Sward height and bite size affect the functional response of barnacle geese Branta leucopsis

Intake rate, the rate in which herbivores can process their food, is presumed to be an important factor in habitat selection down to the scale of the foraging patch. Much attention has been given to the selection of swards of high nutritional quality, but much less has been given to the influences of sward structure on patch selection in small herbivores. In this study we tested the effects of sward density and height on the functional foraging response of barnacle geese, Branta leucopsis. The functional response curve for herbivores describes how intake rate is affected by food availability. We conducted feeding trials to determine intake rate and bite size of barnacle geese on experimentally manipulated swards. Results indicate that intake rate is mainly dependent on sward height and that there is a strong correlation between bite size and intake rate. Sward density does not influence the rate of food consumption; it is, however, a crucial parameter affecting potential total yield. We conclude that bite size is the crucial parameter influencing intake rate. Bite size is explained both by sward height and individual differences in bill morphology. Furthermore, intake rate seems to be dependent on the physical structure of the grass species consumed.     

Increased searching and handling effort in tall swards lead to a Type IV functional response in small grazing herbivores

Understanding the functional response of species is important in comprehending the species’ population dynamics and the functioning of multi-species assemblages. A Type II functional response, where instantaneous intake rate increases asymptotically with sward biomass, is thought to be common in grazers. However, at tall, dense swards, food intake might decline due to mechanical limitations or if animals selectively forage on the most nutritious parts of a sward, leading to a Type IV functional response, especially for smaller herbivores. We tested the predictions that bite mass, cropping time, swallowing time and searching time increase, and bite rate decreases with increasing grass biomass for different-sized Canada geese (Branta canadensis) foraging on grass swards. Bite mass indeed showed an increasing asymptotic relationship with grass biomass. At high biomass, difficulties in handling long leaves and in locating bites were responsible for increasing cropping, swallowing, and searching times. Constant bite mass and decreasing bite rate caused the intake rate to decrease at high sward biomass after reaching an optimum, leading to a Type IV functional response. Grazer body mass affected maximum bite mass and intake rate, but did not change the shape of the functional response. As grass nutrient contents are usually highest in short swards, this Type IV functional response in geese leads to an intake rate that is maximised in these swards. The lower grass biomass at which intake rate was maximised allows resource partitioning between different-sized grazers. We argue that this Type IV functional response is of more importance than previously thought.     

Effects of sward structure on herbivore foraging behaviour in a South African savanna: An investigation of the forage maturation hypothesis

Abstract: The grass layer of savannas is characterized by strong temporal and spatial heterogeneity in the quantity and quality of forage. Besides this, there is strong variation in other aspects of sward structure, here defined as sward chemistry, morphology, architecture and species composition. The forage maturation hypothesis (FMH) is based on the temporal dynamics of forage quantity and quality of grasslands. Recently this hypothesis has been used to explain the foraging behaviour of large herbivores as an optimal solution for balancing forage ingestion and forage digestion, leading to a maximization of daily rates of forage intake in patches of intermediate forage quantity. However, so far studies using this hypothesis have been constrained to mono‐specific forage resources or have ignored inter‐patch variation in sward structure. We studied the foraging behaviour of cattle in a South African savanna. We explicitly addressed forage quantity and quality, and the structure of seven sward types in order to investigate (i) their effects on foraging behaviour; and (ii) the assumptions and predictions of the FMH for foraging behaviour in grasslands with pronounced variation in sward structure. The results indicated that the assumptions of the FMH were met and that forage quantity affected forage intake the most, with sward structure having little or no effect. The predicted maximum rate of daily forage intake agreed with the results of previous studies. However, the forage quantity at which this maximum intake rate was found, was larger than expected based on the results of some studies. It is likely that this discrepancy resulted from the use of artificial leaf‐only grass swards in these studies. The results suggest that the FMH can be used to explain the foraging behaviour of herbivores over a wide range of sward structures.     

