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.     

The allometry of patch selection in ruminants

An axiomatic feature of food consumption by animals is that intake rate and prey abundance are positively related. While this has been demonstrated rigorously for large herbivores, it is apparent from patch selection trials that grazers paradoxically tend to prefer short, sparse swards to tall, dense swards. Indeed, migratory herbivores often shift from areas of high to low sward biomass during the growing season. As nutritional quality is an inverse function of grass abundance, herbivores appear to sacrifice short-term intake for nutritional gains obtainable by eating sparse forage of higher quality. Explicit models of this trade-off suggest that individual ruminants maximize daily rates of energy gain by choosing immature swards of intermediate biomass. As body mass is related positively to both ruminant cropping rates and digestibility, there should be an allometric link between grass abundance and energy maximization, providing a tool for predicting patterns of herbivore habitat selection. We used previously published studies to develop a synthetic model of trade-offs between forage abundance and quality predicting that optimal sward biomass should scale allometrically with body size. The model predicts size-related variation in habitat selection observed in a guild of grazing ungulates in the Serengeti ecosystem.     

Effects of fire on sward structure and grazing in western Serengeti, Tanzania

In Serengeti fire is used as a management tool to improve the forage quality for large herbivores. However, little is known of the effects of fire on grazing resources particularly sward structure, its influence on herbivore forage patch selection and utilization to the relative amount of phytomass consumed in burnt and nonburnt patches. From September 2003 to July 2004, consumption of phytomass by large herbivores was assessed with eight samplings in six grassland sites in the Western Corridor in Serengeti National Park. Each site had burnt and nonburnt plots. Movable cages were used to exclude grazing between samplings and plant material harvests were used to assess phytomass and sward structure changes in time. Nonburnt grasslands had consistently larger phytomass at all sampling events whereas the ratio for live leaf/total phytomass was higher in burnt grassland at early postfire stages, but declined later in the season. Moreover, periodic consumption of both total phytomass and different phytomass components shifted between burnt and nonburnt grasslands, but there were also large site-specific responses. The shift appears to be related to the balance between the amount of phytomass available and the quality of the forage in terms of the ratio between live and total phytomass. The study highlights the significance of maintaining mosaics of burnt and nonburnt areas with an adequate provision of forage amount and quality all year round.     

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.     

Temporary grazing exclusion promotes rapid recovery of species richness and productivity in a long‐term overgrazed Campos grassland

Moderate grazing intensity is considered the basic requirement to enhance ecosystem function in grasslands. Yet, deterioration by overgrazing is common in many biomes, including Campos grasslands in South America. Understanding how grazing management can lead to recovery of ecosystem function is essential to design and implement effective strategies for sustainable use of this resource. In a long‐term field experiment carried out in Southern Brazil, we studied the effects of temporal grazing exclusions (spring or fall) at moderate and severe livestock grazing intensities (maintained by adjusting contrasting forage allowances) on the species richness, botanical composition, forage mass, sward height, and photosynthetic active radiation intercepted. The experiment was arranged in a completely randomized design with three replications of grazing exclusions, applied simultaneously at moderate and severe grazing intensities. Moderate grazing intensity showed a bimodal structure of shorter and taller canopies, and high species richness. Severe grazing created a shorter and homogeneous sward structure characterized by less standing biomass and species loss. In response to grazing exclusions, sward height, standing biomass, and light interception recovered almost to the levels of moderate grazing. Further, within 2 years grass species richness increased and botanical composition changed toward grasses with erect habit prevailing in moderate grazing intensity. Our study confirms that (1) moderate grazing intensities allow the coexistence of high number of species and (2) spring grazing exclusions of long‐term overgrazed grasslands can lead to a quick start to recover the grass species richness, primary productivity, and species composition like that prevailing in well‐managed grasslands.     

Seasonal selection of soil types and grass swards by roan antelope in a South African savanna

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.     

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.     

