexistence of trees and grasses in semi-arid savannas.

The red fox Vulpes vulpes is usually classi?ed as being territorial, dispersing or transient. Past studies have focused almost exclusively on territorial or dispersing foxes, leaving transient foxes out of the analysis. In this paper, we present spatial-statistical methods for the classi?cation of free-ranging foxes, using 95a?xed kernels and 100␖inimum convex polygons. By means of these procedures we classify individual foxes on the basis of their spatial behaviour, using home-range size and home range shift. Also, we make a methodological comparison between these classi?cation procedures and interpret the composition of these classes ethologically. The procedures apply to a sample of 24 foxes, radio-tracked in the dune area of the Netherlands from January 1997 to June 1999. We analysed size of home range and successive 3-month overlap using a geographical information system (GIS). Classifying the sample using 95Wxed kernel home ranges resulted in two classes of foxes: a class of 20 territorial foxes with relatively small home ranges (< 250 ha), and a class of four dispersing and transient foxes with relatively large home ranges (400±600 ha). This study shows that a fox population can be divided into different classes of individuals in a quantitative statistical way, honouring measured characteristics. This is a clear extension of more informal ways relying on expert judgement applied so far.

Roan antelope are distributed mainly in regions characterized by infertile soils, offering food of low quality. We hypothesized that roan may select localities with higher soil nutrient levels and/or grass swards with more favourable properties in terms of food abundance or quality than generally available in those regions. Roan antelope were observed in a savanna region in South Africa where soils of widely varying nutrient status occurred. Roan favoured open grassland over wooded savanna areas. During the wet season, roan preferred sites with felsite-derived soil of intermediate soil nutrient status. Grasslands growing on nutrient enriched alluvial soils were preferred outside of the early wet season, although most of the favourable sward characteristics were present in other landscape units. Food quantity, rather than quality appeared to attract roan to foraging sites in the late wet and early dry seasons. Food quality appeared more important in the early wet and late dry seasons. The higher degree of clustering of leafy material within foraging swards seemed to be an additional discriminating factor. The factors governing the selection of foraging sites by roan did not seem notably different from those influencing other species of grazing ruminant, but roan nevertheless seemed tolerant of stemmy grasslands growing on nutrient richer substrates.

There is growing concern that increasing eutrophication causes degradation of coastal ecosystems. Studies in terrestrial ecosystems have shown that increasing the concentration of nitrogen in soils contributes to the acidification process, which leads to leaching of base cations. To test the effects of eutrophication on the availability of base cations in mangroves, we compared paired leaf and soil nutrient levels sampled in Nypa fruticans and Rhizophora spp. on a severely disturbed, i.e. nutrient loaded, site (Mahakam delta) with samples from an undisturbed, near-pristine site (Berau delta) in East Kalimantan, Indonesia. The findings indicate that under pristine conditions, the availability of base cations in mangrove soils is determined largely by salinity. Anthropogenic disturbances on the Mahakam site have resulted in eutrophication, which is related to lower levels of foliar and soil base cations. Path analysis suggests that increasing soil nitrogen reduces soil pH, which in turn reduces the levels of foliar and soil base cations in mangroves.

Conversion of mangroves to shrimp ponds creates fragmentation and eutrophication. Detection of the spatial variation of foliar nitrogen is essential for understanding the effect of eutrophication on mangroves. We aim (i) to estimate nitrogen variability across mangrove landscapes of the Mahakam delta using airborne hyperspectral remote sensing (HyMap) and (ii) to investigate links between the variation of foliar nitrogen mapped and local environmental variables. In this study, multivariate prediction models achieved a higher level of accuracy than narrow-band vegetation indices, making multivariate modeling the best choice for mapping. The variation of foliar nitrogen concentration in mangroves was significantly influenced by the local environment: (1) position of mangroves (seaward/landward), (2) distance to the shrimp ponds, and (3) predominant mangrove species. The findings suggest that anthropogenic disturbances, in this case shrimp ponds, influence nitrogen variation in mangroves. Mangroves closer to the shrimp ponds had higher foliar nitrogen concentrations.