Feeding patch selection by herbivorous Anatidae: the influence of body size, and of plant quantity and quality

Recent findings suggest that herbivores select feeding sites of intermediate biomass in order to maximise their digestible nutrient intake as the result of the trade-off between forage quality and quantity ('forage maturation hypothesis'). We propose a reformulation of this hypothesis which recognises this trade-off, but also underlines that constraints due to body mass (i.e. metabolism and digestive constraints, size of the feeding apparatus) can lead to variations in grazing patterns. We tested this latter hypothesis experimentally in three species of herbivorous Anatidae of different body mass: the wigeon Anas penelope (in our study c. 620 g), the barnacle goose Branta leucopsis (c. 2000 g), and the greylag goose Anser anser (c. 3500 g). Each species was tested separately from 0600 to 0930 hours, in an enclosure with a mosaic of patches of grass of three different heights: short, medium and tall. The behaviour, and the location (i.e. patch) of each individual were recorded every 5 minutes. Our results show important interspecific differences in intake rates resulting in different feeding site selection: wigeon and barnacle goose fed fastest on the shortest swards, and selected short grass which was also of higher quality. Tall grass provided the highest dry matter intake rate and digestible protein intake for greylag geese, and they preferred these swards. These choices allowed the birds to maximise their digestible nitrogen intake rate rather than dry matter intake rate and our results thus underline the importance of nitrogen as a major currency for foraging decisions in herbivorous Anatidae. Since the birds selected the two extreme sward heights (instead of the medium one), the results give support to our hypothesis and underline the role of body size as an important cause of variations in patch selection in herbivorous Anatidae.     

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.     

Importance of nutritional and anti-parasite strategies in the foraging decisions of horses: an experimental test

The primary goal of foraging herbivores is to maximise the net rate of intake of digestible energy (or of a limiting nutrient). However, foraging strategies of herbivores are also sensitive to other selective forces (e.g. predation, parasites), which may modify their choice of feeding patches. Horses feed in spatially complex swards, and allocate their time among patches which vary both in terms of their nutritional benefits, and the risk of parasitism. It has long been suggested that horses allocate time among patches principally in relation to the risk of parasitism, though the nutritional costs and benefits of different foraging choices must play some role too. In this study, we investigated the roles of nutritional and anti-parasite factors in foraging decisions by horses. Six naturally parasitized and six unparasitized two-year-old geldings were allowed to graze from pairs of trays (112×72 cm) with swards at two heights (tall: 52 cm, medium: 15 cm) and two levels of contamination (280 g m⁻² of faeces, no faeces) in a latin square, pair-wise design with six different choices. In nutritional terms the differences between the swards were slight: the tall sward provided larger bite sizes and higher intake rates (+36%). Feeding motivation was not high, and the faecal stimulus was strong. The horses selected uncontaminated swards when the nutritional benefits were identical, and tall swards in virtually all circumstances. Contamination of the preferred tall sward (i.e. a test of the tradeoff between anti-parasite and nutritional strategies) had little effect on its selection by the horses. Their parasite status also had little effect: both groups selected the tall sward in the tradeoff (though this was only a trend for the non-parasitised group in one analysis). We conclude that patch selection by horses in these particular conditions was driven principally by their nutritional strategy.     

Feeding-patch choice by red deer in relation to foraging efficiency

We tested the idea that ruminants allocate their feeding time to habitat patches in relation to foraging efficiency. We used five tame red deer (Cervus elaphus) in an enclosure planted with four treatment of timothy grass (Phleum pratense) differing in their stage of growth. Older swards offered higher biomass but lower nutritional quality than younger swards. We observed time spent feeding in each treatment during each of seven trials. We measured goodness-of-fit between observed times and predictions from two alternative hypotheses differing in optimization strategy (maximizing versus matching), and a third, null hypothesis. We tested the hypotheses using two alternative currecies: digestible protein, and digestible dry matter or energy. Although digestible protein concentration and dry-matter digestibility were highly correlated (r=0.763, P<0.001), the wider range of digestible protein made it the much more sensitive measure of forage quality. Distributions of feeding time closely matched estimated intake rates of digestible protein (R _infPred ^sup2 =0.899) across all animals and trials. The other hypotheses were rejected. The results have important ecological implications in showing the underlying role of food in the selection of habitat by ruminants, and that simple, mechanistic models of forage intake and digestion can be scaled up to the level of animal behavioural choices.     