Time allocation of bison in meadow patches driven by potential energy gains and group size dynamics

Understanding the behavioural mechanisms involved in broad‐scale spatial organisation of grazing herbivores requires uncovering the factors controlling foraging decisions, such as patch residency time. Foraging theory specifies that rate maximizers must simultaneously consider both the optimal residency time in a food patch and the optimal diet. Specifically, resource depletion or spatial variation in food type availability should not influence food choice, but only patch residency time. Few studies, however, have tested these principles together, and none on free‐ranging large herbivores. We evaluated the combined effects of forage characteristics, predation risk, and group size on residency time by free‐ranging bison Bison bison in summer. We hypothesized that residency time in meadows would increase with the availability of Carex atherodes, a highly profitable plant species maximizing energy intake rate, but decrease with sward complexity (i.e. plant species composition and structure) within foraging stations. We also anticipated that predation risk and group size would influence the relationship between vegetation characteristics and residency time. Residency times were measured in 44 sites using cameras located at meadow edges. We determined that residency time in meadows varied with meadow area, group size, biomass of C. atherodes available on the area, and proportion of C. atherodes within foraging stations. We found that the likelihood of departure decreased with an increase in the total biomass of C. atherodes available over the meadow, an effect attenuated by an increase in group size. Residency time in meadows was also influenced by plant species composition, with higher accessibility of C. atherodes within feeding stations increasing residency time. We found little evidence, however, that sward structure and predation risk influenced residency time. Overall, our study demonstrated that the search for rapid energy gains, together with the constraints imposed by group living, can explain time allocation in habitat patches by large gregarious herbivores.     

Use of trade-off theory to advance understanding of herbivore–parasite interactions

• 1 Trade‐off theory has been extensively used to further our understanding of animal behaviour. In mammalian herbivores, it has been used to advance our understanding of their reproductive, parental care and foraging strategies. Here, we detail how trade‐off theory can be applied to herbivore–parasite interactions, especially in foraging environments.
• 2 Foraging is a common mode of uptake of parasites that represent the most pervasive challenge to mammalian fitness and survival. Hosts are hypothesized to alter their foraging behaviour in the presence of parasites in three ways: (i) hosts avoid foraging in areas that are contaminated with parasites; (ii) hosts select diets that increase their resistance and resilience to parasites; and (iii) hosts select for foods with direct anti‐parasitic properties (self‐medication). We concentrate on the mammalian herbivore literature to detail the recent advances made using trade‐off frameworks to understand the mechanisms behind host–parasite interactions in relation to these three hypotheses.
• 3 In natural systems, animals often face complex foraging decisions including nutrient intake vs. predation risk, nutrient intake vs. sheltering and nutrient intake vs. parasite risk trade‐offs. A trade‐off framework is detailed that can be used to interpret mammal behaviour in complex environments, and may be used to advance the self‐medication hypothesis.
• 4 The use of trade‐off theory has advanced our understanding of the contact process between grazing mammalian hosts and their parasites transmitted via the faecal–oral route. Experimental manipulation of the costs and benefits of a nutrient intake vs. parasite risk trade‐off has shown that environmental conditions (forage quality and quantity) and the physiological state (parasitic and immune status) of a mammalian host can both affect the behavioural decisions of foraging animals.
• 5 Naturally occurring trade‐offs and the potential to manipulate their costs and benefits enables us to identify the abilities and behavioural rules used by mammals when making decisions in complex environments and thus predict animal behaviour.

     

Food quality and quantity are more important in explaining foraging of an intermediate‐sized mammalian herbivore than predation risk or competition

During times of high activity by predators and competitors, herbivores may be forced to forage in patches of low‐quality food. However, the relative importance in determining where and what herbivores forage still remains unclear, especially for small‐ and intermediate‐sized herbivores. Our objective was to test the relative importance of predator and competitor activity, and forage quality and quantity on the proportion of time spent in a vegetation type and the proportion of time spent foraging by the intermediate‐sized herbivore European hare (Lepus europaeus). We studied red fox (Vulpes vulpes) as a predator species and European rabbit (Oryctolagus cuniculus) as a competitor. We investigated the time spent at a location and foraging time of hare using GPS with accelerometers. Forage quality and quantity were analyzed based on hand‐plucked samples of a selection of the locally most important plant species in the diet of hare. Predator activity and competitor activity were investigated using a network of camera traps. Hares spent a higher proportion of time in vegetation types that contained a higher percentage of fibers (i.e., NDF). Besides, hares spent a higher proportion of time in vegetation types that contained relatively low food quantity and quality of forage (i.e., high percentage of fibers) during days that foxes (Vulpes vulpes) were more active. Also during days that rabbits (Oryctolagus cuniculus) were more active, hares spent a higher proportion of time foraging in vegetation types that contained a relatively low quality of forage. Although predation risk affected space use and foraging behavior, and competition affected foraging behavior, our study shows that food quality and quantity more strongly affected space use and foraging behavior than predation risk or competition. It seems that we need to reconsider the relative importance of the landscape of food in a world of fear and competition.     