Multiple factors determine diet selection of herbivores. However, in many diet studies selection of single nutrients is studied or optimization models are developed using only one currency. In this paper, we use linear programming to explain diet selection by African elephant based on plant availability and nutrient and deterrent content over time. Our results indicate that elephant at our study area maximized intake of phosphorus throughout the year, possibly in response to the deficiency of this nutrient in the region. After adjusting the model to incorporate the effects of this deficiency, elephant were found to maximize nitrogen intake during the wet season and energy during the dry season. We reason that the increased energy requirements during the dry season can be explained by seasonal changes in water availability and forage abundance. As forage abundance decrease into the dry season, elephant struggle to satisfy their large absolute food requirements. Adding to this restriction is the simultaneous decrease in plant and surface water availability, which force the elephant to seek out scarce surface water sources at high energy costs. During the wet season when food becomes more abundant and energy requirements are satisfied easier, elephant aim to maximize nitrogen intake for growth and reproduction. Our study contributes to the emerging theory on understanding foraging for multiple resources

Seasonal variation in forage availability and quality is understood to affect the annual timing of parturition in large herbivores. In India–where seasonal monsoonal rains define variation in forage availability and quality–chital Axis axis exhibit stronger seasonality in parturition than the larger gaur Bos gaurus. We hypothesized that this difference can be explained by forage requirements determined by body mass. We developed a model to simulate changes in leaf biomass and nitrogen content based on plant available moisture and nutrients, and calibrated our model with field data. Our results show that the minimum forage nitrogen content required by lactating gaur was available throughout the year, but that required by lactating chital was available for less than 40% of the year, i.e. during the early wet season, which coincides with the annual peak period of chital births. The three to four month spread of chital births, which begins in the dry season, implies that the period of highest quality is also important for females to replenish maternal reserves for future reproduction and help maximize the growth rate of neonates. This spread also indicates low synchrony of chital births and suggests that predator swamping was less important in influencing their timing of parturition. As monsoonal rain exhibits annual temporal variation, we analyzed our model under different rainfall patterns while keeping the total annual rainfall constant. We found that the difference between the durations of how long forage quality is available to satisfy the minimum requirements of lactating gaur and lactating chital is similar for all simulated patterns. This insensitivity to variable rainfall patterns lends support to our hypothesis that forage requirements determined by body mass is one plausible explanation for the variation in parturition strategies among large herbivores species. Many animals live in environments where the availability and quality of food follow cyclical patterns of seasonal variation. In the temperate zone, food of adequate quantity is available only during the summer and not in winter. In tropical regions with seasonal rainfall, abundant food of high quality during the wet season is replaced by less abundant food of low quality during the dry season. This seasonality in food availability and quality is understood to influence the timing of parturition of mammalian herbivores (Rutberg 1987, Wittemyer et al. 2007). The ‘seasonality’ hypothesis proposes

There is little understanding of how large mammalian herbivores in Asia partition habitat and forage resources, and vary their diet and habitat selection seasonally in order to coexist. We studied an assemblage of four large herbivores, chital (Axis axis), sambar (Cervus unicolor), gaur (Bos gaurus) and Asian elephant (Elephas maximus), in the seasonal tropical forests of Bandipur and Mudumalai, South India, and tested predictions regarding the species’ seasonal diet browse : graze ratios, habitat selection and habitat-niche preference and overlap. Field data collected for the study included the seasonal variation in grass quality, the seasonal variation in δ13C in the species’ faeces and the seasonal variation in the species’ habitat selection and overlap using a grid-based survey. Results of the δ13C analyses showed that the chital was more of a grazer in the wet season (−17.9‰ to −21.6‰), but that it increased the proportion of browse in its diet in the dry season (−25.6‰ to −27.7‰); the gaur was a grazer for most of the year (−15.3‰ to −18.6‰); the sambar preferred to browse throughout the year (−21.1‰ to −30.4‰); and that the elephant was a mixed feeder (−14.2‰ to −21.4‰). Elephant habitat-niche breadth was high (0.53 in wet and 0.54 in dry) and overlapped equally with that of the other species in both seasons (0.39–0.94). The gaur had the most restricted habitat-niche breadth in both seasons (0.25 in wet and 0.28 in dry), and it switched from the moist deciduous habitat in the dry season to the dry deciduous habitat in the wet season. These results offer the first insights into the seasonal variation in browse : graze diet ratios and the habitat-niche overlap amongst the common largest-bodied mammalian herbivore species found in South India.