Foraging behaviour of brent geese, Branta b. bernicla, on grasslands: effects of sward length and nitrogen content

Feeding behaviour and preferences of brent geese, Branta b. bernicla were observed on pastures of different sward lengths and nitrogen contents. On swards of 2.0-6.0 cm captive geese took larger bites and had a higher intake rate when feeding on 6-cm swards than when feeding on shorter ones. In the field wild geese chose 6-cm swards in preference to both shorter and longer ones. For unfertilized pastures there was a strong negative relationship between nitrogen content and sward height: 11-cm swards contained 2.8% nitrogen, 3.5-cm swards 4.2% nitrogen. Application of 75 kg N ha^-1 organically based fertilizer at the end of October eliminated this relationship between nitrogen content and sward height, swards of all heights then having a mean content of 4.1% N. On fertilized plots the geese preferred swards longer than 6 cm with no indication of a decrease in preference up to the maximum height investigated, 16 cm. Breaking strain of grass laminae was measured to give an indication of the proportion of strengthening tissues in the leaves and hence their digestibility. Apical laminae from longer swards had a higher breaking strain than those from shorter swards. Unfertilized swards had a higher breaking strain than fertilized swards but the difference in breaking strain between long and short swards was the same on fertilized and control treatments. These results are discussed in relation to the forage maturation hypothesis and are interpreted as indicating that the primary constraint on maximizing energy intake rates is not the reduced energy digestibility of older foliage but the reduced nitrogen content in the higher-biomass swards. We conclude that it is the balance between maximizing energy intake and nitrogen absorption rates which results in the observed preference for intermediate-height swards.     

Grazing decisions of Soay sheep,Ovis aries,on St Kilda: a consequence of parasite distribution?

Relationships between the height of grass swards and herbage intake have been established for a number of large herbivores and sward height plays a major role in determining the selection of herbivore diets. However, tall grass swards also represent a more stable damp microclimate for gastrointestinal parasite larvae and tall swards contain generally greater numbers of parasites than short swards. Herbivores may then trade-off the need to maximise nutrient intake through the selection of tall swards with the need to avoid parasite ingestion. Stratified sward sampling techniques were used to determine the distribution of nutrients and parasites in a heterogeneous sward structure on the island of Hirta, St Kilda, which is grazed by a population of feral Soay sheep. The disparity between the short gap vegetation and the tall tussock vegetation was greatest in spring, when gap vegetation was some three-fold shorter than tussock vegetation; this led to tussocks offering greater nutrient and energy intake rate when compared to gap vegetation in spring. Parasites were concentrated in tussocks in spring, thus creating trade-offs. Such trade-offs were not present in summer when parasites were more evenly distributed across the gap/tussock sward structure and the nutritional advantages associated with grazing tussocks were diminished. The diet selection of the resident population of sheep was determined by recording the number of bites taken from gap and tussock vegetation by randomly selected focal animals over repeated 5-min grazing periods. Overall, all animals avoided grazing tussocks, which were most strongly avoided in the spring, and older animals avoided grazing tussocks to a greater extent. Overall, females with lambs and males avoided tussocks to a similar degree and both avoided them less than barren females. Faecal egg counts (FEC) of female sheep were negatively correlated with tussock selectivity and vice versa for male sheep. The interaction between the grazing behaviour of each sex and FEC is discussed in relation to the immunocompetence and nutrient requirements of the different sexes. The maximum disparity between the costs and benefits of the trade-off studied occurred in late winter/early spring which also represents the time of greatest nutrient demand in the Soay sheep, since many are close to starvation and/or are in the advanced stages of pregnancy. Grazing decisions of individuals at this time, determining relative nutrient and parasite intake, may then be related to subsequent fitness and survival.     

Genetically resistant sheep avoid parasites to a greater extent than do susceptible sheep

Livestock breeding programmes have created resistant (R) and susceptible (S) sheep that differ in their ability to control parasites through their immune function but potentially also their grazing behaviour (i.e. parasite avoidance). Using the Perendale genetic lines, we tested the hypothesis that R-sheep avoid parasites more effectively, reducing their parasite exposure/challenge, compared with S-sheep. However, in grazing systems, parasite-rich areas are also forage rich, suggesting that parasite avoidance behaviours are associated with nutritional penalties. We first created a naturally heterogeneous sward structure of gaps and tussocks and then used focal behavioural observations to quantify the sward selection of R- and S-sheep. Tussock swards were more nitrogen rich (41%), offered increased forage intake rates (32%) and contained 17 times more parasite larvae than gap swards. All the animals avoided grazing the tussock swards. However, the R-sheep grazed the tussock swards to a lesser degree than the S-sheep. We conclude that selection for genetic resistance has resulted in animals that, despite being well armed to fight parasitism through improved immune function, adopt parasite avoidance strategies with associated nutritional disadvantages. This experiment highlights the role of host behaviour in the control of parasitism and suggests that animals can be bred to avoid disease.     