Grazing in heterogeneous environments: infra- and supra-parasite distributions determine herbivore grazing decisions

We tested the hypothesis that the infra-gastrointestinal parasite population of herbivores affects their grazing behaviour in relation to the supra-parasite population of parasites in the environment. Our first objective was to create a naturally heterogeneous sward structure of gaps and tussocks using a continuous grazing scheme. We then demonstrate that a nutrition vs. parasitism grazing trade-off occurs within that sward structure and that infra-gastrointestinal parasite populations affect the grazing decisions of herbivores faced with the trade-off. A pool of 50 naturally parasitised female Soay sheep and their lambs were used to create a heterogeneous tall, faeces-contaminated tussock/short, non-contaminated gap sward structure in a 1-ha experimental plot. Tussocks offered approximately 1.5 times greater forage intake but contained 5.5 times the number of strongyle parasites compared to the gaps. Following a 10-week period in which the heterogeneous sward structure was created, two 5-day periods of observations of sward structure selection (i.e. gap vs. tussock) were carried out. Twenty female Soay lambs were divided into two groups of ten (balanced for live-weight) immediately prior to the start of the observation period. One of the groups of lambs was treated with an anthelmintic drench before the start of the second observation period creating two levels of parasitism (high and low). On each observation day 5-min focal observations were carried out on each animal at least twice a day, during which time the number of bites taken from gaps and tussocks were recorded along with the number of steps. During the first period of observations, all animals rejected the relatively tall, faeces-contaminated tussocks for grazing to a similar extent and had similar bite and step rates. During the second period of observations all animals showed reduced rejection of the tussocks relative to the first week, however, animals with a reduced parasite population showed a greater reduction in rejection as compare to the highly parasitised animals. We conclude that the infra- and supra-distributions of parasites within herbivore hosts and the environment greatly impact on herbivore grazing behaviour and foraging decisions and thus the structure and heterogeneity of grazed ecosystems.     

The relationship between forage cell wall content and voluntary food intake in mammalian herbivores

• 1 It is generally assumed that animals compensate for a declining diet quality with increasing food intake. Differences in the response to decreasing forage quality in herbivores have been postulated particularly between cattle (ruminants) and horses (hindgut fermenters). However, empirical tests for both assumptions in herbivorous mammals are rare.
• 2 We collected data on voluntary food intake in mammals on forage‐only diets and related this to dietary neutral detergent fibre (NDF) content, assuming a nonlinear correlation between these measurements. Generally, the paucity of corresponding data is striking.
• 3 Elephants and pandas showed very high food intakes that appeared unrelated to dietary fibre content. Only in small rodents, and possibly in rabbits, was an increase in food intake on forages of higher NDF content evident. In particular, other large herbivores, including horses, followed patterns of decreasing intake with increasing forage NDF, also observed in domestic cattle or sheep.
• 4 For large herbivores, empirical data therefore do not – so far – support the notion that intake is increased in response to declining diet quality. However, data are in accord with the assumption that most large herbivores have an anticipatory strategy of acquiring body reserves when high‐quality forage is available, and reducing food intake (and potentially metabolic losses) when only low‐quality forage is available.
• 5 Intake studies in which the influence of digestive strategy on food intake capacity is tested should be designed as long‐term studies that outlast an anticipatory strategy and force animals to ingest as much as possible.
• 6 We suggest that a colonic separation mechanism coupled with coprophagy, in order to minimize metabolic faecal losses, is necessary below a body size threshold where an anticipatory strategy (living off body reserves, migration) is not feasible. Future studies aimed at investigating fine‐scale differences, for example between equids and bovids, should focus on non‐domesticated species.

     

Can grazing sheep compensate for a daily foraging time constraint?