This study’s goal was to better understand the growth pattern and limitations of the herbaceous production that supports South India’s rich large herbivore grazer assemblage. We conducted a fully factorial nitrogen and water (three levels each) treatment field experiment in the herbivore rich South Indian Western Ghats region to determine the seasonal pattern and the extent to which nitrogen and water availability limit herbaceous production. Graminoid production was found to be nitrogen limited. Despite low rainfall, additional water did not significantly increase overall biomass production nor extend growth in the dry season. Accumulated standing biomass was highest in the late wet season (November) and lowest in the dry season (May). Leaf nitrogen was highest in the early wet season (June) and lowest in the late dry season (March). Grazing had a positive effect on grass production by extending the growing season. Biomass production and graminoid leaf nitrogen concentration levels in the study area were similar to other tropical areas in the world. Also similar to other tropical large herbivore areas, the dry season poses an annual challenge for large herbivores in the study area —particularly the smaller bodied species—to satisfy their nutrient requirements.

1. Globally, both climatic patterns and nitrogen deposition rates show directional changes over time. It is uncertain how woody seedlings, which coexist with herbaceous plants in savannas, respond to concurrent changes in water and nutrient availability. 2. We investigated competition effects between herbaceous vegetation and tree seedlings (Colophospermum mopane) under changed water and nutrient (fertilized) conditions in a garden experiment situated in a semi-arid savanna. 3. Herbaceous competition significantly suppressed woody seedling growth. The effect of herbaceous competition on woody seedling growth remained constant with both increasing water and nutrient availability. However, during a wet-season drought, herbaceous competition apparently caused premature leaf senescence in non-irrigated treatments. Fertilization exacerbated negative competition effects during the drought, while irrigation prevented leaf loss of tree seedlings in spite of herbaceous competition and fertilization. 4. Based on a conceptual model, we propose that the vigorous response of herbaceous plants to increased nutrient availability leads to faster depletion of soil water, which increasingly causes water stress in woody seedlings if the interval between watering events is prolonged, e.g. during wet-season droughts. 5. Synthesis. Our data support the notion that changes in drought frequency are of greater importance to woody recruitment success than changes in annual rainfall amount. Based on the water and nutrient interactions observed in our experiment, we suggest that the effect of increased nitrogen deposition on woody seedling recruitment is contingent on water availability.

Severe defoliation as occurs in insect outbreaks can alter the chemistry and food value of woody-plant leaves for insects for several years after defoliation ceases. Two hypotheses, the active-defense hypothesis (AD) and the hypothesis of a carbon/nutrient balance (CNB), attempt to explain the mechanism of such responses to defoliation. We tested these hypotheses by studying the responses to severe defoliation by six southern African savanna woody species. Manual defoliation of the three fast-growing deciduous species we studied (Grewia flavescens, Acacia tortilis, Dichrostachys cinerea) resulted in increased resistance to insect attack in the next year. Concomitantly, nitrogen (N) and phosphorus (P) concentrations in leaves decreased and leaf total phenol and condensed tannin concentrations increased. The three slowly growing species that we studied (Burkea africana, Ochna pulchra, Euclea natalensis) responded differently to manual defoliation. In the year following defoliation, leaf resistance to insect...

The functional response as the link between the consumer and its resource is a central issue in herbivore–vegetation interactions. Vegetation often consists of low and high quality tissue due to nutritional or structural differences. It has been observed that the instantaneous intake rate of herbivores decreases with decreasing forage quality. However, so far variation in forage quality is not explicitly considered when modeling this instantaneous intake rate. In this paper, we derive a model for the instantaneous intake rate depending on forage quality, i.e., the proportion of high quality tissue in the vegetation. In the model, a downward deflection of the functional response curve is caused by a reduction of the maximum consumption rate at high proportions of low quality tissue. The model gives a mechanistic explanation for decreasing intake rate with decreasing forage quality. Compared to a conventional functional response model with constant maximum consumption rate, a herbivore–grass model with the forage quality-dependent functional response leads to discontinuous changes in the vegetation and herbivore density. The effects hinge on the positive feedback between herbivore density and the proportion of high quality forage. Our analyses show that this positive feedback can explain the maintenance of lawn grass where a high herbivore density can maintain high quality forage. The model results indicate that depending on the coefficients of trophic conversion from resource to consumer, co-existence, facilitation and competitive exclusion between differently sized herbivore species can emerge.