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

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

Voluntary intake and digestibility in horses: effect of forage quality with emphasis on individual variability

Food intake is a key biological process in animals, as it determines the energy and nutrients available for the physiological and behavioural processes. In herbivores, the abundance, structure and quality of plant resources are known to influence intake strongly. In ruminants, as the forage quality declines, digestibility and total intake decline. Equids are believed to be adapted to consume high-fibre low-quality forages. As hindgut fermenters, it has been suggested that their response to a reduction in food quality is to increase intake to maintain rates of energy and nutrient absorption. All reviews of horse nutrition show that digestibility declines with forage quality; for intake, however, most studies have found no significant relationship with forage quality, and it has even been suggested that horses may eat less with declining forage quality similarly to ruminants. A weakness of these reviews is to combine data from different studies in meta-analyses without allowing the differences between animals and diets to be controlled for. In this study, we analysed a set of 45 trials where intake and digestibility were measured in 21 saddle horses. The dataset was analysed both at the group (to allow comparisons with the literature) and at the individual levels (to control for individual variability). As expected, dry matter digestibility declined with forage quality in both analyses. Intake declined slightly with increasing fibre contents at the group level, and there were no effects of crude protein or dry matter digestibility on intake. Overall, the analysis for individual horses showed a different pattern: intake increased as digestibility and crude protein declined, and increased with increasing fibre. Our analysis at the group level confirms previous reviews and shows that forage quality explains little of the variance in food intake in horses. For the first time, using mixed models, we show that the variable 'individual' clarifies the picture, as the horses showed different responses to a decrease in forage quality: some compensated for the low nutritional value of the forages by increasing intake, few others responded by decreasing intake with declining forage quality, but not enough to cause any deficit in their energy and protein supplies. On the whole, all the animals managed to meet their maintenance requirements. The individual variability may be a by-product of artificial selection for performance in competition in saddle horses.     

The effect of sward height and drainage on Common Starlings Sturnus vulgaris and Northern Lapwings Vanellus vanellus foraging in grassland habitats

Agricultural change is often cited as a causal factor in the decline of the UK's farmland birds because bird declines have mirrored changes in agricultural practices. Although much is known about the mechanisms driving population declines on arable systems, mechanisms in grassland systems are relatively poorly studied, despite receiving a similar degree of intensification. Agricultural intensification may affect bird declines by reducing food abundance or accessibility, forager mobility or predation risk. Here we examine experimentally the effects of sward height on the foraging behaviour of adult Common Starlings Sturnus vulgaris , and the effects of sward height and drainage on the behaviour of Northern Lapwing Vanellus vanellus chicks. Both species are declining across the UK and both forage in farmed grassland habitats, but they differ in their foraging methods because Starlings probe for soil invertebrates whereas Lapwing chicks glean prey from surfaces. Overall, after controlling for prey abundance, short swards were found to be more productive for both species. Prey capture rate within foraging bouts did not differ with sward height for Starlings, but Starlings spent more time foraging on short swards and captured 33.2% more prey. Starlings walked more steps on short swards. Lapwing chick foraging rates declined as sward height increased. Soil moisture was not found to be a predictor of Lapwing chick foraging rates within the observed range. Our results suggest that short swards are a more profitable foraging habitat for soil and surface invertebrate feeders. Short swards may facilitate surface prey detection, improve forager mobility and increase foraging time by altering vigilance patterns. Provision of short swards in areas where these are lacking could be simple method of improving foraging habitats for grassland birds.     