1. Theoretical studies of large herbivore foraging assume that total daily grazing time is a key constraint on daily intake and diet choice. We experimentally tested this assumption and investigated the effects of food availability on the ability of grazing sheep to compensate for restriction of available daily grazing time.
2. Foraging behaviour, intake and diet digestibility by sheep, were measured on grass pastures in a replicated 2 × 2 factorial experiment, in which overnight access to pasture was varied (restricted overnight and continuous access) on two sward heights (5·5 and 3·0 cm), representing high and low food availability.
3. Regardless of food availability, the overnight-restricted sheep fed for almost all of the available grazing time by grazing for fewer, longer foraging bouts, but still had much shorter total daily grazing time than the continuous access sheep.
4. In response to overnight penning, the sheep had a significantly higher instantaneous rate of intake achieved mainly via larger bites. The continuous access sheep were hence not maximizing their short-term rate of intake, whilst grazing according to the daily schedule considered normal for sheep.
5. The behavioural responses to overnight food restriction were able to counteract the reduction in daily grazing time only where food availability was high. In contrast on short swards overnight grazing restriction led to a reduction in total daily intake. We suggest that the interactions between the factors considered as constraints on foraging behaviour of herbivores are, as yet, only poorly quantified.     

Foraging behaviour at multiple temporal scales in a wild alpine equid

Forage abundance, forage quality, and social factors are key elements of the foraging ecology of wild herbivores. For non-ruminant equids, forage-limited environments are likely to impose severe constraints on their foraging behaviour. We used a multi-scale approach to study foraging behaviour in kiang (Equus kiang), a wild equid inhabiting the high-altitude rangelands of the Tibetan Plateau. Using behavioural observations and vegetation sampling, we first assessed how patterns of plant abundance and quality affected (i) the instantaneous forage intake rate (fine scale) and (ii) the proportion of time spent foraging (coarse scale) across seasons. We also tested whether foraging behaviour differed among group types, between sex in adults, and between females of different reproductive status. At a fine scale, intake rate increased linearly with bite size and increased following a type II curvilinear function with biomass on feeding sites. Forage intake rate also increased linearly with plant quality. Male and female kiangs had similar intake rates. Likewise, gravid and lactating females had similar intake rates as barren and non-lactating females. At a coarse scale, kiangs spent longer time feeding in mesic than in xeric habitats, and spent more time feeding in early summer and fall than in late summer. Groups of adults with foals spent less time feeding than male groups and groups of adults without foals. Our findings suggest that kiangs use flexible foraging behaviours in relation to seasonal variations of vegetation quality and abundance, a likely outcome of the extreme seasonal conditions encountered on the Tibetan Plateau.     

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.     

Green‐up selection by red deer in heterogeneous,human‐dominated landscapes of Central Europe

The forage maturation hypothesis (FMH) assumes that herbivores cope with the trade‐off between digestibility and biomass in forage by selecting vegetation at intermediate growth. The green wave hypothesis (GWH) extends the FMH to suggest how spatiotemporal heterogeneity in plant quality shapes migratory movements of herbivores. Growing empirical support for these hypotheses mainly comes from studies in vast landscapes with large‐scale habitat heterogeneity. It is unclear, however, to what extent ungulates surf green waves in human‐altered landscapes with small‐scale heterogeneity in terms of land use and topography. We used plant phenological proxies derived from Sentinel 2 satellite data to analyze the habitat selection of 93 collared red deer (Cervus elaphus) in montane and alpine habitats. Using a step selection analysis, we investigated how plant phenology, that is, the instantaneous rate of green‐up (IRG) and normalized difference vegetation index (NDVI), and a set of variables describing topography and human presence influenced red deer resource selection in open habitats. We learned that red deer selected areas with high biomass at green‐up and avoided habitats with possible exposure to human activity. Additionally, landscape structure and topography strongly influenced spatial behavior of red deer. We further compared cumulative access to high‐quality forage across migrant strategies and found migrants gained better access than residents. Many migratory individuals surfed the green wave, and their surfing behavior, however, became less pronounced with decreasing distance to settlements. Within the constraints of topography and human land use, red deer track spring green‐up on a fine spatiotemporal scale and follow the green wave across landscapes in migration movements. Thus, they benefit from high‐quality forage even in human‐dominated landscapes with small‐scale heterogeneity and vegetation emerging in a heterogenic, dynamic mosaic.     