Influence of sward structure on daily intake and foraging behaviour by horses

The spatial heterogeneity of grasslands determines the abundance and quality of food resources for grazing animals. As plants mature, they increase in mass, which allows greater instantaneous intake rates, but the cell wall concentrations increase too, reducing diet quality. In ruminants, daily intake rates are often constrained by the time needed for the ingesta to pass through the rumen, which is influenced by the rate of digestion. It has been suggested that the digestive constraint should have much less effect on hindgut fermenters such as equids. Horses play an increasing role in the management of grasslands in Europe, but the data on the influence of the heterogeneity of the vegetation on their daily intake and foraging behaviour are sparse. We report here the results of a preliminary study concerning the effects of sward structure on nutrient assimilation and the use of patches of different heights by horses grazing successively a short immature, a tall mature and a heterogeneous pastures (with short and tall swards). Daily nutrient assimilation was higher in the heterogeneous pasture compared to the short (+35%) and the tall (+55%) ones. The digestive constraints may have limited voluntary intake by horses on the tall swards. In the heterogeneous pasture, the mean height used for feeding (6 to 7 cm) by horses was intermediate between the heights used in the short (4 to 5 cm) and tall pastures (22 to 23 cm), and the animals may thus have benefited from both short swards of high quality and tall swards offering a higher instantaneous intake rate.     

Body size dimorphism and sexual segregation in polygynous ungulates: an experimental test with Soay sheep

Sexual segregation in Soay sheep (Ovis aries) was investigated using an experimental approach in order to test the sexual dimorphism-body size hypothesis. Two corollaries of the sexual dimorphism-body size hypothesis were tested: (1) in dimorphic species males, the larger sex, have relatively smaller bite sizes on short swards because of the scaling of incisor arcade with body weight, and (2) they move off earlier to feed on taller but poorer-quality swards when such swards are patchily distributed on a scale which enables the spatial segregation of individuals. Patch choice between sexes was estimated using a matrix of grass patches which differed in both quality and biomass of grass on offer (HQ: high-quality-low-biomass; LQ: low-quality-high-biomass). Sex differences in patch choice and grazing behaviour were tested in short-term preference trials. Incisor breadth showed no significant difference between sexes. On the other hand, muzzle width was dimorphic, with females having a narrower muzzle than males. Bite size was significantly different between the sexes, being smaller in females than in males, although it was not significantly different between sward types. Females had a higher bite rate than males and the bite rate was higher in the HQ sward type than the LQ sward type. When the effect of body mass was removed, no sex differences in muzzle size, bite size or bite rate were found. The intake rate did not differ between the sexes or between sward types. Whilst both sexes preferred the HQ sward type, females spent a significantly longer time feeding on the LQ sward type than did males. The difference detected between the sexes in patch choice was not consistent directly with the sexual dimorphism-body size hypothesis. Alternative explanations based on sex differences in foraging behaviour in relation to body mass sexual dimorphism are discussed to explain the result. Received: 1 February 1999 / Accepted: 12 May 1999     

Changing grass height alters foraging site selection by wintering farmland birds

Grassland is a valuable habitat for many farmland bird species, some of which have declined in Europe. Successful management of this habitat is important to benefit these species. Recent studies have suggested little correlation between food abundance and bird use in grassland; instead providing access to food may be crucial. We tested the importance of manipulating sward height (via mowing) on the distribution of wintering birds in agriculturally managed grass fields using two within-field experimental designs. In experiment 1, we investigated the effect of two treatments (mown once and control). In experiment 2, we investigated the effects of four treatments (mown once early, mown once late, mown early and late, and control).Mown plots supported higher abundances of foraging kestrels, thrushes and starlings. In contrast meadow pipits and grey herons preferred unmown plots with longer grass. These differences are probably explained by differing anti-predation strategies, increased food abundance for species that prefer above-ground invertebrates (e.g. more food for meadow pipits on longer swards) and increased food accessibility on short grass swards (e.g. for starling, thrush spp.). There were limited effects of multiple mowing events on plot use suggesting one early winter mowing treatment may suffice to create favourable conditions for species that prefer shorter swards.Providing grassland heterogeneity should benefit a range of wintering farmland birds. Although we use mowing to change grass sward height, our findings may also apply at other times of year when livestock grazing may be a more practical solution.     

Habitat associations of Yellow Wagtails Motacilla flava flavissima on lowland wet grassland

In common with many other previously widespread British birds, Yellow Wagtail Motacilla flava flavissima populations have declined substantially in the UK in recent decades. Recent national data suggest particular problems in grassland. We investigated how habitat management in a grassland area of East Anglia, England, influenced the habitat selection of this species. Breeding territories were associated with fields previously subject to prolonged winter floods and which contained shallow-edged ponds or wet ditches during summer. Territories were also associated with fields with short, sparse swards and high proportions of bare earth within the sward, and these features were themselves associated with previous winter flooding. However, nests were associated with taller swards than were random points within the same field, suggesting that Yellow Wagtails would benefit from within-field heterogeneity of sward height to provide both nesting and foraging habitat. We discuss how grazing, cutting and water-level management regimes could be used to promote optimal sward structure for Yellow Wagtails.     