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     

Evolutionary irony: evidence that ‘defensive’ plant spines act as a proximate cue to attract a mammalian herbivore

Many plants produce structural defenses to deter feeding by herbivores. However, many previous studies testing whether spines are effective at defending against mammalian herbivores have produced equivocal results. These ambiguous results are hypothesized to be due to herbivore counter‐adaptations. We investigated potential counter‐adaptations in a population of white‐throated woodrats Neotoma albigua that specialize on cactus by investigating feeding behavior and preference for cacti varying in spinescence. Neotoma albigula exhibited a unique behavior of clipping cactus spines, which renders these defenses ineffective. Strikingly, these woodrats chose to collect spiny cacti over experimentally de‐spined cacti, demonstrating that spines act as a proximal cue that attracts woodrats. This attraction is likely due to the higher protein and lower fiber content of spiny cacti compared to naturally non‐spiny cacti. Thus, the ‘defensive’ spines of cacti are ineffective against a specialist herbivore and instead serve as an indicator of nutritional quality that promotes herbivory. Our results support the ‘rule‐of‐thumb’ hypothesis of foraging, which states that herbivores forage according to obvious visual cues that are indicative of nutritional content, rather than sampling nutrient composition of plants. We propose that specialist herbivores are unique systems in which to study other counter‐adaptations to structural defenses and ‘rule‐of‐thumb’ foraging strategies.     

Spatio‐temporal feeding selection of red deer in a mountainous landscape

Understanding the dietary consumption and selection of wild populations of generalist herbivores is hampered by the complex array of factors. Here, we determine the influence of habitat, season, and animal density, sex, and age on the diet consumption and selection of 426 red deer (Cervus elaphus scoticus) culled in Fiordland National Park, New Zealand. Our site differs from studies elsewhere both in habitat (evergreen angiosperm‐dominated forests) and the intensity of hunting pressures. We predicted that deer would not consume forage in proportion to its relative availability, and that dietary consumption would change among and within years in response to hunting pressures that would also limit opportunities for age and sex segregation. Using canonical correspondence analysis, we evaluated the relative importance of different drivers of variation in diet consumption assessed from gut content and related these to available forage in the environment. We found that altitude explained the largest proportion of variation in diet consumption, reflecting the ability of deer to alter their consumption and selection in relation to their foraging grounds. Grasses formed a high proportion of the diet consumption, even for deer culled several kilometres from the alpine grasslands. In the winter months, when the alpine grasslands were largely inaccessible, less grass was eaten and deer resorted to woody plants that were avoided in the summer months. Surprisingly, there were no significant dietary differences between adults and juveniles and only subtle differences between the sexes. Sex‐based differences in diet consumption are commonly observed in ungulate species and we suggest that they may have been reduced in our study area owing to decreased heterogeneity in available forage as the diversity of palatable species decreased under high deer browsing pressures, or by intense hunting pressure.     

Plant Trait Assembly Affects Superiority of Grazer's Foraging Strategies in Species-Rich Grasslands

Background

Current plant – herbivore interaction models and experiments with mammalian herbivores grazing plant monocultures show the superiority of a maximizing forage quality strategy (MFQ) over a maximizing intake strategy (MI). However, there is a lack of evidence whether grazers comply with the model predictions under field conditions.

Methodology/Findings

We assessed diet selection of sheep (Ovis aries) using plant functional traits in productive mesic vs. low-productivity dry species-rich grasslands dominated by resource-exploitative vs. resource-conservative species respectively. Each grassland type was studied in two replicates for two years. We investigated the first grazing cycle in a set of 288 plots with a diameter of 30 cm, i.e. the size of sheep feeding station. In mesic grasslands, high plot defoliation was associated with community weighted means of leaf traits referring to high forage quality, i.e. low leaf dry matter content (LDMC) and high specific leaf area (SLA), with a high proportion of legumes and the most with high community weighted mean of forage indicator value. In contrast in dry grasslands, high community weighted mean of canopy height, an estimate of forage quantity, was the best predictor of plot defoliation. Similar differences in selection on forage quality vs. quantity were detected within plots. Sheep selected plants with higher forage indicator values than the plot specific community weighted mean of forage indicator value in mesic grasslands whereas taller plants were selected in dry grasslands. However, at this scale sheep avoided legumes and plants with higher SLA, preferred plants with higher LDMC while grazing plants with higher forage indicator values in mesic grasslands.

Conclusions

Our findings indicate that MFQ appears superior over MI only in habitats with a predominance of resource-exploitative species. Furthermore, plant functional traits (LDMC, SLA, nitrogen fixer) seem to be helpful correlates of forage quality only at the community level.