Modelling ungulate dependence on higher quality forage under large trees in African savannahs

In African savannahs, large trees improve grass quality, particularly in dry and nutrient poor areas. Enhanced below-canopy grass nutrients, such as nitrogen and phosphorus contents should therefore attract and benefit grazers. To predict whether ungulates really need these forage quality islands we focused on four grazer species, i.e., zebra, buffalo, wildebeest, and warthog, differing in body size and digestive system. We confronted literature estimations of their feeding requirements with forage availability and quality, observed in three South African savannah systems, through linear modelling. The model predicted the proportion of below-canopy grass that grazers should include in their diet to meet their nutritional requirements.During the wet season, the model predicted that all animals could satisfy their daily nutrient requirements when feeding on a combination of below- and outside-canopy grasses. However, wildebeest, having relatively high nutrient demands, could meet their nutrient requirements only by feeding almost exclusively below canopies.During the dry season, all animals could gain almost twice as much digestible protein when feeding on below – compared to outside-canopy forage. Nonetheless, only warthogs could satisfy their nutrient requirements – when feeding almost exclusively on below-canopy grasses. The other ungulate species could not meet their phosphorus demands by feeding at either site without exceeding their maximum fibre intake, indicating the unfavourable conditions during the dry season.We conclude that grazing ungulates, particularly warthog, zebra, and buffalo, actually depend on the available below-canopy grass resources. Our model therefore helps to quantify the importance of higher quality forage patches beneath savannah trees. The composition of grazer communities depending on below-canopy grasses can be anticipated if grazer food requirements and the abundance of large trees in savannahs are known. The model suggests that the conservation of large single-standing trees in savannahs is crucial for maintenance of locally grazing herbivores.     

The herbivores’ dilemma: trade-offs between nutrition and parasitism in foraging decisions

An experiment was carried out using a trade-off framework to determine the rules of sward selection, in relation to gastrointestinal parasite dispersion, used by mammalian herbivores, and the effect of level of feeding motivation and parasitic status on these rules. Twenty-four sheep divided into four animal treatment groups resulting from two levels of feeding motivation (high and moderate) and two parasitic states (parasitised with Ostertagia circumcincta and non-parasitised) were presented with pairs of experimental swards which varied in N content (high and low), sward height (tall and short) and level of contamination with faeces and thus parasites (contaminated and non-contaminated). The selection for tall swards outweighed both the selection for N-rich swards and the avoidance of faecal contaminated swards. The selection for N-rich swards could not completely overcome faecal avoidance. Parasitism in animals with a moderate feeding motivation reduced their bite rates and grazing depths, thereby probably reducing the rate of ingestion of parasitic larvae. In contrast, highly feeding-motivated animals (including those parasitised) increased their bite rates and grazing depths, thereby increasing the rate of ingestion of parasites. The inclusion of parasite distributions, both in the environment and within herbivore host populations, is likely to advance optimal foraging theory by enhancing its predictive power. Received: 30 November 1999 / Accepted: 14 February 2000     

Herbaceous Forage and Selection Patterns by Ungulates across Varying Herbivore Assemblages in a South African Savanna

Herbivores generally have strong structural and compositional effects on vegetation, which in turn determines the plant forage species available. We investigated how selected large mammalian herbivore assemblages use and alter herbaceous vegetation structure and composition in a southern African savanna in and adjacent to the Kruger National Park, South Africa. We compared mixed and mono-specific herbivore assemblages of varying density and investigated similarities in vegetation patterns under wildlife and livestock herbivory. Grass species composition differed significantly, standing biomass and grass height were almost twice as high at sites of low density compared to high density mixed wildlife species. Selection of various grass species by herbivores was positively correlated with greenness, nutrient content and palatability. Nutrient-rich Urochloa mosambicensis Hack. and Panicum maximum Jacq. grasses were preferred forage species, which significantly differed in abundance across sites of varying grazing pressure. Green grasses growing beneath trees were grazed more frequently than dry grasses growing in the open. Our results indicate that grazing herbivores appear to base their grass species preferences on nutrient content cues and that a characteristic grass species abundance and herb layer structure can be matched with mammalian herbivory types.